EVOLUTION

A book of readings from Wikipedia

Volume 1: An Introduction

Khalid Chraibi

04 septembre 2010
Part of the Biology series on Evolution
Mechanisms and processes
Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation
Research and history
Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact
Evolutionary biology fields
Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics
Biology portal · v • d • e
Part of the Biology series on Genetics

Key components
Chromosome
DNA · RNA
Genome
Heredity
Mutation
Nucleotide
Variation
Glossary
Index
Outline
History and topics
Introduction
History
Classical genetics
Evolution · Molecular
Mendelian inheritance
Molecular genetics
Research
DNA sequencing
Genetic engineering
Genomics · Topics
Medical genetics
Branches in genetics
Evolution Wikipedia the main articles
Overview
Introduction to evolution
Evolution
Evolution as theory and fact
Evolutionary history of life
Timeline of evolution
History
History of evolutionary thought
Lamarckism
 Saltationism
Orthogenesis
On the Origin of Species
Darwinism
The Genetical Theory of Natural Selection
Neo-Darwinism
Modern evolutionary synthesis
Base concepts
Heredity
Fitness
Common descent
Evidence of common descent
Mechanisms
Adaptation
Genetic drift
Gene flow
 Mutation
Natural selection
Speciation
Phylogenetics
Phylogenetics
Cladistics
Cladogram
Molecular phylogenetics
Fields
Evolutionary developmental biology
Molecular evolution
Human evolution
Evolutionary psychology
Controversy and social impacts
Creation–evolution controversy
Objections to evolution
 Creationism
Intelligent design
Social effect of evolutionary theory
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Basic topics in evolutionary biology
Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary Canalisation · Modularity · Phenotypic plasticity

developmental
biology (Evo-devo)
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

 Retrieved from "http://en.wikipedia.org/wiki/Evolution"

Index of evolutionary biology articles

From Wikipedia, the free encyclopedia

(Redirected from List of evolutionary biology topics)
Jump to: navigation, search
This is a list of topics in evolutionary biology.
Main article: Evolutionary biology
Contents: A B C D E F G H I J K L M N O P Q R S T U V W X Y Z—
See also
This list is incomplete; you can help by expanding it.
[edit] A
abiogenesis - adaptation - adaptive radiation - allele - allele frequency - allopatric speciation -
altruism - anagenesis - Archaeopteryx - aquatic adaptation - artificial selection - atavism

[edit] B
biological organisation - Brassica oleracea - breed

[edit] C
Cambrian explosion - catagenesis - gene-centered view of evolution - cephalization - Sergei
Chetverikov - Chi square test - chronobiology - chronospecies - clade - cladistics - Climatic adaptation -
coalescent theory - co-evolution - Co-operation (evolution) - coefficient of relationship - common descent -
convergent evolution - creation-evolution controversy - cultivar - current research in evolutionary biology

[edit] D
Darwin (unit) - Charles Darwin - Darwinism - Darwin's finches - Richard Dawkins - directional
selection - Theodosius Dobzhansky - Dog breeding - Domestication - Domestication of the horse

[edit] E
ecological genetics - ecological selection - endosymbiosis - error threshold (evolution) - evidence of
common descent - evolution - evolutionary arms race - evolutionary capacitance - evolution of cetaceans -
evolution of complexity - evolution of the horse - evolution of mammalian auditory ossicles - evolution of
mammals - evolution of sex - evolution of sirenians - evolution of the eye - evolutionary developmental
biology - evolutionary neuroscience - evolutionary psychology - evolutionary radiation - evolutionary stable
strategy - evolutionary tree - evolvability - experimental evolution - exaptation - extinction

[edit] F
Joe Felsenstein - R.A. Fisher - Fisher's reproductive value - fitness - fitness landscape - E.B. Ford -
fossil

[edit] G
Galápagos Islands - gene - gene-centric view of evolution - gene duplication - gene flow - gene pool -
genetic drift - genetic hitchhiking - genetic recombination - genetic variation - genotype - genotype-
environment correlation - genotype-environment interaction - genotype-phenotype distinction - Stephen Jay
Gould - gradualism - Peter and Rosemary Grant - group selection

[edit] H
J. B. S. Haldane - W. D. Hamilton - Hardy-Weinberg principle - heredity - hierarchy of life - history of
evolutionary thought - horizontal gene transfer - human evolution - human evolutionary genetics -
homologous chromosomes - homology (biology) - Julian Huxley - human vestigiality
[edit] I
inclusive fitness - insect evolution - Invertebrate paleontology (a.k.a. invertebrate paleobiology or
paleozoology)

[edit] K
kin selection - Motoo Kimura–Karyotype - Koinophilia

[edit] L
Jean-Baptiste Lamarck - Lamarckism - landrace - Language -Last common ancestor - Last universal
ancestor - Richard Lewontin - list of gene families - List of human evolution fossils - life-history theory - Wen-
Hsiung Li - living fossils - Charles Lyell

[edit] M
macroevolution - macromutation - The Major Transitions in Evolution - maladaptation - mass
extinctions - mating systems - John Maynard Smith - Ernst Mayr - Gregor Mendel - memetics - Mendelian
inheritance - microevolution - Micropaleontology (a.k.a. micropaleobiology) - Mitochondrial Eve - modern
evolutionary synthesis - molecular clock - molecular evolution - molecular phylogeny - molecular systematics
- Monogamy - most recent common ancestor - Hermann Joseph Muller - Muller's ratchet - mutation -
mutational meltdown
[edit] N
natural selection - Nature versus nurture - Neo-Darwinism - neutral theory of molecular evolution -
Baron Franz Nopcsa

[edit] O
Susumu Ohno - Aleksandr Oparin - On The Origin of Species - Origin of birds - orthologous genes

[edit] P
Paleoanthropology - Paleobiology - Paleobotany - Paleontology - Paleozoology - Paleozoology:
Vertebrates - Paleozoology: Invertebrates - parallel evolution - paraphyletic - particulate inheritance -
peppered moth - peppered moth evolution - peripatric speciation - phenotype - phylogenetics - phylogeny -
phylogenetic tree - pikaia - Plant evolution - polymorphism (biology) - population - Population bottleneck -
population dynamics - population genetics - preadaptation - prehistoric archaeology - Principles of Geology -
George R. Price - Price equation - punctuated equilibrium - Plant Evolutionary Developmental Biology

[edit] Q
quasispecies model

[edit] R
race (biology) - Red Queen - recapitulation theory - recombination - Bernhard Rensch
[edit] S
selection - selective breeding - Science of Evolution - sexual selection - sociobiology - species -
speciation - species flock - sperm competition - stabilizing selection - strain (biology) - subspecies - survival
of the fittest - Survival value - symbiogenesis - systematics - George Gaylord Simpson - G. Ledyard Stebbins

[edit] T
Tiktaalik - timeline of evolution - Trait (biological) - transgressive phenotype - transitional fossil -
transposon - Triangle of U

[edit] U
Uniformitarianism (science) - Unit of selection

[edit] V
variety (plant) - Vertebrate paleontology (a.k.a. vertebrate paleobiology or paleozoology) - viral
evolution - The Voyage of the Beagle - vestigiality

[edit] W
Alfred Russel Wallace - Wallace effect - Wallace Line - Wallacea - George C. Williams - Edward O.
Wilson - Sewall Wright
[edit] Y
Y-chromosomal Adam - Y-DNA haplogroups by ethnic groups

[edit] See also

• List of biology topics
• List of biochemistry topics
Retrieved from "http://en.wikipedia.org/wiki/Index_of_evolutionary_biology_articles"

Categories: Biology lists | Evolutionary biology | Indexes of articles

• This page was last modified on 21 August 2010 at 18:28.

Related topics
Evolutionary biology fields— Cladistics • Ecological genetics • Evolutionary development • Human
evolution • Molecular evolution • Phylogenetics • Population genetics
Mechanisms— Natural selection • Genetic drift • Gene flow
Outcomes— Adaptation • Co-evolution • Co-operation • Speciation • Extinction
 Research and history— Evidence of common descent • Evolutionary history of life • History of evolutionary
thought • Modern evolutionary synthesis • Social effect of evolutionary theory • Objections to evolution

Categories
Evolutionary biology
Evolutionary biologists • Eugenics • Evolutionary biology literature • Evolutionary dynamics •
Evolutionary game theory • Extinction • History of evolutionary biology • Human evolution • Microbial
population biology • Molecular evolution • Mutation • Paleontology • Phylogenetics • Population genetics •
Evolutionary psychology • Selection • Sociobiology • Speciation • Symbiosis • Tree of life •
 W000

Universe
From Wikipedia, the free encyclopedia

Jump to: navigation, search

For other uses, see Universe (disambiguation).
Physical cosmology
 Universe · Big Bang
Age of the universe
Timeline of the Big Bang
Ultimate fate of the universe
[show]Early universe

Inflation · Nucleosynthesis
GWB · Neutrino background
Cosmic microwave background

[show]Expanding universe

Redshift · Hubble's law

Metric expansion of space
Friedmann equations
FLRW metric
 [show]Structure Formation

Shape of the Universe

Structure formation
Reionization
Galaxy formation
Large-scale structure
Galaxy filaments

[show]Components

Lambda-CDM model
Dark energy · Dark matter

[show]Timeline

Timeline of cosmological theories

Future of an expanding universe

[show]Experiments

Observational cosmology
2dF · SDSS
COBE · BOOMERanG · WMAP · Planck
 [show]Scientists

Isaac Newton · Einstein · Hawking · Friedman · Lemaître · Hubble · Penzias · Wilson · Gamow ·
Dicke · Zel'dovich · Mather · Rubin · Smoot· others
v•d•e
The universe is commonly defined as the totality of everything that exists,[1] including all physical
matter and energy, the planets, stars, galaxies, and the contents of intergalactic space,[2][3] although this
usage may differ with the context (see definitions, below). The term universe may be used in slightly different
contextual senses, denoting such concepts as the cosmos, the world, or nature.
Observations of earlier stages in the development of the universe, which can be seen at great
distances, suggest that the universe has been governed by the same physical laws and constants throughout
most of its extent and history.
Contents
[hide]
• 1 History
• 2 Etymology, synonyms and definitions
• 2.1 Broadest definition: reality and probability
• 2.2 Definition as reality
• 2.3 Definition as connected space-time
• 2.4 Definition as observable reality
• 3 Size, age, contents, structure, and laws
• 3.1 Fine tuning
• 4 Historical models
• 4.1 Creation
• 4.2 Philosophical models
• 4.3 Astronomical models
• 5 Theoretical models
• 5.1 General theory of relativity
• 5.2 Special relativity and space-time
• 5.3 Solving Einstein's field equations
• 5.4 Big Bang model
• 6 Untestable proposals
• 6.1 Multiverse theory
[edit] History
Throughout recorded history, several cosmologies and cosmogonies have been proposed to account
for observations of the universe. The earliest quantitative geocentric models were developed by the ancient
Greeks, who proposed that the universe possesses infinite space and has existed eternally, but contains a
single set of concentric spheres of finite size – corresponding to the fixed stars, the Sun and various planets –
rotating about a spherical but unmoving Earth. Over the centuries, more precise observations and improved
theories of gravity led to Copernicus's heliocentric model and the Newtonian model of the Solar System,
respectively. Further improvements in astronomy led to the realization that the Solar System is embedded in
a galaxy composed of millions of stars, the Milky Way, and that other galaxies exist outside it, as far as
astronomical instruments can reach. Careful studies of the distribution of these galaxies and their spectral
lines have led to much of modern cosmology. Discovery of the red shift and cosmic microwave background
radiation revealed that the universe is expanding and apparently had a beginning.
 This high-resolution image of the Hubble ultra deep field shows a diverse range of galaxies, each
consisting of billions of stars. The equivalent area of sky that the picture occupies is shown in the lower left
corner. The smallest, reddest galaxies, about 100, are some of the most distant galaxies to have been
imaged by an optical telescope, existing at the time shortly after the Big Bang.
 According to the prevailing scientific model of the universe, known as the Big Bang, the universe
expanded from an extremely hot, dense phase called the Planck epoch, in which all the matter and energy of
the observable universe was concentrated. Since the Planck epoch, the universe has been expanding to its
present form, possibly with a brief period (less than 10 −32 seconds) of cosmic inflation. Several independent
experimental measurements support this theoretical expansion and, more generally, the Big Bang theory.
Recent observations indicate that this expansion is accelerating because of dark energy, and that most of the
matter in the universe may be in a form which cannot be detected by present instruments, and so is not
accounted for in the present models of the universe; this has been named dark matter. The imprecision of
current observations has hindered predictions of the ultimate fate of the universe.
Current interpretations of astronomical observations indicate that the age of the universe is 13.75
±0.17 billion years,[4] and that the diameter of the observable universe is at least 93 billion light years, or
8.80 × 1026 metres.[5] According to general relativity, space can expand faster than the speed of light,
although we can view only a small portion of the universe due to the limitation imposed by light speed. Since
we cannot observe space beyond the limitations of light (or any electromagnetic radiation), it is uncertain
whether the size of the universe is finite or infinite.

[edit] Etymology, synonyms and definitions

See also: Cosmos, Nature, World (philosophy), and Celestial spheres
The word universe derives from the Old French word Univers, which in turn derives from the Latin
word universum.[6] The Latin word was used by Cicero and later Latin authors in many of the same senses
as the modern English word is used.[7] The Latin word derives from the poetic contraction Unvorsum — first
used by Lucretius in Book IV (line 262) of his De rerum natura (On the Nature of Things) — which connects
un, uni (the combining form of unus', or "one") with vorsum, versum (a noun made from the perfect passive
participle of vertere, meaning "something rotated, rolled, changed").[7] Lucretius used the word in the sense
"everything rolled into one, everything combined into one".

Artistic rendition (highly exaggerated) of a Foucault pendulum showing that the Earth is not
stationary, but rotates.
 An alternative interpretation of unvorsum is "everything rotated as one" or "everything rotated by
one". In this sense, it may be considered a translation of an earlier Greek word for the universe, περιφορά,
"something transported in a circle", originally used to describe a course of a meal, the food being carried
around the circle of dinner guests.[8] This Greek word refers to an early Greek model of the universe, in
which all matter was contained within rotating spheres centered on the Earth; according to Aristotle, the
rotation of the outermost sphere was responsible for the motion and change of everything within. It was
natural for the Greeks to assume that the Earth was stationary and that the heavens rotated about the Earth,
because careful astronomical and physical measurements (such as the Foucault pendulum) are required to
prove otherwise.
The most common term for "universe" among the ancient Greek philosophers from Pythagoras
onwards was τὸ πᾶν (The All), defined as all matter (τὸ ὅλον) and all space (τὸ κενόν).[9][10] Other
synonyms for the universe among the ancient Greek philosophers included κόσμος (meaning the world, the
cosmos) and φύσις (meaning Nature, from which we derive the word physics).[11] The same synonyms are
found in Latin authors (totum, mundus, natura)[12] and survive in modern languages, e.g., the German words
Das All, Weltall, and Natur for universe. The same synonyms are found in English, such as everything (as in
the theory of everything), the cosmos (as in cosmology), the world (as in the many-worlds hypothesis), and
Nature (as in natural laws or natural philosophy).[13]

[edit] Broadest definition: reality and probability

See also: Introduction to quantum mechanics, Interpretation of quantum mechanics, and Many-
worlds hypothesis
The broadest definition of the universe can be found in De divisione naturae by the medieval
philosopher and theologian Johannes Scotus Eriugena, who defined it as simply everything: everything that
is created and everything that is not created. Time is not considered in Eriugena's definition; thus, his
definition includes everything that exists, has existed and will exist, as well as everything that does not exist,
has never existed and will never exist. This all-embracing definition was not adopted by most later
philosophers, but something not entirely dissimilar reappears in quantum physics, perhaps most obviously in
the path-integral formulation of Feynman.[14] According to that formulation, the probability amplitudes for the
various outcomes of an experiment given a perfectly defined initial state of the system are determined by
summing over all possible paths by which the system could progress from the initial to final state. Naturally,
an experiment can have only one outcome; in other words, only one possible outcome is made real in this
universe, via the mysterious process of quantum measurement, also known as the collapse of the
wavefunction (but see the many-worlds hypothesis below in the Multiverse section). In this well-defined
mathematical sense, even that which does not exist (all possible paths) can influence that which does finally
exist (the experimental measurement). As a specific example, every electron is intrinsically identical to every
other; therefore, probability amplitudes must be computed allowing for the possibility that they exchange
positions, something known as exchange symmetry. This conception of the universe embracing both the
existent and the non-existent loosely parallels the Buddhist doctrines of shunyata and interdependent
development of reality, and Gottfried Leibniz's more modern concepts of contingency and the identity of
indiscernibles.

[edit] Definition as reality

See also: Reality and Physics
More customarily, the universe is defined as everything that exists, has existed, and will exist[ citation
needed]. According to this definition and our present understanding, the universe consists of three elements:
space and time, collectively known as space-time or the vacuum; matter and various forms of energy and
momentum occupying space-time; and the physical laws that govern the first two. These elements will be
discussed in greater detail below. A related definition of the term universe is everything that exists at a single
moment of cosmological time, such as the present, as in the sentence "The universe is now bathed uniformly
in microwave radiation".
The three elements of the universe (spacetime, matter-energy, and physical law) correspond roughly
to the ideas of Aristotle. In his book The Physics (Φυσικῆς, from which we derive the word "physics"),
Aristotle divided τὸ πᾶν (everything) into three roughly analogous elements: matter (the stuff of which the
universe is made), form (the arrangement of that matter in space) and change (how matter is created,
destroyed or altered in its properties, and similarly, how form is altered). Physical laws were conceived as the
rules governing the properties of matter, form and their changes. Later philosophers such as Lucretius,
Averroes, Avicenna and Baruch Spinoza altered or refined these divisions[citation needed]; for example,
Averroes and Spinoza discern natura naturans (the active principles governing the universe) from natura
naturata, the passive elements upon which the former act.

[edit] Definition as connected space-time

See also: Chaotic Inflation theory
It is possible to conceive of disconnected space-times, each existing but unable to interact with one
another. An easily visualized metaphor is a group of separate soap bubbles, in which observers living on one
soap bubble cannot interact with those on other soap bubbles, even in principle. According to one common
terminology, each "soap bubble" of space-time is denoted as a universe, whereas our particular space-time
is denoted as the universe, just as we call our moon the Moon. The entire collection of these separate space-
times is denoted as the multiverse.[15] In principle, the other unconnected universes may have different
dimensionalities and topologies of space-time, different forms of matter and energy, and different physical
laws and physical constants, although such possibilities are currently speculative.

[edit] Definition as observable reality

See also: Observable universe and Observational cosmology
According to a still-more-restrictive definition, the universe is everything within our connected space-
time that could have a chance to interact with us and vice versa.[ citation needed] According to the general
theory of relativity, some regions of space may never interact with ours even in the lifetime of the universe,
due to the finite speed of light and the ongoing expansion of space. For example, radio messages sent from
Earth may never reach some regions of space, even if the universe would live forever; space may expand
faster than light can traverse it. It is worth emphasizing that those distant regions of space are taken to exist
and be part of reality as much as we are; yet we can never interact with them. The spatial region within which
we can affect and be affected is denoted as the observable universe. Strictly speaking, the observable
universe depends on the location of the observer. By traveling, an observer can come into contact with a
greater region of space-time than an observer who remains still, so that the observable universe for the
former is larger than for the latter. Nevertheless, even the most rapid traveler may not be able to interact with
all of space. Typically, the observable universe is taken to mean the universe observable from our vantage
point in the Milky Way Galaxy.

[edit] Size, age, contents, structure, and laws

Main articles: Observable universe, Age of the universe, Large-scale structure of the universe, and
Abundance of the chemical elements
 The universe is very large and possibly infinite in volume. The region visible from Earth (the
observable universe) is about 92 billion light years across,[16] based on where the expansion of space has
taken the most distant objects observed. For comparison, the diameter of a typical galaxy is only 30,000
light-years, and the typical distance between two neighboring galaxies is only 3 million light-years.[17] As an
example, our Milky Way Galaxy is roughly 100,000 light years in diameter,[18] and our nearest sister galaxy,
the Andromeda Galaxy, is located roughly 2.5 million light years away.[19] There are probably more than 100
billion (1011) galaxies in the observable universe.[20] Typical galaxies range from dwarfs with as few as ten
million[21] (107) stars up to giants with one trillion[22] (1012) stars, all orbiting the galaxy's center of mass.
Thus, a very rough estimate from these numbers would suggest there are around one sextillion (1021) stars
in the observable universe; though a 2003 study by Australian National University astronomers resulted in a
figure of 70 sextillion (7 x 1022)[23].
 The universe is believed to be mostly composed of dark energy and dark matter, both of which are
poorly understood at present. Less than 5% of the universe is ordinary matter, a relatively small perturbation.
The observable matter is spread uniformly (homogeneously) throughout the universe, when
averaged over distances longer than 300 million light-years.[24] However, on smaller length-scales, matter is
observed to form "clumps", i.e., to cluster hierarchically; many atoms are condensed into stars, most stars
into galaxies, most galaxies into clusters, superclusters and, finally, the largest-scale structures such as the
Great Wall of galaxies. The observable matter of the universe is also spread isotropically, meaning that no
direction of observation seems different from any other; each region of the sky has roughly the same content.
[25] The universe is also bathed in a highly isotropic microwave radiation that corresponds to a thermal
equilibrium blackbody spectrum of roughly 2.725 kelvin.[26] The hypothesis that the large-scale universe is
homogeneous and isotropic is known as the cosmological principle,[27] which is supported by astronomical
observations.
The present overall density of the universe is very low, roughly 9.9 × 10−30 grams per cubic
centimetre. This mass-energy appears to consist of 73% dark energy, 23% cold dark matter and 4% ordinary
matter. Thus the density of atoms is on the order of a single hydrogen atom for every four cubic meters of
volume.[28] The properties of dark energy and dark matter are largely unknown. Dark matter gravitates as
ordinary matter, and thus works to slow the expansion of the universe; by contrast, dark energy accelerates
its expansion.
The universe is old and evolving. The most precise estimate of the universe's age is 13.73±0.12
billion years old, based on observations of the cosmic microwave background radiation.[29] Independent
estimates (based on measurements such as radioactive dating) agree, although they are less precise,
ranging from 11–20 billion years[30] to 13–15 billion years.[31] The universe has not been the same at all
times in its history; for example, the relative populations of quasars and galaxies have changed and space
itself appears to have expanded. This expansion accounts for how Earth-bound scientists can observe the
light from a galaxy 30 billion light years away, even if that light has traveled for only 13 billion years; the very
space between them has expanded. This expansion is consistent with the observation that the light from
distant galaxies has been redshifted; the photons emitted have been stretched to longer wavelengths and
lower frequency during their journey. The rate of this spatial expansion is accelerating, based on studies of
Type Ia supernovae and corroborated by other data.
The relative fractions of different chemical elements — particularly the lightest atoms such as
hydrogen, deuterium and helium — seem to be identical throughout the universe and throughout its
observable history.[32] The universe seems to have much more matter than antimatter, an asymmetry
possibly related to the observations of CP violation.[33] The universe appears to have no net electric charge,
and therefore gravity appears to be the dominant interaction on cosmological length scales. The universe
also appears to have neither net momentum nor angular momentum. The absence of net charge and
momentum would follow from accepted physical laws (Gauss's law and the non-divergence of the stress-
energy-momentum pseudotensor, respectively), if the universe were finite.[34]

The elementary particles from which the universe is constructed. Six leptons and six quarks comprise
most of the matter; for example, the protons and neutrons of atomic nuclei are composed of quarks, and the
ubiquitous electron is a lepton. These particles interact via the gauge bosons shown in the middle row, each
corresponding to a particular type of gauge symmetry. The Higgs boson (as yet unobserved) is believed to
confer mass on the particles with which it is connected. The graviton, a supposed gauge boson for gravity, is
not shown.
The universe appears to have a smooth space-time continuum consisting of three spatial dimensions
and one temporal (time) dimension. On the average, space is observed to be very nearly flat (close to zero
curvature), meaning that Euclidean geometry is experimentally true with high accuracy throughout most of
the Universe.[35] Spacetime also appears to have a simply connected topology, at least on the length-scale
of the observable universe. However, present observations cannot exclude the possibilities that the universe
has more dimensions and that its spacetime may have a multiply connected global topology, in analogy with
the cylindrical or toroidal topologies of two-dimensional spaces.[36]
The universe appears to behave in a manner that regularly follows a set of physical laws and
physical constants.[37] According to the prevailing Standard Model of physics, all matter is composed of
three generations of leptons and quarks, both of which are fermions. These elementary particles interact via
at most three fundamental interactions: the electroweak interaction which includes electromagnetism and the
weak nuclear force; the strong nuclear force described by quantum chromodynamics; and gravity, which is
best described at present by general relativity. The first two interactions can be described by renormalized
quantum field theory, and are mediated by gauge bosons that correspond to a particular type of gauge
symmetry. A renormalized quantum field theory of general relativity has not yet been achieved, although
various forms of string theory seem promising. The theory of special relativity is believed to hold throughout
the universe, provided that the spatial and temporal length scales are sufficiently short; otherwise, the more
general theory of general relativity must be applied. There is no explanation for the particular values that
physical constants appear to have throughout our universe, such as Planck's constant h or the gravitational
constant G. Several conservation laws have been identified, such as the conservation of charge, momentum,
angular momentum and energy; in many cases, these conservation laws can be related to symmetries or
mathematical identities.

[edit] Fine tuning

Main article: fine-tuned universe
It appears that many of properties of the universe have special values in the sense that a universe
where these properties only differ slightly would not be able to support intelligent life.[38][39] Not all scientists
agree that this fine-tuning exists.[40][41] In particular, it is not known under what conditions intelligent life
could form and what form or shape that would take. A relevant observation in this discussion is that existence
of an observer to observe fine-tuning, requires that the universe supports intelligent life. As such the
conditional probability of observing a universe that is fine-tuned to support intelligent life is 1. This
observation is known as the anthropic principle and is particularly relevant if the creation of the universe was
probabilistic or if multiple universes with a variety of properties exist (see below).

[edit] Historical models

See also: Cosmology and Timeline of cosmology
Many models of the cosmos (cosmologies) and its origin (cosmogonies) have been proposed, based
on the then-available data and conceptions of the universe. Historically, cosmologies and cosmogonies were
based on narratives of gods acting in various ways. Theories of an impersonal universe governed by physical
laws were first proposed by the Greeks and Indians. Over the centuries, improvements in astronomical
observations and theories of motion and gravitation led to ever more accurate descriptions of the universe.
The modern era of cosmology began with Albert Einstein's 1915 general theory of relativity, which made it
possible to quantitatively predict the origin, evolution, and conclusion of the universe as a whole. Most
modern, accepted theories of cosmology are based on general relativity and, more specifically, the predicted
Big Bang; however, still more careful measurements are required to determine which theory is correct.

[edit] Creation
Main articles: Creation myth and Creator deity

Sumerian account of the creatrix goddess Nammu, the precursor of the Assyrian goddess Tiamat;
perhaps the earliest surviving creation myth.
 Many cultures have stories describing the origin of the world, which may be roughly grouped into
common types. In one type of story, the world is born from a world egg; such stories include the Finnish epic
poem Kalevala, the Chinese story of Pangu or the Indian Brahmanda Purana. In related stories, the creation
idea is caused by a single entity emanating or producing something by his or herself, as in the Tibetan
Buddhism concept of Adi-Buddha, the ancient Greek story of Gaia (Mother Earth), the Aztec goddess
Coatlicue myth, the ancient Egyptian god Atum story, or the Genesis creation narrative. In another type of
story, the world is created from the union of male and female deities, as in the Maori story of Rangi and
Papa. In other stories, the universe is created by crafting it from pre-existing materials, such as the corpse of
a dead god — as from Tiamat in the Babylonian epic Enuma Elish or from the giant Ymir in Norse mythology –
or from chaotic materials, as in Izanagi and Izanami in Japanese mythology. In other stories, the universe
emanates from fundamental principles, such as Brahman and Prakrti, or the yin and yang of the Tao.

[edit] Philosophical models

Further information: Cosmology
See also: Pre-Socratic philosophy, Physics (Aristotle), Hindu cosmology, Islamic cosmology, and
Time
From the 6th century BCE, the pre-Socratic Greek philosophers developed the earliest known
philosophical models of the universe. The earliest Greek philosophers noted that appearances can be
deceiving, and sought to understand the underlying reality behind the appearances. In particular, they noted
the ability of matter to change forms (e.g., ice to water to steam) and several philosophers proposed that all
the apparently different materials of the world (wood, metal, etc.) are all different forms of a single material,
the arche. The first to do so was Thales, who called this material Water. Following him, Anaximenes called it
Air, and posited that there must be attractive and repulsive forces that cause the arche to condense or
dissociate into different forms. Empedocles proposed that multiple fundamental materials were necessary to
explain the diversity of the universe, and proposed that all four classical elements (Earth, Air, Fire and Water)
existed, albeit in different combinations and forms. This four-element theory was adopted by many of the
subsequent philosophers. Some philosophers before Empedocles advocated less material things for the
arche; Heraclitus argued for a Logos, Pythagoras believed that all things were composed of numbers,
whereas Thales' student, Anaximander, proposed that everything was composed of a chaotic substance
known as apeiron, roughly corresponding to the modern concept of a quantum foam. Various modifications of
the apeiron theory were proposed, most notably that of Anaxagoras, which proposed that the various matter
in the world was spun off from a rapidly rotating apeiron, set in motion by the principle of Nous (Mind). Still
other philosophers — most notably Leucippus and Democritus — proposed that the universe was composed of
indivisible atoms moving through empty space, a vacuum; Aristotle opposed this view ("Nature abhors a
vacuum") on the grounds that resistance to motion increases with density; hence, empty space should offer
no resistance to motion, leading to the possibility of infinite speed.
Although Heraclitus argued for eternal change, his quasi-contemporary Parmenides made the radical
suggestion that all change is an illusion, that the true underlying reality is eternally unchanging and of a
single nature. Parmenides denoted this reality as τὸ ἐν (The One). Parmenides' theory seemed implausible
to many Greeks, but his student Zeno of Elea challenged them with several famous paradoxes. Aristotle
resolved these paradoxes by developing the notion of an infinitely divisible continuum, and applying it to
space and time.
The Indian philosopher Kanada, founder of the Vaisheshika school, developed a theory of atomism
and proposed that light and heat were varieties of the same substance.[42] In the 5th century AD, the
Buddhist atomist philosopher Dignāga proposed atoms to be point-sized, durationless, and made of energy.
They denied the existence of substantial matter and proposed that movement consisted of momentary
flashes of a stream of energy.[43]
The theory of temporal finitism was inspired by the doctrine of creation shared by the three
Abrahamic religions: Judaism, Christianity and Islam. The Christian philosopher, John Philoponus, presented
the philosophical arguments against the ancient Greek notion of an infinite past. Philoponus' arguments
against an infinite past were used by the early Muslim philosopher, Al-Kindi (Alkindus); the Jewish
philosopher, Saadia Gaon (Saadia ben Joseph); and the Muslim theologian, Al-Ghazali (Algazel). They
employed two logical arguments against an infinite past, the first being the "argument from the impossibility
of the existence of an actual infinite", which states:[44]
"An actual infinite cannot exist."
"An infinite temporal regress of events is an actual infinite."

" An infinite temporal regress of events cannot exist."

The second argument, the "argument from the impossibility of completing an actual infinite by
successive addition", states:[44]
"An actual infinite cannot be completed by successive addition."
"The temporal series of past events has been completed by successive addition."

" The temporal series of past events cannot be an actual infinite."

 Both arguments were adopted by later Christian philosophers and theologians, and the second
argument in particular became more famous after it was adopted by Immanuel Kant in his thesis of the first
antinomy concerning time.[44]

[edit] Astronomical models

Main article: History of astronomy
Astronomical models of the universe were proposed soon after astronomy began with the Babylonian
astronomers, who viewed the universe as a flat disk floating in the ocean, and this forms the premise for early
Greek maps like those of Anaximander and Hecataeus of Miletus.
Later Greek philosophers, observing the motions of the heavenly bodies, were concerned with
developing models of the universe based more profoundly on empirical evidence. The first coherent model
was proposed by Eudoxus of Cnidos. According to this model, space and time are infinite and eternal, the
Earth is spherical and stationary, and all other matter is confined to rotating concentric spheres. This model
was refined by Callippus and Aristotle, and brought into nearly perfect agreement with astronomical
observations by Ptolemy. The success of this model is largely due to the mathematical fact that any function
(such as the position of a planet) can be decomposed into a set of circular functions (the Fourier modes).
However, not all Greek scientists accepted the geocentric model of the universe. The Pythagorean
philosopher Philolaus postulated that at the center of the universe was a "central fire" around which the
Earth, Sun, Moon and Planets revolved in uniform circular motion.[45] The Greek astronomer Aristarchus of
Samos was the first known individual to propose a heliocentric model of the universe. Though the original
text has been lost, a reference in Archimedes' book The Sand Reckoner describes Aristarchus' heliocentric
theory. Archimedes wrote: (translated into English)
 You King Gelon are aware the 'universe' is the name given by most astronomers to the sphere
the center of which is the center of the Earth, while its radius is equal to the straight line between
the center of the Sun and the center of the Earth. This is the common account as you have
heard from astronomers. But Aristarchus has brought out a book consisting of certain
hypotheses, wherein it appears, as a consequence of the assumptions made, that the universe
is many times greater than the 'universe' just mentioned. His hypotheses are that the fixed stars
and the Sun remain unmoved, that the Earth revolves about the Sun on the circumference of a
circle, the Sun lying in the middle of the orbit, and that the sphere of fixed stars, situated about
the same center as the Sun, is so great that the circle in which he supposes the Earth to revolve
bears such a proportion to the distance of the fixed stars as the center of the sphere bears to its
surface.

Aristarchus thus believed the stars to be very far away, and saw this as the reason why there was no
visible parallax, that is, an observed movement of the stars relative to each other as the Earth moved around
the Sun. The stars are in fact much farther away than the distance that was generally assumed in ancient
times, which is why stellar parallax is only detectable with telescopes. The geocentric model, consistent with
planetary parallax, was assumed to be an explanation for the unobservability of the parallel phenomenon,
stellar parallax. The rejection of the heliocentric view was apparently quite strong, as the following passage
from Plutarch suggests (On the Apparent Face in the Orb of the Moon):
Cleanthes [a contemporary of Aristarchus and head of the Stoics] thought it was the duty of the
Greeks to indict Aristarchus of Samos on the charge of impiety for putting in motion the Hearth
of the universe [i.e. the earth], . . . supposing the heaven to remain at rest and the earth to
revolve in an oblique circle, while it rotates, at the same time, about its own axis. [1]
 The only other astronomer from antiquity known by name who supported Aristarchus' heliocentric
model was Seleucus of Seleucia, a Hellenized Babylonian astronomer who lived a century after Aristarchus.
[46][47][48] According to Plutarch, Seleucus was the first to prove the heliocentric system through reasoning,
but it is not known what arguments he used. Seleucus' arguments for a heliocentric theory were probably
related to the phenomenon of tides.[49] According to Strabo (1.1.9), Seleucus was the first to state that the
tides are due to the attraction of the Moon, and that the height of the tides depends on the Moon's position
relative to the Sun.[50] Alternatively, he may have proved the heliocentric theory by determining the
constants of a geometric model for the heliocentric theory and by developing methods to compute planetary
positions using this model, like what Nicolaus Copernicus later did in the 16th century.[51] During the Middle
Ages, heliocentric models may have also been proposed by the Indian astronomer, Aryabhata,[52] and by
the Persian astronomers, Albumasar[53] and Al-Sijzi.[54]
 Model of the Copernican universe by Thomas Digges in 1576, with the amendment that the stars are
no longer confined to a sphere, but spread uniformly throughout the space surrounding the planets.
The Aristotelian model was accepted in the Western world for roughly two millennia, until Copernicus
revived Aristarchus' theory that the astronomical data could be explained more plausibly if the earth rotated
on its axis and if the sun were placed at the center of the universe.
“ In the center rests the sun. For who would place this lamp of a very beautiful ”
temple in another or better place than this wherefrom it can illuminate everything at the
 same time?

—Copernicus, in Chapter 10, Book 1 of De Revolutionibus Orbium Coelestrum (1543)

As noted by Copernicus himself, the suggestion that the Earth rotates was very old, dating at least to
Philolaus (c. 450 BC), Heraclides Ponticus (c. 350 BC) and Ecphantus the Pythagorean. Roughly a century
before Copernicus, Christian scholar Nicholas of Cusa also proposed that the Earth rotates on its axis in his
book, On Learned Ignorance (1440).[55] Aryabhata (476–550), Brahmagupta (598–668), Albumasar and Al-
Sijzi, also proposed that the Earth rotates on its axis.[ citation needed] The first empirical evidence for the
Earth's rotation on its axis, using the phenomenon of comets, was given by Tusi (1201–1274) and Ali Kuşçu
(1403–1474).[citation needed] Tusi, however, continued to support the Aristotelian universe, thus Kuşçu was
the first to refute the Aristotelian notion of a stationary Earth on an empirical basis, similar to how Copernicus
later justified the Earth's rotation. Al-Birjandi (d. 1528) further developed a theory of "circular inertia" to
explain the Earth's rotation, similar to how Galileo Galilei explained it.[56][57]
 Johannes Kepler published the Rudolphine Tables containing a star catalog and planetary tables
using Tycho Brahe's measurements.
Copernicus' heliocentric model allowed the stars to be placed uniformly through the (infinite) space
surrounding the planets, as first proposed by Thomas Digges in his Perfit Description of the Caelestiall Orbes
according to the most aunciente doctrine of the Pythagoreans, latelye revived by Copernicus and by
Geometricall Demonstrations approved (1576).[58] Giordano Bruno accepted the idea that space was infinite
and filled with solar systems similar to our own; for the publication of this view, he was burned at the stake in
the Campo dei Fiori in Rome on 17 February 1600.[58]
This cosmology was accepted provisionally by Isaac Newton, Christiaan Huygens and later
scientists,[58] although it had several paradoxes that were resolved only with the development of general
relativity. The first of these was that it assumed that space and time were infinite, and that the stars in the
universe had been burning forever; however, since stars are constantly radiating energy, a finite star seems
inconsistent with the radiation of infinite energy. Secondly, Edmund Halley (1720)[59] and Jean-Philippe de
Cheseaux (1744)[60] noted independently that the assumption of an infinite space filled uniformly with stars
would lead to the prediction that the nighttime sky would be as bright as the sun itself; this became known as
Olbers' paradox in the 19th century.[61] Third, Newton himself showed that an infinite space uniformly filled
with matter would cause infinite forces and instabilities causing the matter to be crushed inwards under its
own gravity.[58] This instability was clarified in 1902 by the Jeans instability criterion.[62] One solution to
these latter two paradoxes is the Charlier universe, in which the matter is arranged hierarchically (systems of
orbiting bodies that are themselves orbiting in a larger system, ad infinitum) in a fractal way such that the
universe has a negligibly small overall density; such a cosmological model had also been proposed earlier in
1761 by Johann Heinrich Lambert.[63] A significant astronomical advance of the 18th century was the
realization by Thomas Wright, Immanuel Kant and others that stars are not distributed uniformly throughout
space; rather, they are grouped into galaxies.[64]
The modern era of physical cosmology began in 1917, when Albert Einstein first applied his general
theory of relativity to model the structure and dynamics of the universe.[65] This theory and its implications
will be discussed in more detail in the following section.
[edit] Theoretical models
 High-precision test of general relativity by the Cassini space probe (artist's impression): radio signals
sent between the Earth and the probe (green wave) are delayed by the warping of space and time (blue
lines) due to the Sun's mass.
Of the four fundamental interactions, gravitation is dominant at cosmological length scales; that is,
the other three forces are believed to play a negligible role in determining structures at the level of planets,
stars, galaxies and larger-scale structures. Since all matter and energy gravitate, gravity's effects are
cumulative; by contrast, the effects of positive and negative charges tend to cancel one another, making
electromagnetism relatively insignificant on cosmological length scales. The remaining two interactions, the
weak and strong nuclear forces, decline very rapidly with distance; their effects are confined mainly to sub-
atomic length scales.

[edit] General theory of relativity

Main articles: Introduction to general relativity, General relativity, and Einstein's field equations
Given gravitation's predominance in shaping cosmological structures, accurate predictions of the
universe's past and future require an accurate theory of gravitation. The best theory available is Albert
Einstein's general theory of relativity, which has passed all experimental tests hitherto. However, since
rigorous experiments have not been carried out on cosmological length scales, general relativity could
conceivably be inaccurate. Nevertheless, its cosmological predictions appear to be consistent with
observations, so there is no compelling reason to adopt another theory.
General relativity provides of a set of ten nonlinear partial differential equations for the spacetime
metric (Einstein's field equations) that must be solved from the distribution of mass-energy and momentum
throughout the universe. Since these are unknown in exact detail, cosmological models have been based on
the cosmological principle, which states that the universe is homogeneous and isotropic. In effect, this
principle asserts that the gravitational effects of the various galaxies making up the universe are equivalent to
those of a fine dust distributed uniformly throughout the universe with the same average density. The
assumption of a uniform dust makes it easy to solve Einstein's field equations and predict the past and future
of the universe on cosmological time scales.
Einstein's field equations include a cosmological constant (Λ),[65][66] that corresponds to an energy
density of empty space.[67] Depending on its sign, the cosmological constant can either slow (negative Λ) or
accelerate (positive Λ) the expansion of the universe. Although many scientists, including Einstein, had
speculated that Λ was zero,[68] recent astronomical observations of type Ia supernovae have detected a
large amount of "dark energy" that is accelerating the universe's expansion.[69] Preliminary studies suggest
that this dark energy corresponds to a positive Λ, although alternative theories cannot be ruled out as yet.[70]
Russian physicist Zel'dovich suggested that Λ is a measure of the zero-point energy associated with virtual
particles of quantum field theory, a pervasive vacuum energy that exists everywhere, even in empty space.
[71] Evidence for such zero-point energy is observed in the Casimir effect.

[edit] Special relativity and space-time

Main articles: Introduction to special relativity and Special relativity
 Only its length L is intrinsic to the rod (shown in black); coordinate differences between its endpoints
(such as Δx, Δy or Δξ, Δη) depend on their frame of reference (depicted in blue and red, respectively).
 The universe has at least three spatial and one temporal (time) dimension. It was long thought that
the spatial and temporal dimensions were different in nature and independent of one another. However,
according to the special theory of relativity, spatial and temporal separations are interconvertible (within
limits) by changing one's motion.
To understand this interconversion, it is helpful to consider the analogous interconversion of spatial
separations along the three spatial dimensions. Consider the two endpoints of a rod of length L. The length
can be determined from the differences in the three coordinates Δx, Δy and Δz of the two endpoints in a
given reference frame
L2 = Δx2 + Δy2 + Δz2
using the Pythagorean theorem. In a rotated reference frame, the coordinate differences differ, but
they give the same length
L2 = Δξ2 + Δη2 + Δζ2.
Thus, the coordinates differences (Δx, Δy, Δz) and (Δξ, Δη, Δζ) are not intrinsic to the rod, but merely
reflect the reference frame used to describe it; by contrast, the length L is an intrinsic property of the rod. The
coordinate differences can be changed without affecting the rod, by rotating one's reference frame.
The analogy in spacetime is called the interval between two events; an event is defined as a point in
spacetime, a specific position in space and a specific moment in time. The spacetime interval between two
events is given by
 where c is the speed of light. According to special relativity, one can change a spatial and time
separation (L1, Δt1) into another (L2, Δt2) by changing one's reference frame, as long as the change
maintains the spacetime interval s. Such a change in reference frame corresponds to changing one's motion;
in a moving frame, lengths and times are different from their counterparts in a stationary reference frame.
The precise manner in which the coordinate and time differences change with motion is described by the
Lorentz transformation.

[edit] Solving Einstein's field equations

See also: Big Bang and Ultimate fate of the universe
The distances between the spinning galaxies increase with time, but the distances between the stars
within each galaxy stay roughly the same, due to their gravitational interactions. This animation illustrates a
closed Friedmann universe with zero cosmological constant Λ; such a universe oscillates between a Big
Bang and a Big Crunch.
 Animation illustrating the metric expansion of the universe
In non-Cartesian (non-square) or curved coordinate systems, the Pythagorean theorem holds only on
infinitesimal length scales and must be augmented with a more general metric tensor gμν, which can vary
from place to place and which describes the local geometry in the particular coordinate system. However,
assuming the cosmological principle that the universe is homogeneous and isotropic everywhere, every point
in space is like every other point; hence, the metric tensor must be the same everywhere. That leads to a
single form for the metric tensor, called the Friedmann-Lemaître-Robertson-Walker metric

where (r, θ, φ) correspond to a spherical coordinate system. This metric has only two undetermined
parameters: an overall length scale R that can vary with time, and a curvature index k that can be only 0, 1 or
−1, corresponding to flat Euclidean geometry, or spaces of positive or negative curvature. In cosmology,
solving for the history of the universe is done by calculating R as a function of time, given k and the value of
the cosmological constant Λ, which is a (small) parameter in Einstein's field equations. The equation
describing how R varies with time is known as the Friedmann equation, after its inventor, Alexander
Friedmann.[72]
The solutions for R(t) depend on k and Λ, but some qualitative features of such solutions are general.
First and most importantly, the length scale R of the universe can remain constant only if the universe is
perfectly isotropic with positive curvature (k=1) and has one precise value of density everywhere, as first
noted by Albert Einstein. However, this equilibrium is unstable and since the universe is known to be
inhomogeneous on smaller scales, R must change, according to general relativity. When R changes, all the
spatial distances in the universe change in tandem; there is an overall expansion or contraction of space
itself. This accounts for the observation that galaxies appear to be flying apart; the space between them is
stretching. The stretching of space also accounts for the apparent paradox that two galaxies can be 40 billion
light years apart, although they started from the same point 13.7 billion years ago and never moved faster
than the speed of light.
 Second, all solutions suggest that there was a gravitational singularity in the past, when R goes to
zero and matter and energy became infinitely dense. It may seem that this conclusion is uncertain since it is
based on the questionable assumptions of perfect homogeneity and isotropy (the cosmological principle) and
that only the gravitational interaction is significant. However, the Penrose-Hawking singularity theorems show
that a singularity should exist for very general conditions. Hence, according to Einstein's field equations, R
grew rapidly from an unimaginably hot, dense state that existed immediately following this singularity (when
R had a small, finite value); this is the essence of the Big Bang model of the universe. A common
misconception is that the Big Bang model predicts that matter and energy exploded from a single point in
space and time; that is false. Rather, space itself was created in the Big Bang and imbued with a fixed
amount of energy and matter distributed uniformly throughout; as space expands (i.e., as R(t) increases), the
density of that matter and energy decreases.
Space has no boundary – that is empirically more certain than any external observation. However,
that does not imply that space is infinite...(translated, original German)

Bernhard Riemann (Habilitationsvortrag, 1854)

Third, the curvature index k determines the sign of the mean spatial curvature of spacetime averaged
over length scales greater than a billion light years. If k=1, the curvature is positive and the universe has a
finite volume. Such universes are often visualized as a three-dimensional sphere S3 embedded in a four-
dimensional space. Conversely, if k is zero or negative, the universe may have infinite volume, depending on
its overall topology. It may seem counter-intuitive that an infinite and yet infinitely dense universe could be
created in a single instant at the Big Bang when R=0, but exactly that is predicted mathematically when k
does not equal 1. For comparison, an infinite plane has zero curvature but infinite area, whereas an infinite
cylinder is finite in one direction and a torus is finite in both. A toroidal universe could behave like a normal
universe with periodic boundary conditions, as seen in "wrap-around" video games such as Asteroids; a
traveler crossing an outer "boundary" of space going outwards would reappear instantly at another point on
the boundary moving inwards.
 Prevailing model of the origin and expansion of spacetime and all that it contains.
The ultimate fate of the universe is still unknown, because it depends critically on the curvature index
k and the cosmological constant Λ. If the universe is sufficiently dense, k equals +1, meaning that its average
curvature throughout is positive and the universe will eventually recollapse in a Big Crunch, possibly starting
a new universe in a Big Bounce. Conversely, if the universe is insufficiently dense, k equals 0 or −1 and the
universe will expand forever, cooling off and eventually becoming inhospitable for all life, as the stars die and
all matter coalesces into black holes (the Big Freeze and the heat death of the universe). As noted above,
recent data suggests that the expansion speed of the universe is not decreasing as originally expected, but
increasing; if this continues indefinitely, the universe will eventually rip itself to shreds (the Big Rip).
Experimentally, the universe has an overall density that is very close to the critical value between recollapse
and eternal expansion; more careful astronomical observations are needed to resolve the question.

[edit] Big Bang model

Main articles: Big Bang, Timeline of the Big Bang, Nucleosynthesis, and Lambda-CDM model
The prevailing Big Bang model accounts for many of the experimental observations described above,
such as the correlation of distance and redshift of galaxies, the universal ratio of hydrogen:helium atoms, and
the ubiquitous, isotropic microwave radiation background. As noted above, the redshift arises from the metric
expansion of space; as the space itself expands, the wavelength of a photon traveling through space likewise
increases, decreasing its energy. The longer a photon has been traveling, the more expansion it has
undergone; hence, older photons from more distant galaxies are the most red-shifted. Determining the
correlation between distance and redshift is an important problem in experimental physical cosmology.
 Chief nuclear reactions responsible for the relative abundances of light atomic nuclei observed
throughout the universe.
Other experimental observations can be explained by combining the overall expansion of space with
nuclear and atomic physics. As the universe expands, the energy density of the electromagnetic radiation
decreases more quickly than does that of matter, since the energy of a photon decreases with its wavelength.
Thus, although the energy density of the universe is now dominated by matter, it was once dominated by
radiation; poetically speaking, all was light. As the universe expanded, its energy density decreased and it
became cooler; as it did so, the elementary particles of matter could associate stably into ever larger
combinations. Thus, in the early part of the matter-dominated era, stable protons and neutrons formed, which
then associated into atomic nuclei. At this stage, the matter in the universe was mainly a hot, dense plasma
of negative electrons, neutral neutrinos and positive nuclei. Nuclear reactions among the nuclei led to the
present abundances of the lighter nuclei, particularly hydrogen, deuterium, and helium. Eventually, the
electrons and nuclei combined to form stable atoms, which are transparent to most wavelengths of radiation;
at this point, the radiation decoupled from the matter, forming the ubiquitous, isotropic background of
microwave radiation observed today.
Other observations are not answered definitively by known physics. According to the prevailing
theory, a slight imbalance of matter over antimatter was present in the universe's creation, or developed very
shortly thereafter, possibly due to the CP violation that has been observed by particle physicists. Although
the matter and antimatter mostly annihilated one another, producing photons, a small residue of matter
survived, giving the present matter-dominated universe. Several lines of evidence also suggest that a rapid
cosmic inflation of the universe occurred very early in its history (roughly 10−35 seconds after its creation).
Recent observations also suggest that the cosmological constant (Λ) is not zero and that the net mass-
energy content of the universe is dominated by a dark energy and dark matter that have not been
characterized scientifically. They differ in their gravitational effects. Dark matter gravitates as ordinary matter
does, and thus slows the expansion of the universe; by contrast, dark energy serves to accelerate the
universe's expansion.

[edit] Untestable proposals

[edit] Multiverse theory
Main articles: Multiverse, Many-worlds hypothesis, Bubble universe theory, and Parallel universe
(fiction)
 Depiction of a multiverse of seven "bubble" universes, which are separate spacetime continua, each
having different physical laws, physical constants, and perhaps even different numbers of dimensions or
topologies.
Some speculative theories have proposed that this universe is but one of a set of disconnected
universes, collectively denoted as the multiverse, altering the concept that the universe encompasses
everything.[15][73] By definition, there is no possible way for anything in one universe to affect another; if two
"universes" could affect one another, they would be part of a single universe. Thus, although some fictional
characters travel between parallel fictional "universes", this is, strictly speaking, an incorrect usage of the
term universe. The disconnected universes are conceived as being physical, in the sense that each should
have its own space and time, its own matter and energy, and its own physical laws — that also challenges the
definition of parallelity as these universes don't exist synchronously (since they have their own time) or in a
geometrically parallel way (since there's no interpretable relation between spatial positions of the different
universes). Such physically disconnected universes should be distinguished from the metaphysical
conception of alternate planes of consciousness, which are not thought to be physical places and are
connected through the flow of information. The concept of a multiverse of disconnected universes is very old;
for example, Bishop Étienne Tempier of Paris ruled in 1277 that God could create as many universes as he
saw fit, a question that was being hotly debated by the French theologians.[74]
There are two scientific senses in which multiple universes are discussed. First, disconnected
spacetime continua may exist; presumably, all forms of matter and energy are confined to one universe and
cannot "tunnel" between them. An example of such a theory is the chaotic inflation model of the early
universe.[75] Second, according to the many-worlds hypothesis, a parallel universe is born with every
quantum measurement; the universe "forks" into parallel copies, each one corresponding to a different
outcome of the quantum measurement. However, both senses of the term "multiverse" are speculative and
may be considered unscientific; no known experimental test in one universe could reveal the existence or
properties of another non-interacting universe.

[edit] Shape of the universe

For more details on this topic, see Shape of the universe.
The shape or geometry of the universe includes both local geometry in the observable universe and
global geometry, which we may or may not be able to measure. Shape can refer to curvature and topology.
More formally, the subject in practice investigates which 3-manifold corresponds to the spatial section in
comoving coordinates of the four-dimensional space-time of the universe. Analysis of data from WMAP
implies that the universe is spatially flat with only a 2% margin of error.[76]
Cosmologists normally work with a given space-like slice of spacetime called the comoving
coordinates. In terms of observation, the section of spacetime that can be observed is the backward light
cone (points within the cosmic light horizon, given time to reach a given observer). If the observable universe
is smaller than the entire universe (in some models it is many orders of magnitude smaller), one cannot
determine the global structure by observation: one is limited to a small patch.
In October 2001, NASA began collecting data with the Wilkinson Microwave Anisotropy Probe
(WMAP) on cosmic background radiation. Like visible light from distant stars and galaxies, cosmic
background radiation allows scientists to peer into the past to the time when the universe was in its infancy.
Density fluctuations in this radiation can also tell scientists much about the physical nature of space.[77]
NASA released the first WMAP cosmic background radiation data in February 2003. In 2009 the Planck
observatory was launched which will be able to analyze the microwave background at higher resolution,
providing more information on the shape of the early universe. The preliminary data will be released in
December 2010.

[edit] See also

Astronomy portal

Space portal

• Anthropic principle
• Big Bang
• Big Crunch
• Cosmic latte
• Cosmology
• Dyson's eternal intelligence
• Esoteric cosmology
• False vacuum
• Final anthropic principle
• Fine-tuned universe
• Heat death of the universe
• Hindu cycle of the universe
• Kardashev scale
• Multiverse
• Nucleocosmochronology
• Non-standard cosmology
• Omega Point
• Omniverse
• Rare Earth hypothesis
• Reality
• Shape of the universe
• Ultimate fate of the universe
• Vacuum genesis
• World view
• Zero-energy universe
[edit] Notes and references
1. ^ Webster's New World College Dictionary. Wiley Publishing, Inc.. 2010.
http://www.yourdictionary.com/universe.
2. ^ The American Heritage® Dictionary of the English Language (4th ed.). Houghton Mifflin
Harcourt Publishing Company. 2010. http://www.yourdictionary.com/universe.
3. ^ Cambridge Advanced Learner's Dictionary.
http://dictionary.cambridge.org/dictionary/british/universe.
4. ^ S. H. Suyu, P. J. Marshall, M. W. Auger, S. Hilbert, R. D. Blandford, L. V. E. Koopmans, C.
D. Fassnacht and T. Treu. Dissecting the Gravitational Lens B1608+656. II. Precision Measurements
of the Hubble Constant, Spatial Curvature, and the Dark Energy Equation of State. The Astrophysical
Journal, 2010; 711 (1): 201 DOI: 10.1088/0004-637X/711/1/201
5. ^ Lineweaver, Charles; Tamara M. Davis (2005). "Misconceptions about the Big Bang".
Scientific American. http://www.sciam.com/article.cfm?id=misconceptions-about-the-2005-
03&page=5. Retrieved 2008-11-06.
6. ^ The Compact Edition of the Oxford English Dictionary , volume II, Oxford: Oxford University
Press, 1971, p.3518.
7. ^ a b Lewis and Short, A Latin Dictionary, Oxford University Press, ISBN 0-19-864201-6, pp.
1933, 1977–1978.
8. ^ Liddell and Scott, A Greek-English Lexicon, Oxford University Press, ISBN 0-19-864214-8,
p.1392.
9. ^ Liddell and Scott, pp.1345–1346.
10.^ Yonge, Charles Duke (1870). An English-Greek lexicon. New York: American Bok
Company. pp. 567.
 11.^ Liddell and Scott, pp.985, 1964.
12.^ Lewis and Short, pp. 1881–1882, 1175, 1189–1190.
13.^ OED, pp. 909, 569, 3821–3822, 1900.
14.^ Feynman RP, Hibbs AR (1965). Quantum Physics and Path Integrals. New York: McGraw–
Hill. ISBN 0-07-020650-3.
Zinn Justin J (2004). Path Integrals in Quantum Mechanics. Oxford University Press. ISBN 0-19-
856674-3. OCLC 212409192.
15.^ a b Ellis, George F.R.; U. Kirchner, W.R. Stoeger (2004). "Multiverses and physical
cosmology" (subscription required). Monthly Notices of the Royal Astronomical Society 347: 921–936.
doi:10.1111/j.1365-2966.2004.07261.x. http://arxiv.org/abs/astro-ph/0305292. Retrieved 2007-01-
09.
16.^ Lineweaver, Charles; Tamara M. Davis (2005). "Misconceptions about the Big Bang".
Scientific American. http://www.sciam.com/article.cfm?articleID=0009F0CA-C523-1213-
852383414B7F0147&pageNumber=5. Retrieved 2007-03-05.
17.^ Rindler (1977), p.196.
18.^ Christian, Eric; Samar, Safi-Harb. "How large is the Milky Way?".
http://imagine.gsfc.nasa.gov/docs/ask_astro/answers/980317b.html. Retrieved 2007-11-28.
19.^ I. Ribas, C. Jordi, F. Vilardell, E.L. Fitzpatrick, R.W. Hilditch, F. Edward (2005). "First
Determination of the Distance and Fundamental Properties of an Eclipsing Binary in the Andromeda
Galaxy". Astrophysical Journal 635: L37–L40. doi:10.1086/499161.
http://adsabs.harvard.edu/abs/2005ApJ...635L..37R.
McConnachie, A. W.; Irwin, M. J.; Ferguson, A. M. N.; Ibata, R. A.; Lewis, G. F.; Tanvir, N. (2005).
"Distances and metallicities for 17 Local Group galaxies". Monthly Notices of the Royal Astronomical
Society 356 (4): 979–997. doi:10.1111/j.1365-2966.2004.08514.x. http://adsabs.harvard.edu/cgi-
bin/nph-bib_query?bibcode=2005MNRAS.356..979M.
20.^ Mackie, Glen (February 1, 2002). "To see the Universe in a Grain of Taranaki Sand".
Swinburne University. http://astronomy.swin.edu.au/~gmackie/billions.html. Retrieved 2006-12-20.
21.^ "Unveiling the Secret of a Virgo Dwarf Galaxy". ESO. 2000-05-03.
http://www.eso.org/outreach/press-rel/pr-2000/pr-12-00.html. Retrieved 2007-01-03.
22.^ "Hubble's Largest Galaxy Portrait Offers a New High-Definition View". NASA. 2006-02-28.
http://www.nasa.gov/mission_pages/hubble/science/hst_spiral_m10.html. Retrieved 2007-01-03.
23.^ "Star Count: ANU Astronomer makes best yet". 2003-07-17.
http://info.anu.edu.au/ovc/media/Media_Releases/2003/030717StarCount. Retrieved 2010-02-19.
24.^ N. Mandolesi, P. Calzolari, S. Cortiglioni, F. Delpino, G. Sironi (1986). "Large-scale
homogeneity of the Universe measured by the microwave background". Letters to Nature 319: 751–
753. doi:10.1038/319751a0.
25.^ Hinshaw, Gary (November 29, 2006). "New Three Year Results on the Oldest Light in the
Universe". NASA WMAP. http://map.gsfc.nasa.gov/m_mm.html. Retrieved 2006-08-10.
26.^ Hinshaw, Gary (December 15, 2005). "Tests of the Big Bang: The CMB". NASA WMAP.
http://map.gsfc.nasa.gov/m_uni/uni_101bbtest3.html. Retrieved 2007-01-09.
27.^ Rindler (1977), p. 202.
28.^ Hinshaw, Gary (February 10, 2006). "What is the Universe Made Of?". NASA WMAP.
http://map.gsfc.nasa.gov/m_uni/uni_101matter.html. Retrieved 2007-01-04.
29.^ "Five-Year Wilkinson Microwave Anisotropy Probe (WMAP) Observations: Data
Processing, Sky Maps, and Basic Results" (PDF). nasa.gov.
http://lambda.gsfc.nasa.gov/product/map/dr3/pub_papers/fiveyear/basic_results/wmap5basic.pdf.
Retrieved 2008-03-06.
 30.^ Britt RR (2003-01-03). "Age of Universe Revised, Again". space.com.
http://www.space.com/scienceastronomy/age_universe_030103.html. Retrieved 2007-01-08.
31.^ Wright EL (2005). "Age of the Universe". UCLA.
http://www.astro.ucla.edu/~wright/age.html. Retrieved 2007-01-08.
Krauss LM, Chaboyer B (3 January 2003). "Age Estimates of Globular Clusters in the Milky Way:
Constraints on Cosmology". Science (American Association for the Advancement of Science) 299
(5603): 65–69. doi:10.1126/science.1075631. PMID 12511641.
http://www.sciencemag.org/cgi/content/abstract/299/5603/65?
ijkey=3D7y0Qonz=GO7ig.&keytype=3Dref&siteid=3Dsci. Retrieved 2007-01-08.
32.^ Wright, Edward L. (September 12, 2004). "Big Bang Nucleosynthesis". UCLA.
http://www.astro.ucla.edu/~wright/BBNS.html. Retrieved 2007-01-05.
M. Harwit, M. Spaans (2003). "Chemical Composition of the Early Universe". The Astrophysical
Journal 589 (1): 53–57. doi:10.1086/374415. http://adsabs.harvard.edu/abs/2003ApJ...589...53H.
C. Kobulnicky, E. D. Skillman (1997). "Chemical Composition of the Early Universe". Bulletin of the
American Astronomical Society 29: 1329. http://adsabs.harvard.edu/abs/1997AAS...191.7603K.
33.^ "Antimatter". Particle Physics and Astronomy Research Council. October 28, 2003.
http://www.pparc.ac.uk/ps/bbs/bbs_antimatter.asp. Retrieved 2006-08-10.
34.^ Landau and Lifshitz (1975), p.361.
35.^ WMAP Mission: Results – Age of the Universe
36.^ Luminet, Jean-Pierre; Boudewijn F. Roukema (1999). "Topology of the Universe: Theory
and Observations". Proceedings of Cosmology School held at Cargese, Corsica, August 1998 .
http://arxiv.org/abs/astro-ph/9901364. Retrieved 2007-01-05.
Luminet, Jean-Pierre; J. Weeks, A. Riazuelo, R. Lehoucq, J.-P. Uzan (2003). "Dodecahedral space
topology as an explanation for weak wide-angle temperature correlations in the cosmic microwave
background" (subscription required). Nature 425 (6958): 593. doi:10.1038/nature01944.
PMID 14534579. http://arxiv.org/abs/astro-ph/0310253. Retrieved 2007-01-09.
37.^ Strobel, Nick (May 23, 2001). "The Composition of Stars". Astronomy Notes.
http://www.astronomynotes.com/starprop/s7.htm. Retrieved 2007-01-04.
"Have physical constants changed with time?". Astrophysics (Astronomy Frequently Asked
Questions). http://www.faqs.org/faqs/astronomy/faq/part4/section-4.html. Retrieved 2007-01-04.
38.^ Stephen Hawking (1988). A Brief History of Time. Bantam Books. p. 125. ISBN 0-553-
05340-X.
39.^ Martin Rees (1999). Just Six Numbers. HarperCollins Publishers. ISBN 0-465-03672-4.
40.^ Adams, F.C. (2008). "Stars in other universes: stellar structure with different fundamental
constants". Journal of Cosmology and Astroparticle Physics 2008 (08): 010. doi:10.1088/1475-
7516/2008/08/010. http://arxiv.org/abs/0807.3697.
41.^ Harnik, R.; Kribs, G.D. and Perez, G. (2006). "A universe without weak interactions".
Physical Review D 74: 035006. doi:10.1103/PhysRevD.74.035006. http://arxiv.org/abs/hep-
ph/0604027.
42.^ Will Durant, Our Oriental Heritage:
"Two systems of Hindu thought propound physical theories suggestively similar to those
of Greece. Kanada, founder of the Vaisheshika philosophy, held that the world was
composed of atoms as many in kind as the various elements. The Jains more nearly
approximated to Democritus by teaching that all atoms were of the same kind,
producing different effects by diverse modes of combinations. Kanada believed light and
heat to be varieties of the same substance; Udayana taught that all heat comes from the
sun; and Vachaspati, like Newton, interpreted light as composed of minute particles
emitted by substances and striking the eye."
 43.^ F. Th. Stcherbatsky (1930, 1962), Buddhist Logic, Volume 1, p.19, Dover, New York:
"The Buddhists denied the existence of substantial matter altogether. Movement
consists for them of moments, it is a staccato movement, momentary flashes of a
stream of energy... "Everything is evanescent“,... says the Buddhist, because there is no
stuff... Both systems [Sānkhya, and later Indian Buddhism] share in common a tendency
to push the analysis of Existence up to its minutest, last elements which are imagined as
absolute qualities, or things possessing only one unique quality. They are called
“qualities” (guna-dharma) in both systems in the sense of absolute qualities, a kind of
atomic, or intra-atomic, energies of which the empirical things are composed. Both
systems, therefore, agree in denying the objective reality of the categories of Substance
and Quality,... and of the relation of Inference uniting them. There is in Sānkhya
philosophy no separate existence of qualities. What we call quality is but a particular
manifestation of a subtle entity. To every new unit of quality corresponds a subtle
quantum of matter which is called guna “quality”, but represents a subtle substantive
entity. The same applies to early Buddhism where all qualities are substantive... or,
more precisely, dynamic entities, although they are also called dharmas ('qualities')."

44.^ a b c Craig, William Lane (June 1979). "Whitrow and Popper on the Impossibility of an
Infinite Past". The British Journal for the Philosophy of Science 30 (2): 165–170 [165–6].
doi:10.1093/bjps/30.2.165.
45.^ Boyer, C. A History of Mathematics. Wiley, p. 54.
46.^ Otto E. Neugebauer (1945). "The History of Ancient Astronomy Problems and Methods",
Journal of Near Eastern Studies 4 (1), p. 1–38.
 "the Chaldaean Seleucus from Seleucia"

47.^ George Sarton (1955). "Chaldaean Astronomy of the Last Three Centuries B. C.", Journal
of the American Oriental Society 75 (3), pp. 166–173 [169]:
"the heliocentrical astronomy invented by Aristarchos of Samos and still defended a
century later by Seleucos the Babylonian"

48.^ William P. D. Wightman (1951, 1953), The Growth of Scientific Ideas, Yale University Press
p.38, where Wightman calls him Seleukos the Chaldean.
49.^ Lucio Russo, Flussi e riflussi, Feltrinelli, Milano, 2003, ISBN 88-07-10349-4.
50.^ Bartel Leendert van der Waerden (1987), "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [527]
51.^ Bartel Leendert van der Waerden (1987), "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [527–9]
52.^ Bartel Leendert van der Waerden (1987). "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [529–34]
53.^ Bartel Leendert van der Waerden (1987). "The Heliocentric System in Greek, Persian and
Hindu Astronomy", Annals of the New York Academy of Sciences 500 (1): 525–545 [534–7]
54.^ Nasr, Seyyed H. (1st edition in 1964, 2nd edition in 1993). An Introduction to Islamic
Cosmological Doctrines (2nd ed.). 1st edition by Harvard University Press, 2nd edition by State
University of New York Press. pp. 135–6. ISBN 0791415155.
55.^ Misner, Thorne and Wheeler (1973), p. 754.
 56.^ Ragep, F. Jamil (2001a). "Tusi and Copernicus: The Earth's Motion in Context". Science in
Context (Cambridge University Press) 14 (1–2): 145–63.
57.^ Ragep, F. Jamil (2001b). "Freeing Astronomy from Philosophy: An Aspect of Islamic
Influence on Science". Osiris, 2nd Series 16 (Science in Theistic Contexts: Cognitive Dimensions):
49–64 & 66–71.
58.^ a b c d Misner, Thorne, and Wheeler (1973), p.755.
59.^ Misner, Thorne, and Wheeler (1973), p. 756.
60.^ de Cheseaux JPL (1744). Traité de la Comète. Lausanne. pp. 223ff. . Reprinted as
Appendix II in Dickson FP (1969). The Bowl of Night: The Physical Universe and Scientific Thought .
Cambridge, MA: M.I.T. Press. ISBN 978-0262540032.
61.^ Olbers HWM (1826). "Unknown title". Bode's Jahrbuch 111. . Reprinted as Appendix I in
Dickson FP (1969). The Bowl of Night: The Physical Universe and Scientific Thought . Cambridge,
MA: M.I.T. Press. ISBN 978-0262540032.
62.^ Jeans, J. H. (1902) Philosophical Transactions Royal Society of London, Series A , 199, 1.
63.^ Rindler, p. 196; Misner, Thorne, and Wheeler (1973), p. 757.
64.^ Misner, Thorne and Wheeler, p.756.
65.^ a b Einstein, A (1917). "Kosmologische Betrachtungen zur allgemeinen Relativitätstheorie".
Preussische Akademie der Wissenschaften, Sitzungsberichte 1917 (part 1): 142–152.
66.^ Rindler (1977), pp. 226–229.
67.^ Landau and Lifshitz (1975), pp. 358–359.
68.^ Einstein, A (1931). "Zum kosmologischen Problem der allgemeinen Relativitätstheorie".
Sitzungsberichte der Preussischen Akademie der Wissenschaften, Physikalisch-mathematische
Klasse 1931: 235–237.
Einstein A., de Sitter W. (1932). "On the relation between the expansion and the mean density of the
 universe". Proceedings of the National Academy of Sciences 18 (3): 213–214.
doi:10.1073/pnas.18.3.213. PMID 16587663.
69.^ Hubble Telescope news release
70.^ BBC News story: Evidence that dark energy is the cosmological constant
71.^ Zel'dovich YB (1967). "Cosmological constant and elementary particles". Zh. Eksp. & Teor.
Fiz. Pis'ma 6: 883–884. English translation in Sov. Phys. — JTEP Lett., 6, pp. 316–317 (1967).
72.^ Friedmann A. (1922). "Über die Krümmung des Raumes". Zeitschrift für Physik 10: 377–
386. doi:10.1007/BF01332580.
73.^ Munitz MK (1959). "One Universe or Many?". Journal of the History of Ideas 12 (2): 231–
255. doi:10.2307/2707516. http://links.jstor.org/sici?sici=0022-
5037(195104)12%3A2%3C231%3AOUOM%3E2.0.CO%3B2-F.
74.^ Misner, Thorne and Wheeler (1973), p.753.
75.^ Linde A. (1986). "Eternal chaotic inflation". Mod. Phys. Lett. A1: 81.
Linde A. (1986). "Eternally existing self-reproducing chaotic inflationary universe". Phys. Lett. B175:
395–400.
76.^ Shape of the Universe, WMAP website at NASA.
77.^ http://en.wikipedia.org/wiki/Homology_sphere#Cosmology

[edit] Further reading

• Landau, Lev, Lifshitz, E.M. (1975). The Classical Theory of Fields (Course of Theoretical
Physics, Vol. 2) (revised 4th English ed.). New York: Pergamon Press. pp. 358–397.
ISBN 9780080181769.
• Edward Robert Harrison (2000) Cosmology 2nd ed. Cambridge University Press. Gentle.
 • Misner, C.W., Thorne, Kip, Wheeler, J.A. (1973). Gravitation. San Francisco: W. H. Freeman.
pp. 703–816. ISBN 978-0-7167-0344-0. The classic text for a generation.
• Rindler, W. (1977). Essential Relativity: Special, General, and Cosmological . New York:
Springer Verlag. pp. 193–244. ISBN 0-387-10090-3.
• Weinberg, S. (1993). The First Three Minutes: A Modern View of the Origin of the Universe
(2nd updated ed.). New York: Basic Books. ISBN 978-0465024377. OCLC 28746057. For lay
readers.
• -------- (2008) Cosmology. Oxford University Press. Challenging.
• Oscar Monchito (1987) Universe. What a concept. Colton, 23rd edition. For advanced
readers.
• Nussbaumer, Harry; Bieri, Lydia; Sandage, Allan (2009). Discovering the Expanding
Universe. Cambridge University Press. ISBN 978-0-521-51484-2. http://books.google.com/books?
id=RaNOJkQ4l14C.

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 Universum - C. Flammarion, gravure sur bois, Paris 1888, coloris : Heikenwaelder Hugo, Wien 1998.
L'Univers est l'ensemble de tout ce qui existe et les lois qui le régissent.
La cosmologie cherche à appréhender l'Univers d'un point de vue scientifique, comme l'ensemble de
la matière distribuée dans le temps et dans l'espace. Pour sa part, la cosmogonie vise à établir une théorie
de la création de l'Univers sur des bases philosophiques ou religieuses.
Selon le modèle standard, on ne connaît au plus que 5 % de la matière de l'Univers[1] ; le reste se
composerait de 25 % de matière noire et de 70 % d'énergie noire. Selon le modèle alternatif avec antimatière
soutenu notamment par Gabriel Chardin et Stephen Hawking, la quantité de matière présente dans l'univers
est 15 fois plus abondante que dans le modèle conventionnel[2].
Sommaire
[masquer]
• 1 Découverte de l'Univers dans l'Histoire
• 2 Naissance de l'Univers
• 2.1 L'expansion de l'Univers, son âge et le Big Bang
• 2.2 Taille de l'Univers, Univers observable
• 2.3 Forme de l'Univers
• 2.4 Modèle dimensionnel de l'Univers
• 3 Avenir de l'Univers
• 4 Notes et références
• 5 Voir aussi
• 5.1 Articles connexes
• 5.2 Liens externes

Découverte de l'Univers dans l'Histoire [modifier]

Articles détaillés : Monde (univers) et Révolution copernicienne.
Les Sciences grecques tentèrent de comprendre le monde et de l'expliquer :
 • les philosophes Parménide, Platon, et Aristote avaient intégré l'idée d'une Terre sphérique,
mais ils la voyaient au centre de l'univers physique, alors que l'école de Milet se représentait la Terre
plate ;
• Ératosthène tenta de réaliser des calculs précis, notamment la mesure de la circonférence
d'un méridien terrestre ;
• Aristarque de Samos est le premier à envisager un modèle de système planétaire
héliocentré. Cette découverte ne fut alors pas suivie[3], pour des raisons philosophiques surtout
parce qu'une telle cosmologie est en désaccord avec la conception géocentrée du monde qui était
retenue par de grands philosophes comme Parménide, Platon, et Aristote. Il calcule aussi la distance
Terre-Lune pour laquelle il trouve une valeur discutée, mais qui se situe en tout état de cause dans
un ordre de grandeur acceptable[4], ainsi qu'une distance Terre-Soleil.[5] ;
• Hipparque poursuit ce travail : il recalcule, selon des méthodes nouvelles, la distance Terre-
Soleil ainsi que distance Terre-Lune (pour la quelle il retient la valeur de 67 1/3 rayons terrestres,
contre 60,2 en réalité[6]), recense 1 500 étoiles[réf. nécessaire], retrouve approximativement la
période de précession des équinoxes, qui était déjà connue des Babyloniens.[réf. nécessaire]
• Ptolémée poursuit le travail d'Hipparque. Son Almageste sera la référence astronomique
essentielle pendant treize siècles.
 L'Univers selon le système de Ptolémée, vu par Andreas Cellarius en 1660/61.
Ces connaissances du monde grec perdureront et influenceront les sciences arabes après
l'effondrement de l'Empire romain d'Occident. Elles resteront présentes en Orient (particulièrement, avec des
hauts et des bas, à Byzance[7]), même si Cosmas d'Alexandrie tente, sans succès, de restaurer le modèle
d'un monde plat.
La Renaissance porte à son apogée cette représentation du monde, grâce aux explorations et aux
grandes découvertes qui eurent lieu du XIIIe au XVIe siècles, à partir de systèmes géographiques et
cosmologiques très élaborés (projection de Mercator).
La révolution copernicienne bouleverse cette cosmologie en trois étapes :
 1. Copernic redécouvre l'héliocentrisme. Toutefois, cette redécouverte n'est que partiellement
révolutionnaire : en effet, Copernic reste attaché aux sphères transparentes du modèle d'Aristote
(pourtant délaissé par Ptolémée) censées soutenir les planètes et leur imprimer leur mouvement ; il
présente son système comme un simple artifice destiné à simplifier les calculs, ce qui lui évite des
ennuis avec le clergé.
2. Le dominicain Giordano Bruno défend la réalité du modèle héliocentrique et l'étend à toutes
les étoiles, ouvrant la dimension de l'univers physique à l'infini. Pour cette raison, entre autres, il sera
brulé au bûcher en tant qu'hérétique.
3. Kepler, Galilée et Newton posent les bases fondamentales de la mécanique à partir du
mouvement des planètes, grâce à leurs études respectivement du mouvement elliptique des
planètes autour du Soleil, l'affinement des observations astronomiques avec la définition du
mouvement uniformément accéléré, et la formalisation mathématique de la force de gravité.
L'Univers, toutefois, reste confiné dans le système solaire.
Des modèles physiques tels que la sphère armillaire ou l'astrolabe ont été élaborés. Ils permettent
d'enseigner et de calculer la position des astres dans le ciel visible. Aujourd'hui encore, la carte du ciel
mobile aide les astronomes amateurs à se repérer dans le ciel, c'est une ré-incarnation de l'astrolabe.

Naissance de l'Univers [modifier]

L'expansion de l'Univers, son âge et le Big Bang [modifier]

Articles détaillés : Frise chronologique du Big Bang, Expansion de l'Univers et Big Bang.
 Les observations du décalage vers le rouge des rayonnements électromagnétiques en provenance
d'autres galaxies suggèrent que celles-ci s'éloignent de notre galaxie, à une vitesse radiale d'éloignement
supposée proportionnelle à ce décalage.
En étudiant les galaxies proches, Edwin Hubble s'est aperçu que la vitesse d'éloignement d'une
galaxie était proportionnelle à sa distance par rapport à l'observateur (loi de Hubble); une telle loi correspond
à un Univers visible en expansion.
Bien que la constante de Hubble ait été révisée par le passé dans d'importantes proportions (dans
un rapport de 10 à 1), la loi de Hubble a été extrapolée aux galaxies éloignées, pour lesquelles la distance
ne peut être calculée au moyen de la parallaxe ; cette loi est ainsi utilisée pour déterminer la distance des
galaxies les plus lointaines.
En extrapolant l'expansion de l'Univers dans le passé, on arrive à une époque où celui-ci a dû être
beaucoup plus chaud et beaucoup plus dense qu'aujourd'hui. C'est le modèle du Big Bang qui est un
ingrédient essentiel du modèle standard de la cosmologie actuelle et possède aujourd'hui un grand nombre
de confirmations expérimentales. La description du début de l'histoire de l'Univers par ce modèle ne
commence cependant qu'après qu'il fût sorti d'une période appelée ère de Planck durant laquelle l'échelle
d'énergie de l'Univers était si grande que le modèle standard n'est pas en mesure de décrire les
phénomènes quantiques qui s'y sont déroulés. Durant cette époque, seule une théorie de la gravitation
quantique pourrait expliquer le comportement microscopique de la matière sous l'influence importante de la
gravité. Mais les physiciens ne disposent pas encore (en 2010) d'une telle théorie. Pour des raisons de
cohérence avec les observations, après l'ère de Planck le modèle du Big Bang privilégie aujourd'hui
l'existence d'une phase d'inflation cosmique très brève mais durant laquelle l'Univers aurait grandi de façon
extrêmement rapide. C'est suite à cette phase que l'essentiel des particules de l'Univers auraient été créées
avec une haute température, enclenchant un grand nombre de processus importants[8] qui ont finalement
abouti à l'émission d'une grande quantité de lumière, appelé fond diffus cosmologique, qui peut être
aujourd'hui observé avec une grande précision par toute une série d'instruments (ballons-sondes, sondes
spatiales).
C'est l'observation de ce rayonnement fossile micro-onde, remarquablement uniforme dans toutes
les directions qui constitue aujourd'hui l'élément capital qui assoit le modèle du Big Bang comme description
correcte de l'Univers dans son passé lointain. Beaucoup d'éléments du modèle devraient encore être
affinés[9], mais il y a aujourd'hui consensus de la communauté scientifique autour du modèle du Big Bang.
Dans le cadre du modèle ΛCDM, qui est le plus simple incorporant tous les éléments que l'on vient
d'évoquer, les contraintes issues des observations de la sonde WMAP[10] sur les paramètres
cosmologiques indiquent une valeur la plus probable pour l'âge de l'Univers à environ 13,7 milliards
d'années[11] avec une incertitude de 0,2 milliard d'années, ce qui est en accord avec les données
indépendantes issues de l'observation des amas globulaires[12] ainsi que celle des naines blanches[13].

Taille de l'Univers, Univers observable [modifier]

Article détaillé : Univers observable.
À ce jour, rien ne nous permet de confirmer que l'Univers est soit fini, soit infini. Certains théoriciens
penchent pour un Univers infini, d'autres pour un Univers fini mais non borné.
Les articles populaires et professionnels de recherche en cosmologie emploient souvent le terme
« Univers » dans le sens d'« Univers observable ». Nous vivons au centre de l'Univers observable, en
contradiction apparente avec le principe de Copernic qui dit que l'Univers est plus ou moins uniforme et ne
possède aucun centre en particulier. C'est simplement parce que la lumière ne se déplace pas à une vitesse
infinie et que les observations que nous faisons proviennent donc du passé. En effet, en regardant de plus
en plus loin, nous voyons des choses qui se sont passées à une époque de plus en plus proche du Big-
Bang. Et puisque la lumière se déplace à la même vitesse dans toutes les directions, tous les observateurs
vivent au centre de leur Univers observable (sur Terre, nous avons pratiquement tous le même). On appelle
« horizon cosmologique » la première lumière émise par le Big-Bang il y a 13,7 milliards d'années. Il nous est
impossible de voir plus loin.
On estime que le diamètre de cet Univers observable est de 100 milliards d'années lumière[14].
Celui-ci contient environ 7×1022 étoiles, répandues dans environ 1011 galaxies, elles-mêmes organisées en
amas et superamas de galaxies[14]. Mais le nombre de galaxies pourrait être encore plus grand, selon le
champ profond observé avec le télescope spatial Hubble.
Il est cependant probable que l'Univers que nous voyons n'est qu'une infime partie d'un Univers réel
beaucoup plus grand. Selon les derniers modèles cosmologiques, la taille minimale de l'Univers réel serait

de [Quoi ?] soit un 1 suivi de dix milliards de zéros[14]. (A priori, l'unité n'a pas d'importance:
métre, millimetre, kilometre ou UA. Ca représente seulement quelques zéro en plus ou en moins...)
On pourrait raisonner que, l'Univers contenant par définition tout ce qui existe, y compris l'espace-
temps (et c'est une précision essentielle), il ne peut avoir de « bord » tel que nous concevons intuitivement
cette notion. En effet, l'existence d'un bord impliquerait qu'au-delà de ce bord, on ne serait plus dans
l'Univers, ce qui serait absurde. Mais si l'Univers n'a pas de bord au sens intuitif de ce terme, alors son
expansion n'est pas intuitive non plus : si elle l'était, dans quoi l'Univers serait-il en expansion ?
On voit ici les apparents paradoxes entraînés par l'utilisation de notions dites « intuitives »... qui ne
sont que le reflet d'une perception locale de la réalité. Donner une définition précise à ces notions intuitives
permet de faire disparaître ces paradoxes. On voit que ce problème échappe à nos raisonnements
simplistes, qui se fondent sur des hypothèses fausses telles que « l'Univers est galiléen », ou « un espace
courbe est nécessairement inclus dans un espace de dimension supérieure ». En définitive, une définition
précise de la notion d'Univers permet de tenter de résoudre ces apparents paradoxes. Les paradoxes
restants permettent de démontrer... que la définition donnée à l'Univers n'a pas sens. On sait en fait
aujourd'hui qu'une définition, même formelle, d'un ensemble n'implique pas son existence.
En bref, le mot « Univers » reste à définir.

Forme de l'Univers [modifier]

Articles détaillés : Topologie de l'Univers et Courbure spatiale.
Une importante question de cosmologie qui reste sans réponse est la topologie de l'Univers.
1. Est-ce que l'Univers est « plat » ? C'est-à-dire : est-ce que le théorème de Pythagore pour
les triangles droits est valide à de plus grandes échelles ? Actuellement, la plupart des cosmologues
croient que l'Univers observable est (presque) plat, juste comme la Terre est (presque) plate.
2. Est-ce que l'Univers est simplement connexe ? Selon le modèle standard du Big Bang,
l'Univers n'a aucune frontière spatiale, mais peut néanmoins être de taille finie.
Ceci peut être compris par une analogie bidimensionnelle : la surface de la Terre n'a aucun bord,
mais possède une aire bien déterminée.
Vous pouvez également penser à un cylindre et imaginer de coller les 2 extrémités du cylindre ensemble,
mais sans plier le cylindre.
C'est aussi un espace bidimensionnel avec une surface finie, mais au contraire de la surface de la Terre, il
est plat, et peut ainsi servir de meilleur modèle.
 Par conséquent, à proprement parler, nous devrions appeler les étoiles et les galaxies mentionnées
ci-dessus « images » d'étoiles et de galaxies, puisqu'il est possible que l'Univers soit fini et si petit que nous
pouvons voir une ou plusieurs fois autour de lui, et le vrai nombre d'étoiles et de galaxies physiquement
distinctes pourrait être plus petit. Des hypothèses d'Univers multiconnexe ont été proposées et sont en cours
d'étude.

Modèle dimensionnel de l'Univers [modifier]

L’Univers a-t-il 3, 6, 10 dimensions ou plus ?
La théorie des cordes prédit qu’espace et matière sont consubstantiels. Il n’y a pas de « contenant »
(l’espace) mais un fond d’espace-temps qui interagit avec la matière. Dans certains cas particuliers, la notion
de « nombre de dimensions de l’espace » dépend de l’intensité avec laquelle les cordes réagissent entre
elles. Si cette interaction est faible, elles semblent se propager dans un espace à neuf dimensions -
auxquelles il faut rajouter celle du temps. Si cette interaction croît, cela développe une dimension de plus (ou
plus en fonction de l’intensité de l’interaction) à laquelle il faut toujours rajouter celle du temps. Supposons
maintenant qu’on enferme l’Univers dans un espace « fini » (une boîte pour être concret) et que cet espace
rapetisse jusqu’à 10-32 centimètre de côté, la théorie des cordes le prédit équivalent à un Univers très grand.
La conception d’espace est fondamentalement bouleversée. La mise en route du grand collisionneur de
hadrons de Genève, Large Hadron Collider (plus communément appelé LHC), viendra peut être confirmer
cette théorie. Elle ne pourra en revanche pas l'infirmer, car aucun ordre de grandeur n'a été prédit par la
théorie des cordes. Ainsi, si le phénomène n'est pas détecté, cela pourrait signifier que trop peu d'énergie a
été générée pour rendre le phénomène observable, sans impliquer pour autant que la théorie soit
nécessairement erronée.
Avenir de l'Univers [modifier]
Les objets galactiques auront une fin : le Soleil, par exemple, s'éteindra dans 5 (à 7) milliards
d'années, lorsqu'il aura consumé tout son combustible. À terme, les autres étoiles suivront elles aussi dans
des cataclysmes cosmologiques (explosions, effondrements). Déjà les naissances d'étoiles se
ralentissent[15] faute de matière, qui se raréfie au fil du temps. Dans 20 milliards d'années environ, aucun
astre ne s'allumera plus. L'Univers sera peuplé d'étoiles éteintes (étoiles à neutrons, naines blanches, trous
noirs) et des naines rouges résiduelles. À bien plus longues échéances, les galaxies se désagrègeront dans
des collisions géantes par leurs interactions gravitationnelles internes et externes[16].
En ce qui concerne le contenant (l'espace), certains[Qui ?] pensent que le processus d'expansion
sera gravitationnellement ralenti et s'inversera selon le scénario du Big Crunch[17]. Pour d'autres[Qui ?],
l'expansion, qui semble à présent accélérée par la présence d'une énergie répulsive de nature inconnue
(l'énergie sombre), continuera à jamais. Peu à peu, les astres éteints s'agglutineront en trous noirs.
L'Univers, sans aucune structure, ne sera plus qu'un bain de photons de plus en plus froids[18]. Toute
activité dans l'Univers s'éteindra ainsi à jamais. Si au contraire la quantité d'énergie sombre croît, l'Univers
continuera son expansion à une vitesse toujours plus grande pour exploser à toutes les échelles : toute la
matière qui le compose (y compris les atomes) se déchirera par dilatation de l'espace. C'est le Big Rip
(littéralement : « grand déchirement »). Certains modèles prévoient une telle fin dans 22 milliards d'années.
Chacun de ces scénarios dépend donc de la quantité d'énergie sombre que contiendra l'Univers à un
moment donné. Actuellement, l'état des connaissances suggère non seulement qu'il y a insuffisamment de
masse et d'énergie pour provoquer ce Big Rip, mais que l'expansion de l'Univers semble s'accélérer et
continuera donc pour toujours[réf. nécessaire].
Notes et références [modifier]
1. ↑ NASA WMAP What is the universe made of ? [archive]
2. ↑ Antimatière : est-elle la clé de l'Univers ?, Science et vie, octobre 2009, n°1105, p.60.
3. ↑ À notre connaissance, un seul autre savant de l'antiquité fut de cet avis, Séleucos de
Séleucie.
4. ↑ Otto Neugebauer, A history of ancient mathematical astronomy, Berlin ; New York :
Springer-Verlag, 1975, p. 634 ss. Aristarque ne donne pas le résultat de ses calculs, mais de ses
données (diamètre apparent angulaire de la Lune : 2° ; diamètre de la Lune : 1/3 de diamètre
lunaire), on peut déduire une distance Terre-Lune de 40 rayons terrestres environ, contre 60,2 en
réalité. Mais Neugebauer estime que c'est un angle de 1/2° et non de 2° qu'Aristarque tenait pour
correct, ce qui aboutirait à 80 rayons terrestres pour la distance Terre-Lune. Voir Aristarque.
5. ↑ Neugebauer, Loc. cit..
6. ↑ Cf. Des grandeurs et des distances du Soleil et de la Lune.
7. ↑ Cf. sciences grecques.
8. ↑ Comme la nucléosynthèse primordiale par exemple ou encore la baryogénèse.
9. ↑ Il n'est pas encore possible de bien discriminer entre plusieurs modèles d'inflation pour
donner un exemple.
10.↑ Lancée par la NASA.
11.↑ (en) D.N. Spergel et al., Wilkinson microwave anisotropy probe (wmap) three year
results : implications for cosmology. [archive] soumis à Astrophys. J., prépublication disponible sur la
base de données arXiv.
12.↑ (en) Chaboyer, B. & Krauss, Theoretical Uncertainties in the Subgiant--Mass Age
Relation and the Absolute Age of Omega Cen [archive]L. M. 2002, ApJ, 567, L45.
 13.↑ (en)B. Hanser et al HST Observations of the White Dwarf Cooling Sequence of
M4 [archive] The Astrophysical Journal Supplement Series, Volume 155, Issue 2, p. 551-576.,
prépublication disponible sur l'arXiv.
14.↑ a, b et c Science-et-Vie Hors-Série n°242, mars 2008. "L'Univers en chiffres".
15.↑ Actuellement, l'observation de notre galaxie dénombre la naissance d'une ou deux étoiles
par an.
16.↑ Jean-Pierre Luminet, Astrophysicien, CNRS, Observatoire de Paris-Meudon, in Sciences
& Avenir N°729, Novembre 2007.
17.↑ Littéralement : « grand écrasement ».
18.↑ D'après une théorie de Stephen Hawking (dans son livre Une brève histoire du temps), si
l'Univers continue indéfiniment à s'étendre, les particules issues d'explosions successives ne seront
plus assez proches les unes des autres pour recréer des étoiles après leur explosion.

Voir aussi [modifier]

Sur les autres projets Wikimédia :
• Univers sur le Wiktionnaire (dictionnaire universel)
• Univers sur Wikibooks (livres pédagogiques)
• Univers sur Wikisource (bibliothèque universelle)
• Modèle cosmologique
• Cosmologie non standard
Articles connexes [modifier]
• Voie lactée
• Monde (univers)
• Radiotélescope
• Observatoire astronomique
• Destin de l'Univers
• Âge de l'Univers
• Masse de l'Univers
• Théorie d'Everett
• Table des constantes astrophysiques
• Densité critique
• Énergie sombre
• Matière noire

Liens externes [modifier]

• (fr) Univers : page de l'Institut national des sciences de l'Univers

• Portail de l’astronomie

• Portail de la cosmologie
 W000

Earth
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about the planet. For other uses, see Earth (disambiguation).

Earth
 "Blue Marble" photograph of Earth,
taken from Apollo 17
Designations
Pronunciation i /ˈɜrθ/

Adjective earthly, tellurian, telluric, terran, terrestrial.

 Orbital characteristics
Epoch J2000.0[note 1]

152,098,232 km
Aphelion
1.01671388 AU[note 2]

147,098,290 km
Perihelion
0.98329134 AU[note 2]

149,598,261 km
Semi-major axis
1.00000261 AU[1]

Eccentricity 0.01671123[1]

365.256363004 days[2]
Orbital period
1.000017421 yr

29.78 km/s[3]
Average orbital speed
107,200 km/h

7.155° to Sun's equator

Inclination
1.57869°[4] to invariable plane

Longitude of ascending node 348.73936°[3][note 3]

Argument of perihelion 114.20783°[3][note 4]

 Satellites 1 (the Moon)

Physical characteristics
Mean radius 6,371.0 km[5]

Equatorial radius 6,378.1 km[6]

Polar radius 6,356.8 km[7]

Flattening 0.0033528[6]

40,075.16 km (equatorial)[8]
Circumference
40,008.00 km (meridional)[8]

510,072,000 km2[9][10][note 5]
Surface area 148,940,000 km2 land (29.2 %)
361,132,000 km2 water (70.8 %)

Volume 1.08321 × 1012 km3[3]

Mass 5.9736 × 1024 kg[3]

 Mean density 5.515 g/cm3[3]

Equatorial surface gravity 9.780327 m/s2[11]

0.99732 g

Escape velocity 11.186 km/s[3]

Sidereal rotation 0.99726968 d[12]

period 23h 56m 4.100s

Equatorial rotation velocity 1,674.4 km/h (465.1 m/s)[13]

Axial tilt 23°26'21".4119[2]

Albedo 0.367[3]

Surface temp.
Kelvin
Celsius
min mean max
184 K[14 287.2 K[15 331 K[16
] ] ]
-89.2 °C 14 °C 57.8 °C
 Atmosphere
Surface pressure 101.325 kPa (MSL)

78.08% nitrogen (N2)[3]

20.95% oxygen (O2)
Composition 0.93% argon
0.038% carbon dioxide
About 1% water vapor (varies with climate)
Earth (or the Earth) is the third planet from the Sun, and the densest and fifth-largest of the eight
planets in the Solar System. It is also the largest of the Solar System's four terrestrial planets. It is sometimes
referred to as the World, the Blue Planet,[note 6] or by its Latin name, Terra.[note 7]
Home to millions of species[17] including humans, Earth is currently the only place in the universe
where life is known to exist. The planet formed 4.54 billion years ago,[18] and life appeared on its surface
within a billion years. Earth's biosphere has significantly altered the atmosphere and other abiotic conditions
on the planet, enabling the proliferation of aerobic organisms as well as the formation of the ozone layer
which, together with Earth's magnetic field, blocks harmful solar radiation, permitting life on land.[19] The
physical properties of the Earth, as well as its geological history and orbit, have allowed life to persist during
this period. The planet is expected to continue supporting life for at least another 500 million years.[20][21]
Earth's outer surface is divided into several rigid segments, or tectonic plates, that migrate across the
surface over periods of many millions of years. About 71% of the surface is covered with salt water oceans,
the remainder consisting of continents and islands which together have many lakes and other sources of
water contributing to the hydrosphere. Liquid water, necessary for all known life, is not known to exist on any
other planet's surface.[note 8][note 9] Earth's poles are mostly covered with solid ice (Antarctic ice sheet) or
sea ice (Arctic ice cap). The planet's interior remains active, with a thick layer of relatively solid mantle, a
liquid outer core that generates a magnetic field, and a solid iron inner core.
Earth interacts with other objects in space, especially the Sun and the Moon. At present, Earth orbits
the Sun once for every roughly 366.26 times it rotates about its axis, which is equal to 365.26 solar days, or
one sidereal year.[note 10] The Earth's axis of rotation is tilted 23.4° away from the perpendicular to its orbital
plane,[22] producing seasonal variations on the planet's surface with a period of one tropical year (365.24
solar days). Earth's only known natural satellite, the Moon, which began orbiting it about 4.53 billion years
ago, provides ocean tides, stabilizes the axial tilt and gradually slows the planet's rotation. Between
approximately 3.8 billion and 4.1 billion years ago, numerous asteroid impacts during the Late Heavy
Bombardment caused significant changes to the greater surface environment.
Both the mineral resources of the planet, as well as the products of the biosphere, contribute
resources that are used to support a global human population. These inhabitants are grouped into about 200
independent sovereign states, which interact through diplomacy, travel, trade, and military action. Human
cultures have developed many views of the planet, including personification as a deity, a belief in a flat Earth
or in Earth as the center of the universe, and a modern perspective of the world as an integrated environment
that requires stewardship.
Contents
[hide]
• 1 Chronology
• 1.1 Evolution of life
• 1.2 Future
• 2 Composition and structure
• 2.1 Shape
• 2.2 Chemical composition
• 2.3 Internal structure
• 2.4 Heat
• 2.5 Tectonic plates
• 2.6 Surface
• 2.7 Hydrosphere
• 2.8 Atmosphere
• 2.8.1 Weather and climate
• 2.8.2 Upper atmosphere
• 2.9 Magnetic field
• 3 Orbit and rotation
• 3.1 Rotation
• 3.2 Orbit
• 3.3 Axial tilt and seasons
[edit] Chronology
Main article: History of the Earth
See also: Geological history of Earth
Scientists have been able to reconstruct detailed information about the planet's past. The earliest
dated Solar System material is dated to 4.5672 ± 0.0006 billion years ago,[23] and by 4.54 billion years ago
(within an uncertainty of 1%)[18] the Earth and the other planets in the Solar System had formed out of the
solar nebula—a disk-shaped mass of dust and gas left over from the formation of the Sun. This assembly of
the Earth through accretion was thus largely completed within 10–20 million years.[24] Initially molten, the
outer layer of the planet Earth cooled to form a solid crust when water began accumulating in the
atmosphere. The Moon formed shortly thereafter, 4.53 billion years ago.[25]
The current consensus model[26] for the formation of the Moon is the giant impact hypothesis, in
which the Moon formed as a result of a Mars-sized object (sometimes called Theia) with about 10% of the
Earth's mass[27] impacting the Earth in a glancing blow.[28] In this model, some of this object's mass would
have merged with the Earth and a portion would have been ejected into space, but enough material would
have been sent into orbit to form the Moon.
Outgassing and volcanic activity produced the primordial atmosphere. Condensing water vapor,
augmented by ice and liquid water delivered by asteroids and the larger proto-planets, comets, and trans-
Neptunian objects produced the oceans.[29] The newly formed Sun was only 70% of its present luminosity,
yet evidence shows that the early oceans remained liquid—a contradiction dubbed the faint young Sun
paradox. A combination of greenhouse gases and higher levels of solar activity served to raise the Earth's
surface temperature, preventing the oceans from freezing over.[30] By 3.5 billion years ago, the Earth's
magnetic field was established, which helped prevent the atmosphere from being stripped away by the solar
wind.[31]
Two major models have been proposed for the rate of continental growth:[32] steady growth to the
present-day[33] and rapid growth early in Earth history.[34] Current research shows that the second option is
most likely, with rapid initial growth of continental crust[35] followed by a long-term steady continental area.
[36][37][38] On time scales lasting hundreds of millions of years, the surface continually reshaped as
continents formed and broke up. The continents migrated across the surface, occasionally combining to form
a supercontinent. Roughly 750 million years ago (Ma), one of the earliest known supercontinents, Rodinia,
began to break apart. The continents later recombined to form Pannotia, 600–540 Ma, then finally Pangaea,
which broke apart 180 Ma.[39]

[edit] Evolution of life

Main article: Evolutionary history of life
At present, Earth provides the only example of an environment that has given rise to the evolution of
life.[40] Highly energetic chemistry is believed to have produced a self-replicating molecule around
4 billion years ago and half a billion years later the last common ancestor of all life existed.[41] The
development of photosynthesis allowed the Sun's energy to be harvested directly by life forms; the resultant
oxygen accumulated in the atmosphere and formed a layer of ozone (a form of molecular oxygen [O3]) in the
upper atmosphere. The incorporation of smaller cells within larger ones resulted in the development of
complex cells called eukaryotes.[42] True multicellular organisms formed as cells within colonies became
increasingly specialized. Aided by the absorption of harmful ultraviolet radiation by the ozone layer, life
colonized the surface of Earth.[43]
 Since the 1960s, it has been hypothesized that severe glacial action between 750 and 580 Ma,
during the Neoproterozoic, covered much of the planet in a sheet of ice. This hypothesis has been termed
"Snowball Earth", and is of particular interest because it preceded the Cambrian explosion, when
multicellular life forms began to proliferate.[44]
Following the Cambrian explosion, about 535 Ma, there have been five major mass extinctions.[45]
The most recent such event was 65 Ma, when an asteroid impact triggered the extinction of the (non-avian)
dinosaurs and other large reptiles, but spared some small animals such as mammals, which then resembled
shrews. Over the past 65 million years, mammalian life has diversified, and several million years ago, an
African ape-like animal such as Orrorin tugenensis gained the ability to stand upright.[46] This enabled tool
use and encouraged communication that provided the nutrition and stimulation needed for a larger brain. The
development of agriculture, and then civilization, allowed humans to influence the Earth in a short time span
as no other life form had,[47] affecting both the nature and quantity of other life forms.
The present pattern of ice ages began about 40 Ma and then intensified during the Pleistocene about
3 Ma. High-latitude regions have since undergone repeated cycles of glaciation and thaw, repeating every
40–100,000 years. The last continental glaciation ended 10,000 years ago.[48]

[edit] Future
Main article: Future of the Earth
See also: Risks to civilization, humans and planet Earth
 The life cycle of the Sun
The future of the planet is closely tied to that of the Sun. As a result of the steady accumulation of
helium at the Sun's core, the star's total luminosity will slowly increase. The luminosity of the Sun will grow by
10% over the next 1.1 Gyr (1.1 billion years) and by 40% over the next 3.5 Gyr.[49] Climate models indicate
that the rise in radiation reaching the Earth is likely to have dire consequences, including the loss of the
planet's oceans.[50]
The Earth's increasing surface temperature will accelerate the inorganic CO2 cycle, reducing its
concentration to levels lethally low for plants (10 ppm for C4 photosynthesis) in approximately 500 million[20]
to 900 million years. The lack of vegetation will result in the loss of oxygen in the atmosphere, so animal life
will become extinct within several million more years.[51] After another billion years all surface water will
have disappeared[21] and the mean global temperature will reach 70 °C[51] (158 °F). The Earth is expected
to be effectively habitable for about another 500 million years from that point,[20] although this may be
extended up to 2.3 billion years if the nitrogen is removed from the atmosphere.[52] Even if the Sun were
eternal and stable, the continued internal cooling of the Earth would result in a loss of much of its CO 2 due to
reduced volcanism,[53] and 35% of the water in the oceans would descend to the mantle due to reduced
steam venting from mid-ocean ridges.[54]
The Sun, as part of its evolution, will become a red giant in about 5 Gyr. Models predict that the Sun
will expand out to about 250 times its present radius, roughly 1 AU (150,000,000 km).[49][55] Earth's fate is
less clear. As a red giant, the Sun will lose roughly 30% of its mass, so, without tidal effects, the Earth will
move to an orbit 1.7 AU (250,000,000 km) from the Sun when the star reaches it maximum radius. The
planet was therefore initially expected to escape envelopment by the expanded Sun's sparse outer
atmosphere, though most, if not all, remaining life would have been destroyed by the Sun's increased
luminosity (peaking at about 5000 times its present level).[49] However, a more recent simulation indicates
that Earth's orbit will decay due to tidal effects and drag, causing it to enter the red giant Sun's atmosphere
and be vaporized.[55]

[edit] Composition and structure

Main article: Earth science
Further information: Earth physical characteristics tables
Earth is a terrestrial planet, meaning that it is a rocky body, rather than a gas giant like Jupiter. It is
the largest of the four solar terrestrial planets in size and mass. Of these four planets, Earth also has the
highest density, the highest surface gravity, the strongest magnetic field, and fastest rotation.[56] It also is the
only terrestrial planet with active plate tectonics.[57]

[edit] Shape
Main article: Figure of the Earth

Size comparison of inner planets (left to right): Mercury, Venus, Earth and Mars
The shape of the Earth is very close to that of an oblate spheroid, a sphere flattened along the axis
from pole to pole such that there is a bulge around the equator.[58] This bulge results from the rotation of the
Earth, and causes the diameter at the equator to be 43 km larger than the pole to pole diameter.[59] The
average diameter of the reference spheroid is about 12,742 km, which is approximately 40,000 km/π, as the
meter was originally defined as 1/10,000,000 of the distance from the equator to the North Pole through
Paris, France.[60]
 Local topography deviates from this idealized spheroid, though on a global scale, these deviations
are very small: Earth has a tolerance of about one part in about 584, or 0.17%, from the reference spheroid,
which is less than the 0.22% tolerance allowed in billiard balls.[61] The largest local deviations in the rocky
surface of the Earth are Mount Everest (8848 m above local sea level) and the Mariana Trench (10,911 m
below local sea level). Because of the equatorial bulge, the surface locations farthest from the center of the
Earth are the summits of Mount Chimborazo in Ecuador and Huascarán in Peru.[62][63][64]
Chemical composition of the crust[65]
Composition
Compound Formula
Continental Oceanic

silica SiO2 60.2% 48.6%

alumina Al2O3 15.2% 16.5%

lime CaO 5.5% 12.3%

magnesia MgO 3.1% 6.8%

iron(II) oxide FeO 3.8% 6.2%

sodium oxide Na2O 3.0% 2.6%

potassium oxide K2O 2.8% 0.4%

 iron(III) oxide Fe2O3 2.5% 2.3%

water H2O 1.4% 1.1%

carbon dioxide CO2 1.2% 1.4%

titanium dioxide TiO2 0.7% 1.4%

phosphorus pentoxide P2O5 0.2% 0.3%

Total 99.6% 99.9%

[edit] Chemical composition

See also: Abundance of elements on Earth
The mass of the Earth is approximately 5.98 × 1024 kg. It is composed mostly of iron (32.1%), oxygen
(30.1%), silicon (15.1%), magnesium (13.9%), sulfur (2.9%), nickel (1.8%), calcium (1.5%), and aluminium
(1.4%); with the remaining 1.2% consisting of trace amounts of other elements. Due to mass segregation, the
core region is believed to be primarily composed of iron (88.8%), with smaller amounts of nickel (5.8%),
sulfur (4.5%), and less than 1% trace elements.[66]
The geochemist F. W. Clarke calculated that a little more than 47% of the Earth's crust consists of
oxygen. The more common rock constituents of the Earth's crust are nearly all oxides; chlorine, sulfur and
fluorine are the only important exceptions to this and their total amount in any rock is usually much less than
1%. The principal oxides are silica, alumina, iron oxides, lime, magnesia, potash and soda. The silica
functions principally as an acid, forming silicates, and all the commonest minerals of igneous rocks are of this
nature. From a computation based on 1,672 analyses of all kinds of rocks, Clarke deduced that 99.22% were
composed of 11 oxides (see the table at right). All the other constituents occur only in very small quantities.
[67]

[edit] Internal structure

Main article: Structure of the Earth
The interior of the Earth, like that of the other terrestrial planets, is divided into layers by their
chemical or physical (rheological) properties. The outer layer of the Earth is a chemically distinct silicate solid
crust, which is underlain by a highly viscous solid mantle. The crust is separated from the mantle by the
Mohorovičić discontinuity, and the thickness of the crust varies: averaging 6 km under the oceans and 30–
50 km on the continents. The crust and the cold, rigid, top of the upper mantle are collectively known as the
lithosphere, and it is of the lithosphere that the tectonic plates are comprised. Beneath the lithosphere is the
asthenosphere, a relatively low-viscosity layer on which the lithosphere rides. Important changes in crystal
structure within the mantle occur at 410 and 660 kilometers below the surface, spanning a transition zone
that separates the upper and lower mantle. Beneath the mantle, an extremely low viscosity liquid outer core
lies above a solid inner core.[68] The inner core may rotate at a slightly higher angular velocity than the
remainder of the planet, advancing by 0.1–0.5° per year.[69]
Geologic layers of the Earth[70]
 Depth[ Density
Component
71]
km
Layer g/cm3

Lithosphere[not
0–60 —
e 11]

0–35 Crust[note 12] 2.2–2.9

35–60 Upper mantle 3.4–4.4

35–
Mantle 3.4–5.6
2890

100–
Asthenosphere —
700

Earth cutaway from core to exosphere. Not to 2890–

Outer core 9.9–12.2
scale. 5100

5100– 12.8–
Inner core
6378 13.1
[edit] Heat
Earth's internal heat comes from a combination of residual heat from planetary accretion (about 20%)
and heat produced through radioactive decay (80%).[72] The major heat-producing isotopes in the Earth are
potassium-40, uranium-238, uranium-235, and thorium-232.[73] At the center of the planet, the temperature
may be up to 7,000 K and the pressure could reach 360 GPa.[74] Because much of the heat is provided by
radioactive decay, scientists believe that early in Earth history, before isotopes with short half-lives had been
depleted, Earth's heat production would have been much higher. This extra heat production, twice present-
day at approximately 3 billion years ago,[72] would have increased temperature gradients within the Earth,
increasing the rates of mantle convection and plate tectonics, and allowing the production of igneous rocks
such as komatiites that are not formed today.[75]
Present-day major heat-producing isotopes[76]
Heat Half-life Mean mantle Heat
Isotope release concentration release
W/kg isotope years kg isotope/kg mantle W/kg mantle

238U 4.47 × 2.91 ×

9.46 × 10−5 30.8 × 10−9
109 10−12

235U 7.04 × 1.25 ×

5.69 × 10−4 0.22 × 10−9
108 10−13

232Th 1.40 × 3.27 ×

2.64 × 10−5 124 × 10−9
1010 10−12
 40K 1.25 × 1.08 ×
2.92 × 10−5 36.9 × 10−9
109 10−12

Total heat loss from the Earth is 4.2 × 1013 watts.[77] A portion of the core's thermal energy is
transported toward the crust by mantle plumes; a form of convection consisting of upwellings of higher-
temperature rock. These plumes can produce hotspots and flood basalts.[78] More of the heat in the Earth is
lost through plate tectonics, by mantle upwelling associated with mid-ocean ridges. The final major mode of
heat loss is through conduction through the lithosphere, the majority of which occurs in the oceans because
the crust there is much thinner than that of the continents.[77]

[edit] Tectonic plates

Earth's main plates[79]
 Area
Plate name
106 km2

African Plate[note 13] 78.0

Antarctic Plate 60.9

Indo-Australian Plate 47.2

Eurasian Plate 67.8

North American Plate 75.9

South American Plate 43.6

Pacific Plate 103.3

Main article: Plate tectonics
The mechanically rigid outer layer of the Earth, the lithosphere, is broken into pieces called tectonic
plates. These plates are rigid segments that move in relation to one another at one of three types of plate
boundaries: Convergent boundaries, at which two plates come together, Divergent boundaries, at which two
plates are pulled apart, and Transform boundaries, in which two plates slide past one another laterally.
Earthquakes, volcanic activity, mountain-building, and oceanic trench formation can occur along these plate
boundaries.[80] The tectonic plates ride on top of the asthenosphere, the solid but less-viscous part of the
upper mantle that can flow and move along with the plates,[81] and their motion is strongly coupled with
convection patterns inside the Earth's mantle.
As the tectonic plates migrate across the planet, the ocean floor is subducted under the leading
edges of the plates at convergent boundaries. At the same time, the upwelling of mantle material at divergent
boundaries creates mid-ocean ridges. The combination of these processes continually recycles the oceanic
crust back into the mantle. Because of this recycling, most of the ocean floor is less than 100 million years in
age. The oldest oceanic crust is located in the Western Pacific, and has an estimated age of about 200
million years.[82][83] By comparison, the oldest dated continental crust is 4030 million years old.[84]
Other notable plates include the Indian Plate, the Arabian Plate, the Caribbean Plate, the Nazca
Plate off the west coast of South America and the Scotia Plate in the southern Atlantic Ocean. The Australian
Plate fused with the Indian Plate between 50 and 55 million years ago. The fastest-moving plates are the
oceanic plates, with the Cocos Plate advancing at a rate of 75 mm/yr[85] and the Pacific Plate moving 52–
69 mm/yr. At the other extreme, the slowest-moving plate is the Eurasian Plate, progressing at a typical rate
of about 21 mm/yr.[86]

[edit] Surface
Main articles: Landform and Extreme points of Earth
The Earth's terrain varies greatly from place to place. About 70.8%[87] of the surface is covered by
water, with much of the continental shelf below sea level. The submerged surface has mountainous features,
including a globe-spanning mid-ocean ridge system, as well as undersea volcanoes,[59] oceanic trenches,
submarine canyons, oceanic plateaus and abyssal plains. The remaining 29.2% not covered by water
consists of mountains, deserts, plains, plateaus, and other geomorphologies.
 The planetary surface undergoes reshaping over geological time periods because of tectonics and
erosion. The surface features built up or deformed through plate tectonics are subject to steady weathering
from precipitation, thermal cycles, and chemical effects. Glaciation, coastal erosion, the build-up of coral
reefs, and large meteorite impacts[88] also act to reshape the landscape.

Present day Earth altimetry and bathymetry. Data from the National Geophysical Data Center's
TerrainBase Digital Terrain Model.
The continental crust consists of lower density material such as the igneous rocks granite and
andesite. Less common is basalt, a denser volcanic rock that is the primary constituent of the ocean floors.
[89] Sedimentary rock is formed from the accumulation of sediment that becomes compacted together.
Nearly 75% of the continental surfaces are covered by sedimentary rocks, although they form only about 5%
of the crust.[90] The third form of rock material found on Earth is metamorphic rock, which is created from the
transformation of pre-existing rock types through high pressures, high temperatures, or both. The most
abundant silicate minerals on the Earth's surface include quartz, the feldspars, amphibole, mica, pyroxene
and olivine.[91] Common carbonate minerals include calcite (found in limestone) and dolomite.[92]
The pedosphere is the outermost layer of the Earth that is composed of soil and subject to soil
formation processes. It exists at the interface of the lithosphere, atmosphere, hydrosphere and biosphere.
Currently the total arable land is 13.31% of the land surface, with only 4.71% supporting permanent crops.
[10] Close to 40% of the Earth's land surface is presently used for cropland and pasture, or an estimated
1.3 × 107 km2 of cropland and 3.4 × 107 km2 of pastureland.[93]
The elevation of the land surface of the Earth varies from the low point of −418 m at the Dead Sea, to
a 2005-estimated maximum altitude of 8,848 m at the top of Mount Everest. The mean height of land above
sea level is 840 m.[94]

[edit] Hydrosphere
Main article: Hydrosphere
 Elevation histogram of the surface of the Earth
The abundance of water on Earth's surface is a unique feature that distinguishes the "Blue Planet"
from others in the Solar System. The Earth's hydrosphere consists chiefly of the oceans, but technically
includes all water surfaces in the world, including inland seas, lakes, rivers, and underground waters down to
a depth of 2,000 m. The deepest underwater location is Challenger Deep of the Mariana Trench in the Pacific
Ocean with a depth of −10,911.4 m.[note 14][95]
 The mass of the oceans is approximately 1.35 × 1018 metric tons, or about 1/4400 of the total mass
of the Earth. The oceans cover an area of 361.8 × 106 km2 with a mean depth of 3,682 m, resulting in an
estimated volume of 1.332 × 109 km3.[96] If all the land on Earth were spread evenly, water would rise to an
altitude of more than 2.7 km.[note 15] About 97.5% of the water is saline, while the remaining 2.5% is fresh
water. Most fresh water, about 68.7%, is currently ice.[97]
The average salinity of the Earth's oceans is about 35 grams of salt per kilogram of sea water
(35 ‰).[98] Most of this salt was released from volcanic activity or extracted from cool, igneous rocks.[99]
The oceans are also a reservoir of dissolved atmospheric gases, which are essential for the survival of many
aquatic life forms.[100] Sea water has an important influence on the world's climate, with the oceans acting
as a large heat reservoir.[101] Shifts in the oceanic temperature distribution can cause significant weather
shifts, such as the El Niño-Southern Oscillation.[102]

[edit] Atmosphere
Main article: Atmosphere of Earth
The atmospheric pressure on the surface of the Earth averages 101.325 kPa, with a scale height of
about 8.5 km.[3] It is 78% nitrogen and 21% oxygen, with trace amounts of water vapor, carbon dioxide and
other gaseous molecules. The height of the troposphere varies with latitude, ranging between 8 km at the
poles to 17 km at the equator, with some variation resulting from weather and seasonal factors.[103]
Earth's biosphere has significantly altered its atmosphere. Oxygenic photosynthesis evolved 2.7
billion years ago, forming the primarily nitrogen-oxygen atmosphere of today. This change enabled the
proliferation of aerobic organisms as well as the formation of the ozone layer which blocks ultraviolet solar
radiation, permitting life on land. Other atmospheric functions important to life on Earth include transporting
water vapor, providing useful gases, causing small meteors to burn up before they strike the surface, and
moderating temperature.[104] This last phenomenon is known as the greenhouse effect: trace molecules
within the atmosphere serve to capture thermal energy emitted from the ground, thereby raising the average
temperature. Carbon dioxide, water vapor, methane and ozone are the primary greenhouse gases in the
Earth's atmosphere. Without this heat-retention effect, the average surface temperature would be −18 °C and
life would likely not exist.[87]

[edit] Weather and climate

Main articles: Weather and Climate

Satellite cloud cover image of Earth using NASA's Moderate-Resolution Imaging Spectroradiometer.
The Earth's atmosphere has no definite boundary, slowly becoming thinner and fading into outer
space. Three-quarters of the atmosphere's mass is contained within the first 11 km of the planet's surface.
This lowest layer is called the troposphere. Energy from the Sun heats this layer, and the surface below,
causing expansion of the air. This lower density air then rises, and is replaced by cooler, higher density air.
The result is atmospheric circulation that drives the weather and climate through redistribution of heat
energy.[105]
The primary atmospheric circulation bands consist of the trade winds in the equatorial region below
30° latitude and the westerlies in the mid-latitudes between 30° and 60°.[106] Ocean currents are also
important factors in determining climate, particularly the thermohaline circulation that distributes heat energy
from the equatorial oceans to the polar regions.[107]

Source regions of global air masses

Water vapor generated through surface evaporation is transported by circulatory patterns in the
atmosphere. When atmospheric conditions permit an uplift of warm, humid air, this water condenses and
settles to the surface as precipitation.[105] Most of the water is then transported to lower elevations by river
systems and usually returned to the oceans or deposited into lakes. This water cycle is a vital mechanism for
supporting life on land, and is a primary factor in the erosion of surface features over geological periods.
Precipitation patterns vary widely, ranging from several meters of water per year to less than a millimeter.
Atmospheric circulation, topological features and temperature differences determine the average
precipitation that falls in each region.[108]
The amount of solar energy reaching the Earth's decreases with increasing latitude. At higher
latitudes the sunlight reaches the surface at a lower angles and it must pass through thicker columns of the
atmosphere. As a result, the mean annual air temperature at sea level decreases by about 0.4° C per per
degree of latitude away from the equator.[109] The Earth can be sub-divided into specific latitudinal belts of
approximately homogeneous climate. Ranging from the equator to the polar regions, these are the tropical
(or equatorial), subtropical, temperate and polar climates.[110] Climate can also be classified based on the
temperature and precipitation, with the climate regions characterized by fairly uniform air masses. The
commonly used Köppen climate classification system (as modified by Wladimir Köppen's student Rudolph
Geiger) has five broad groups (humid tropics, arid, humid middle latitudes, continental and cold polar), which
are further divided into more specific subtypes.[106]
 [edit] Upper atmosphere

This view from orbit shows the full Moon partially obscured and deformed by the Earth's atmosphere.
NASA image.
See also: Outer space
Above the troposphere, the atmosphere is usually divided into the stratosphere, mesosphere, and
thermosphere.[104] Each layer has a different lapse rate, defining the rate of change in temperature with
height. Beyond these, the exosphere thins out into the magnetosphere, where the Earth's magnetic fields
interact with the solar wind.[111] An important part of the atmosphere for life on Earth is the ozone layer, a
component of the stratosphere that partially shields the surface from ultraviolet light. The Kármán line,
defined as 100 km above the Earth's surface, is a working definition for the boundary between atmosphere
and space.[112]
Thermal energy causes some of the molecules at the outer edge of the Earth's atmosphere have
their velocity increased to the point where they can escape from the planet's gravity. This results in a slow but
steady leakage of the atmosphere into space. Because unfixed hydrogen has a low molecular weight, it can
achieve escape velocity more readily and it leaks into outer space at a greater rate than other gasses.[113]
The leakage of hydrogen into space contributes to the pushing of the Earth from an initially reducing state to
its current oxidizing one. Photosynthesis provided a source of free oxygen, but the loss of reducing agents
such as hydrogen is believed to have been a necessary precondition for the widespread accumulation of
oxygen in the atmosphere.[114] Hence the ability of hydrogen to escape from the Earth's atmosphere may
have influenced the nature of life that developed on the planet.[115] In the current, oxygen-rich atmosphere
most hydrogen is converted into water before it has an opportunity to escape. Instead, most of the hydrogen
loss comes from the destruction of methane in the upper atmosphere.[116]
[edit] Magnetic field

The Earth's magnetic field, which approximates a dipole

Main article: Earth's magnetic field
The Earth's magnetic field is shaped roughly as a magnetic dipole, with the poles currently located
proximate to the planet's geographic poles. According to dynamo theory, the field is generated within the
molten outer core region where heat creates convection motions of conducting materials, generating electric
currents. These in turn produce the Earth's magnetic field. The convection movements in the core are
chaotic; the magnetic poles drift and periodically change alignment. This results in field reversals at irregular
intervals averaging a few times every million years. The most recent reversal occurred approximately
700,000 years ago.[117][118]
The field forms the magnetosphere, which deflects particles in the solar wind. The sunward edge of
the bow shock is located at about 13 times the radius of the Earth. The collision between the magnetic field
and the solar wind forms the Van Allen radiation belts, a pair of concentric, torus-shaped regions of energetic
charged particles. When the plasma enters the Earth's atmosphere at the magnetic poles, it forms the aurora.
[119]

[edit] Orbit and rotation

[edit] Rotation
Main article: Earth's rotation
 Earth's axial tilt (or obliquity) and its relation to the rotation axis and plane of orbit.
Earth's rotation period relative to the Sun—its mean solar day—is 86,400 seconds of mean solar time.
Each second is slightly longer than an SI second because Earth's solar day is now slightly longer than it was
during the 19th century because of tidal acceleration.[120]
Earth's rotation period relative to the fixed stars, called its stellar day by the International Earth
Rotation and Reference Systems Service (IERS), is 86164.098903691 seconds of mean solar time (UT1), or
23h 56m 4.098903691s. [2][note 16] Earth's rotation period relative to the precessing or moving mean vernal
equinox, misnamed its sidereal day, is 86164.09053083288 seconds of mean solar time (UT1) (23 h 56m
4.09053083288s).[2] Thus the sidereal day is shorter than the stellar day by about 8.4 ms.[121] The length of
the mean solar day in SI seconds is available from the IERS for the periods 1623–2005[122] and 1962–2005.
[123]
Apart from meteors within the atmosphere and low-orbiting satellites, the main apparent motion of
celestial bodies in the Earth's sky is to the west at a rate of 15°/h = 15'/min. For bodies near the celestial
equator, this is equivalent to an apparent diameter of the Sun or Moon every two minutes; from the planet's
surface, the apparent sizes of the Sun and the Moon are approximately the same.[124][125]

[edit] Orbit
Main article: Earth's orbit
Earth orbits the Sun at an average distance of about 150 million kilometers every
365.2564 mean solar days, or one sidereal year. From Earth, this gives an apparent movement of the Sun
eastward with respect to the stars at a rate of about 1°/day, or a Sun or Moon diameter every 12 hours.
Because of this motion, on average it takes 24 hours—a solar day—for Earth to complete a full rotation about
its axis so that the Sun returns to the meridian. The orbital speed of the Earth averages about 30 km/s
(108,000 km/h), which is fast enough to cover the planet's diameter (about 12,600 km) in seven minutes, and
the distance to the Moon (384,000 km) in four hours.[3]
The Moon revolves with the Earth around a common barycenter every 27.32 days relative to the
background stars. When combined with the Earth–Moon system's common revolution around the Sun, the
period of the synodic month, from new moon to new moon, is 29.53 days. Viewed from the celestial north
pole, the motion of Earth, the Moon and their axial rotations are all counter-clockwise. Viewed from a vantage
point above the north poles of both the Sun and the Earth, the Earth appears to revolve in a counterclockwise
direction about the Sun. The orbital and axial planes are not precisely aligned: Earth's axis is tilted some
23.5 degrees from the perpendicular to the Earth–Sun plane, and the Earth–Moon plane is tilted about
5 degrees against the Earth-Sun plane. Without this tilt, there would be an eclipse every two weeks,
alternating between lunar eclipses and solar eclipses.[3][126]
The Hill sphere, or gravitational sphere of influence, of the Earth is about 1.5 Gm (or 1,500,000
kilometers) in radius.[127][note 17] This is maximum distance at which the Earth's gravitational influence is
stronger than the more distant Sun and planets. Objects must orbit the Earth within this radius, or they can
become unbound by the gravitational perturbation of the Sun.
 Illustration of the Milky Way Galaxy, showing the location of the Sun
Earth, along with the Solar System, is situated in the Milky Way galaxy, orbiting about 28,000 light
years from the center of the galaxy. It is currently about 20 light years above the galaxy's equatorial plane in
the Orion spiral arm.[128]

[edit] Axial tilt and seasons

Main article: Axial tilt
 Because of the axial tilt of the Earth, the amount of sunlight reaching any given point on the surface
varies over the course of the year. This results in seasonal change in climate, with summer in the northern
hemisphere occurring when the North Pole is pointing toward the Sun, and winter taking place when the pole
is pointed away. During the summer, the day lasts longer and the Sun climbs higher in the sky. In winter, the
climate becomes generally cooler and the days shorter. Above the Arctic Circle, an extreme case is reached
where there is no daylight at all for part of the year—a polar night. In the southern hemisphere the situation is
exactly reversed, with the South Pole oriented opposite the direction of the North Pole.

Earth and Moon from Mars, imaged by Mars Reconnaissance Orbiter. From space, the Earth can be
seen to go through phases similar to the phases of the Moon.
 By astronomical convention, the four seasons are determined by the solstices—the point in the orbit of
maximum axial tilt toward or away from the Sun—and the equinoxes, when the direction of the tilt and the
direction to the Sun are perpendicular. In the northern hemisphere, Winter Solstice occurs on about
December 21, Summer Solstice is near June 21, Spring Equinox is around March 20 and Autumnal Equinox
is about September 23. In the Southern hemisphere, the situation is reversed, with the Summer and Winter
Solstices exchanged and the Spring and Autumnal Equinox dates switched.[129]
The angle of the Earth's tilt is relatively stable over long periods of time. However, the tilt does
undergo nutation; a slight, irregular motion with a main period of 18.6 years.[130] The orientation (rather than
the angle) of the Earth's axis also changes over time, precessing around in a complete circle over each
25,800 year cycle; this precession is the reason for the difference between a sidereal year and a tropical
year. Both of these motions are caused by the varying attraction of the Sun and Moon on the Earth's
equatorial bulge. From the perspective of the Earth, the poles also migrate a few meters across the surface.
This polar motion has multiple, cyclical components, which collectively are termed quasiperiodic motion. In
addition to an annual component to this motion, there is a 14-month cycle called the Chandler wobble. The
rotational velocity of the Earth also varies in a phenomenon known as length of day variation.[131]
In modern times, Earth's perihelion occurs around January 3, and the aphelion around July 4.
However, these dates change over time due to precession and other orbital factors, which follow cyclical
patterns known as Milankovitch cycles. The changing Earth-Sun distance results in an increase of about
6.9%[132] in solar energy reaching the Earth at perihelion relative to aphelion. Since the southern
hemisphere is tilted toward the Sun at about the same time that the Earth reaches the closest approach to
the Sun, the southern hemisphere receives slightly more energy from the Sun than does the northern over
the course of a year. However, this effect is much less significant than the total energy change due to the
axial tilt, and most of the excess energy is absorbed by the higher proportion of water in the southern
hemisphere.[133]

[edit] Moon
Characteristics
Diameter 3,474.8 km

Mass 7.349 × 1022 kg

Semi-major axis 384,400 km

Orbital period 27 d 7 h 43.7 m

Main article: Moon
The Moon is a relatively large, terrestrial, planet-like satellite, with a diameter about one-quarter of
the Earth's. It is the largest moon in the Solar System relative to the size of its planet, although Charon is
larger relative to the dwarf planet Pluto. The natural satellites orbiting other planets are called "moons" after
Earth's Moon.
The gravitational attraction between the Earth and Moon causes tides on Earth. The same effect on
the Moon has led to its tidal locking: its rotation period is the same as the time it takes to orbit the Earth. As a
result, it always presents the same face to the planet. As the Moon orbits Earth, different parts of its face are
illuminated by the Sun, leading to the lunar phases; the dark part of the face is separated from the light part
by the solar terminator.
 Because of their tidal interaction, the Moon recedes from Earth at the rate of approximately 38 mm a
year. Over millions of years, these tiny modifications—and the lengthening of Earth's day by about 23 µs a
year—add up to significant changes.[134] During the Devonian period, for example, (approximately 410
million years ago) there were 400 days in a year, with each day lasting 21.8 hours.[135]

Details of the Earth-Moon system. Besides the radius of each object, the radius to the Earth-Moon
barycenter is shown. Photos from NASA. Data from NASA. The Moon's axis is located by Cassini's third law.
The Moon may have dramatically affected the development of life by moderating the planet's climate.
Paleontological evidence and computer simulations show that Earth's axial tilt is stabilized by tidal
interactions with the Moon.[136] Some theorists believe that without this stabilization against the torques
applied by the Sun and planets to the Earth's equatorial bulge, the rotational axis might be chaotically
unstable, exhibiting chaotic changes over millions of years, as appears to be the case for Mars.[137] If
Earth's axis of rotation were to approach the plane of the ecliptic, extremely severe weather could result from
the resulting extreme seasonal differences. One pole would be pointed directly toward the Sun during
summer and directly away during winter. Planetary scientists who have studied the effect claim that this
might kill all large animal and higher plant life.[138] However, this is a controversial subject, and further
studies of Mars—whose rotation period and axial tilt are similar to those of Earth, but which lacks a large
moon or liquid core—may settle the matter.
Viewed from Earth, the Moon is just far enough away to have very nearly the same apparent-sized
disk as the Sun. The angular size (or solid angle) of these two bodies match because, although the Sun's
diameter is about 400 times as large as the Moon's, it is also 400 times more distant.[125] This allows total
and annular solar eclipses to occur on Earth.
The most widely accepted theory of the Moon's origin, the giant impact theory, states that it formed
from the collision of a Mars-size protoplanet called Theia with the early Earth. This hypothesis explains
(among other things) the Moon's relative lack of iron and volatile elements, and the fact that its composition is
nearly identical to that of the Earth's crust.[139]
Earth has at least two co-orbital asteroids, 3753 Cruithne and 2002 AA29.[140]

A scale representation of the relative sizes of, and average distance between, Earth and Moon
[edit] Habitability
See also: Planetary habitability

A range of theoretical habitable zones with stars of different mass (our Solar System at center). Scale
is logarithmic, and planet sizes are not to scale.
A planet that can sustain life is termed habitable, even if life did not originate there. The Earth
provides the (currently understood) requisite conditions of liquid water, an environment where complex
organic molecules can assemble, and sufficient energy to sustain metabolism.[141] The distance of the Earth
from the Sun, as well as its orbital eccentricity, rate of rotation, axial tilt, geological history, sustaining
atmosphere and protective magnetic field all contribute to the conditions believed necessary to originate and
sustain life on this planet.[142]

[edit] Biosphere
Main article: Biosphere
The planet's life forms are sometimes said to form a "biosphere". This biosphere is generally believed
to have begun evolving about 3.5 billion years ago. Earth is the only place in the universe where life is known
to exist. Some scientists believe that Earth-like biospheres might be rare.[143]
The biosphere is divided into a number of biomes, inhabited by broadly similar plants and animals.
On land, biomes are separated primarily by differences in latitude, height above sea level and humidity.
Terrestrial biomes lying within the Arctic or Antarctic Circles, at high altitudes or in extremely arid areas are
relatively barren of plant and animal life; species diversity reaches a peak in humid lowlands at equatorial
latitudes.[144]

[edit] Natural resources and land use

Main article: Natural resource
The Earth provides resources that are exploitable by humans for useful purposes. Some of these are
non-renewable resources, such as mineral fuels, that are difficult to replenish on a short time scale.
Large deposits of fossil fuels are obtained from the Earth's crust, consisting of coal, petroleum,
natural gas and methane clathrate. These deposits are used by humans both for energy production and as
feedstock for chemical production. Mineral ore bodies have also been formed in Earth's crust through a
process of Ore genesis, resulting from actions of erosion and plate tectonics.[145] These bodies form
concentrated sources for many metals and other useful elements.
The Earth's biosphere produces many useful biological products for humans, including (but far from
limited to) food, wood, pharmaceuticals, oxygen, and the recycling of many organic wastes. The land-based
ecosystem depends upon topsoil and fresh water, and the oceanic ecosystem depends upon dissolved
nutrients washed down from the land.[146] Humans also live on the land by using building materials to
construct shelters. In 1993, human use of land is approximately:
Arable Permanent Permanent Forests Urban
Land use O
land crops pastures and woodland areas

13.13%
Percentage 4.71%[10] 26% 32% 1.5%
[10]

The estimated amount of irrigated land in 1993 was 2,481,250 km2.[10]

[edit] Natural and environmental hazards

Large areas are subject to extreme weather such as tropical cyclones, hurricanes, or typhoons that
dominate life in those areas. Many places are subject to earthquakes, landslides, tsunamis, volcanic
eruptions, tornadoes, sinkholes, blizzards, floods, droughts, and other calamities and disasters.
Many localized areas are subject to human-made pollution of the air and water, acid rain and toxic
substances, loss of vegetation (overgrazing, deforestation, desertification), loss of wildlife, species extinction,
soil degradation, soil depletion, erosion, and introduction of invasive species.
 According to the United Nations, a scientific consensus exists linking human activities to global
warming due to industrial carbon dioxide emissions. This is predicted to produce changes such as the
melting of glaciers and ice sheets, more extreme temperature ranges, significant changes in weather and a
global rise in average sea levels.[147]

[edit] Human geography

Main article: Human geography
See also: World
 Cartography, the study and practice of map making, and vicariously geography, have historically
been the disciplines devoted to depicting the Earth. Surveying, the determination of locations and distances,
and to a lesser extent navigation, the determination of position and direction, have developed alongside
cartography and geography, providing and suitably quantifying the requisite information.
Earth has approximately 6,803,000,000 human inhabitants as of December 12, 2009.[148]
Projections indicate that the world's human population will reach seven billion in 2013 and 9.2 billion in 2050.
[149] Most of the growth is expected to take place in developing nations. Human population density varies
widely around the world, but a majority live in Asia. By 2020, 60% of the world's population is expected to be
living in urban, rather than rural, areas.[150]
It is estimated that only one-eighth of the surface of the Earth is suitable for humans to live on—three-
quarters is covered by oceans, and half of the land area is either desert (14%),[151] high mountains (27%),
[152] or other less suitable terrain. The northernmost permanent settlement in the world is Alert, on
Ellesmere Island in Nunavut, Canada.[153] (82°28′N) The southernmost is the Amundsen-Scott South Pole
Station, in Antarctica, almost exactly at the South Pole. (90°S)
 The Earth at night, a composite of DMSP/OLS ground illumination data on a simulated night-time
image of the world. This image is not photographic and many features are brighter than they would appear to
a direct observer.
Independent sovereign nations claim the planet's entire land surface, except for some parts of
Antarctica and the odd unclaimed area of Bir Tawil between Egypt and Sudan. As of 2007 there are 201
sovereign states, including the 192 United Nations member states. In addition, there are 59 dependent
territories, and a number of autonomous areas, territories under dispute and other entities.[10] Historically,
Earth has never had a sovereign government with authority over the entire globe, although a number of
nation-states have striven for world domination and failed.[154]
 The United Nations is a worldwide intergovernmental organization that was created with the goal of
intervening in the disputes between nations, thereby avoiding armed conflict.[155] It is not, however, a world
government. The U.N. serves primarily as a forum for international diplomacy and international law. When
the consensus of the membership permits, it provides a mechanism for armed intervention.[156]
The first human to orbit the Earth was Yuri Gagarin on April 12, 1961.[157] In total, about 400 people
visited outer space and reached Earth orbit as of 2004, and, of these, twelve have walked on the Moon.[158]
[159][160] Normally the only humans in space are those on the International Space Station. The station's
crew, currently six people, is usually replaced every six months.[161] The furthest humans have travelled
from Earth is 400,171 km, achieved during the 1970 Apollo 13 mission.[162]
[edit] Cultural viewpoint

The first photograph ever taken by astronauts of an "Earthrise", from Apollo 8

Main article: Earth in culture
The name "Earth" derives from the Anglo-Saxon word erda, which means ground or soil, and is
related to the German word erde. It became eorthe later, and then erthe in Middle English.[163] The standard
astronomical symbol of the Earth consists of a cross circumscribed by a circle.[164]
 Unlike the rest of the planets in the Solar System, mankind did not perceive the Earth as a planet
until the 16th century.[165] Earth has often been personified as a deity, in particular a goddess. In many
cultures the mother goddess is also portrayed as a fertility deity. Creation myths in many religions recall a
story involving the creation of the Earth by a supernatural deity or deities. A variety of religious groups, often
associated with fundamentalist branches of Protestantism[166] or Islam,[167] assert that their interpretations
of these creation myths in sacred texts are literal truth and should be considered alongside or replace
conventional scientific accounts of the formation of the Earth and the origin and development of life.[168]
Such assertions are opposed by the scientific community[169][170] and by other religious groups.[171][172]
[173] A prominent example is the creation-evolution controversy.
In the past there were varying levels of belief in a flat Earth,[174] but this was displaced by the
concept of a spherical Earth due to observation and circumnavigation.[175] The human perspective
regarding the Earth has changed following the advent of spaceflight, and the biosphere is now widely viewed
from a globally integrated perspective.[176][177] This is reflected in a growing environmental movement that
is concerned about humankind's effects on the planet.[178]

[edit] See also

Solar System portal

Earth sciences portal

 Book:Solar System

Books are collections of articles that can be downloaded or ordered in print.

• List of Earth-related topics
• Earth science
• Outline of earth science
• Index of earth science articles
• Geodesy
• History of geodesy
• Geography
• List of basic geography topics
• Index of geography articles
• Geology
• Outline of geology
• Index of geology articles

[edit] Notes
1. ^ All astronomical quantities vary, both secularly and periodically. The quantities given are
the values at the instant J2000.0 of the secular variation, ignoring all periodic variations.
2. ^ a b aphelion = a × (1 + e); perihelion = a × (1 - e), where a is the semi-major axis and e is
the eccentricity.
 3. ^ The reference lists the longitude of the ascending node as -11.26064°, which is equivalent
to 348.73936° by the fact that any angle is equal to itself plus 360°.
4. ^ The reference lists the longitude of perihelion, which is the sum of the longitude of the
ascending node and the argument of perihelion. That is, 114.20783° + (-11.26064°) = 102.94719°.
5. ^ Due to natural fluctuations, ambiguities surrounding ice shelves, and mapping conventions
for vertical datums, exact values for land and ocean coverage are not meaningful. Based on data
from the Vector Map and Global Landcover datasets, extreme values for coverage of lakes and
streams are 0.6% and 1.0% of the earth's surface. The ice shields of Antarctica and Greenland are
counted as land, even though much of the rock which supports them lies below sea level.
6. ^ Blue Planet is used as the title of several films Blue Planet and The Blue Planet, in the Life
issue The Incredible Year '68 featuring the Earthrise photo with lines from poet James Dickey
Behold/The blue planet steeped in its dream/Of reality , and in the title of the European Space Agency
bulletin report Exploring the water cycle of the 'Blue Planet'
7. ^ By International Astronomical Union convention, the term terra is used only for naming
extensive land masses on celestial bodies other than the Earth. Cf. Blue, Jennifer (2007-07-05).
"Descriptor Terms (Feature Types)". Gazetteer of Planetary Nomenclature. USGS.
http://planetarynames.wr.usgs.gov/jsp/append5.jsp. Retrieved 2007-07-05.
8. ^ Other planets in the Solar System are either too hot or too cold to support liquid water.
However, it is confirmed to have existed on the surface of Mars in the past, and may still appear
today. See:
• Malik, Tariq (2007-03-02). "Rover reveals Mars was once wet enough for life".
Space.com (via MSNBC). http://www.msnbc.msn.com/id/4202901/. Retrieved 2007-08-28.
 • Staff (2005-11-07). "Simulations Show Liquid Water Could Exist on Mars". Daily
Headlines (University of Arkansas). http://dailyheadlines.uark.edu/5717.htm. Retrieved 2007-
08-08.
9. ^ As of 2007, water vapor has been detected in the atmosphere of only one extrasolar planet,
and it is a gas giant. See: Tinetti, G.; Vidal-Madjar, A.; Liang, M.C.; Beaulieu, J. P.; Yung, Y.; Carey,
S.; Barber, R. J.; Tennyson, J.; Ribas, I (July 2007). "Water vapour in the atmosphere of a transiting
extrasolar planet". Nature 448 (7150): 169–171. doi:10.1038/nature06002. PMID 17625559.
http://www.nature.com/nature/journal/v448/n7150/abs/nature06002.html.
10.^ The number of solar days is one less than the number of sidereal days because the orbital
motion of the Earth about the Sun results in one additional revolution of the planet about its axis.
11.^ Locally varies between 5 and 200 km.
12.^ Locally varies between 5 and 70 km.
13.^ Including the Somali Plate, which is currently in the process of formation out of the African
Plate. See: Chorowicz, Jean (October 2005). "The East African rift system". Journal of African Earth
Sciences 43 (1–3): 379–410. doi:10.1016/j.jafrearsci.2005.07.019.
14.^ This is the measurement taken by the vessel Kaikō in March 1995 and is believed to be the
most accurate measurement to date. See the Challenger Deep article for more details.
15.^ The total surface area of the Earth is 5.1 × 108 km2. To first approximation, the average
depth would be the ratio of the two, or 2.7 km.
16.^ Aoki, the ultimate source of these figures, uses the term "seconds of UT1" instead of
"seconds of mean solar time".—Aoki, S. (1982). "The new definition of universal time". Astronomy and
Astrophysics 105 (2): 359–361. http://adsabs.harvard.edu/abs/1982A&A...105..359A. Retrieved 2008-
09-23.
17.^ For the Earth, the Hill radius is
 ,
where m is the mass of the Earth, a is an Astronomical Unit, and M is the mass of the Sun. So the

radius in A.U. is about: .

[edit] References
1. ^ a b Standish, E. Myles; Williams, James C. "Orbital Ephemerides of the Sun, Moon, and
Planets" (PDF). International Astronomical Union Commission 4: (Ephemerides). http://iau-
comm4.jpl.nasa.gov/XSChap8.pdf. Retrieved 2010-04-03. See table 8.10.2. Calculation based upon
1 AU = 149,597,870,700(3) m.
2. ^ a b c d Staff (2007-08-07). "Useful Constants". International Earth Rotation and Reference
Systems Service (IERS). http://hpiers.obspm.fr/eop-pc/models/constants.html. Retrieved 2008-09-
23.
3. ^ a b c d e f g h i j k l Williams, David R. (2004-09-01). "Earth Fact Sheet". NASA.
http://nssdc.gsfc.nasa.gov/planetary/factsheet/earthfact.html. Retrieved 2010-08-09.
4. ^ Allen, Clabon Walter; Cox, Arthur N. (2000). Allen's Astrophysical Quantities. Springer.
p. 294. ISBN 0387987460. http://books.google.com/?id=w8PK2XFLLH8C&pg=PA294.
5. ^ Various (2000). David R. Lide. ed. Handbook of Chemistry and Physics (81st ed.). CRC.
ISBN 0849304814.
 6. ^ a b IERS Working Groups (2003). "General Definitions and Numerical Standards". in
McCarthy, Dennis D.; Petit, Gérard. IERS Technical Note No. 32. U.S. Naval Observatory and
Bureau International des Poids et Mesures. http://www.iers.org/MainDisp.csl?pid=46-25776.
Retrieved 2008-08-03.
7. ^ Cazenave, Anny (1995). "Geoid, Topography and Distribution of Landforms". in Ahrens,
Thomas J (PDF). Global earth physics a handbook of physical constants . Washington, DC: American
Geophysical Union. ISBN 0-87590-851-9. http://www.agu.org/reference/gephys/5_cazenave.pdf.
Retrieved 2008-08-03. [dead link]
8. ^ a b Rosenberg, Matt. "What is the circumference of the earth?". About.com.
http://geography.about.com/library/faq/blqzcircumference.htm. Retrieved 2010-04-22.
9. ^ Pidwirny, Michael (2006-02-02). Surface area of our planet covered by oceans and
continents.(Table 8o-1). University of British Columbia, Okanagan.
http://www.physicalgeography.net/fundamentals/8o.html. Retrieved 2007-11-26.
10.^ a b c d e f Staff (2008-07-24). "World". The World Factbook. Central Intelligence Agency.
https://www.cia.gov/library/publications/the-world-factbook/geos/xx.html. Retrieved 2008-08-05.
11.^ Yoder, Charles F. (1995). T. J. Ahrens. ed. Global Earth Physics: A Handbook of Physical
Constants. Washington: American Geophysical Union. p. 12. ISBN 0875908519.
http://www.agu.org/reference/gephys.html. Retrieved 2007-03-17. [dead link]
12.^ Allen, Clabon Walter; Cox, Arthur N. (2000). Allen's Astrophysical Quantities. Springer.
p. 296. ISBN 0387987460. http://books.google.com/?id=w8PK2XFLLH8C&pg=PA296. Retrieved
2010-08-17.
13.^ Arthur N. Cox, ed (2000). Allen's Astrophysical Quantities (4th ed.). New York: AIP Press.
p. 244. ISBN 0-387-98746-0. http://books.google.com/?id=w8PK2XFLLH8C&pg=PA244. Retrieved
2010-08-17.
 14.^ "World: Lowest Temperature". WMO Weather and Climate Extremes Archive. Arizona
State University. http://wmo.asu.edu/world-lowest-temperature. Retrieved 2010-08-07.
15.^ Kinver, Mark (December 10, 2009). "Global average temperature may hit record level in
2010". BBC Online. http://news.bbc.co.uk/2/hi/science/nature/8406839.stm. Retrieved 2010-04-22.
16.^ "World: Highest Temperature". WMO Weather and Climate Extremes Archive. Arizona
State University. http://wmo.asu.edu/world-highest-temperature. Retrieved 2010-08-07.
17.^ May, Robert M. (1988). "How many species are there on earth?". Science 241 (4872):
1441–1449. doi:10.1126/science.241.4872.1441. PMID 17790039.
http://adsabs.harvard.edu/abs/1988Sci...241.1441M. Retrieved 2007-08-14.
18.^ a b See:
• Dalrymple, G.B. (1991). The Age of the Earth. California: Stanford University Press.
ISBN 0-8047-1569-6.
• Newman, William L. (2007-07-09). "Age of the Earth". Publications Services, USGS.
http://pubs.usgs.gov/gip/geotime/age.html. Retrieved 2007-09-20.
• Dalrymple, G. Brent (2001). "The age of the Earth in the twentieth century: a problem
(mostly) solved". Geological Society, London, Special Publications 190: 205–221.
doi:10.1144/GSL.SP.2001.190.01.14.
http://sp.lyellcollection.org/cgi/content/abstract/190/1/205. Retrieved 2007-09-20.
• Stassen, Chris (2005-09-10). "The Age of the Earth". TalkOrigins Archive.
http://www.talkorigins.org/faqs/faq-age-of-earth.html. Retrieved 2008-12-30.
19.^ Harrison, Roy M.; Hester, Ronald E. (2002). Causes and Environmental Implications of
Increased UV-B Radiation. Royal Society of Chemistry. ISBN 0854042652.
20.^ a b c Britt, Robert (2000-02-25). "Freeze, Fry or Dry: How Long Has the Earth Got?".
http://www.space.com/scienceastronomy/solarsystem/death_of_earth_000224.html.
 21.^ a b Carrington, Damian (2000-02-21). "Date set for desert Earth". BBC News.
http://news.bbc.co.uk/1/hi/sci/tech/specials/washington_2000/649913.stm. Retrieved 2007-03-31.
22.^ Yoder, Charles F. (1995). T. J. Ahrens. ed. Global Earth Physics: A Handbook of Physical
Constants. Washington: American Geophysical Union. p. 8. ISBN 0875908519.
http://www.agu.org/reference/gephys.html. Retrieved 2007-03-17. [dead link]
23.^ Bowring, S.; Housh, T. (1995). "The Earth's early evolution". Science 269 (5230): 1535.
doi:10.1126/science.7667634. PMID 7667634.
24.^ Yin, Qingzhu; Jacobsen, S. B.; Yamashita, K.; Blichert-Toft, J.; Télouk, P.; Albarède, F.
(2002). "A short timescale for terrestrial planet formation from Hf-W chronometry of meteorites".
Nature 418 (6901): 949–952. doi:10.1038/nature00995. PMID 12198540.
25.^ Kleine, Thorsten; Palme, Herbert; Mezger, Klaus; Halliday, Alex N. (2005-11-24). "Hf-W
Chronometry of Lunar Metals and the Age and Early Differentiation of the Moon". Science 310
(5754): 1671–1674. doi:10.1126/science.1118842. PMID 16308422.
26.^ Reilly, Michael (October 22, 2009). "Controversial Moon Origin Theory Rewrites History".
http://news.discovery.com/space/moon-earth-formation.html. Retrieved 2010-01-30.
27.^ Canup, R. M.; Asphaug, E. (Fall Meeting 2001). "An impact origin of the Earth-Moon
system". Abstract #U51A-02. American Geophysical Union.
http://adsabs.harvard.edu/abs/2001AGUFM.U51A..02C. Retrieved 2007-03-10.
28.^ Canup, R.; Asphaug, E. (2001). "Origin of the Moon in a giant impact near the end of the
Earth's formation". Nature 412 (6848): 708–712. doi:10.1038/35089010. PMID 11507633.
http://www.nature.com/nature/journal/v412/n6848/abs/412708a0.html.
29.^ Morbidelli, A.; Chambers, J.; Lunine, J. I.; Petit, J. M.; Robert, F.; Valsecchi, G. B.; Cyr, K.
E. (2000). "Source regions and time scales for the delivery of water to Earth". Meteoritics & Planetary
Science 35 (6): 1309–1320. doi:10.1111/j.1945-5100.2000.tb01518.x.
http://adsabs.harvard.edu/abs/2000M&PS...35.1309M. Retrieved 2007-03-06.
30.^ Guinan, E. F.; Ribas, I. "Our Changing Sun: The Role of Solar Nuclear Evolution and
Magnetic Activity on Earth's Atmosphere and Climate". in Benjamin Montesinos, Alvaro Gimenez and
Edward F. Guinan. ASP Conference Proceedings: The Evolving Sun and its Influence on Planetary
Environments. San Francisco: Astronomical Society of the Pacific. ISBN 1-58381-109-5.
http://adsabs.harvard.edu/abs/2002ASPC..269...85G. Retrieved 2009-07-27.
31.^ Staff (March 4, 2010). "Oldest measurement of Earth's magnetic field reveals battle
between Sun and Earth for our atmosphere". Physorg.news.
http://www.physorg.com/news186922627.html. Retrieved 2010-03-27.
32.^ Rogers, John James William; Santosh, M. (2004). Continents and Supercontinents. Oxford
University Press US. p. 48. ISBN 0195165896.
33.^ Hurley, P. M. (Jun 1969). "Pre-drift continental nuclei". Science 164 (3885): 1229–1242.
doi:10.1126/science.164.3885.1229. ISSN 0036-8075. PMID 17772560.
34.^ Armstrong, R. L. (1968). "A model for the evolution of strontium and lead isotopes in a
dynamic earth". Reviews of Geophysics 6: 175–199. doi:10.1029/RG006i002p00175.
35.^ De Smet, J. (2000). "Early formation and long-term stability of continents resulting from
decompression melting in a convecting mantle". Tectonophysics 322: 19. doi:10.1016/S0040-
1951(00)00055-X.
36.^ Harrison, T.; Blichert-Toft, J.; Müller, W.; Albarede, F.; Holden, P.; Mojzsis, S. (December
2005). "Heterogeneous Hadean hafnium: evidence of continental crust at 4.4 to 4.5 ga". Science 310
(5756): 1947–50. doi:10.1126/science.1117926. PMID 16293721.
 37.^ Hong, D. (2004). "Continental crustal growth and the supercontinental cycle: evidence from
the Central Asian Orogenic Belt". Journal of Asian Earth Sciences 23: 799. doi:10.1016/S1367-
9120(03)00134-2.
38.^ Armstrong, R. L. (1991). "The persistent myth of crustal growth". Australian Journal of
Earth Sciences 38: 613–630. doi:10.1080/08120099108727995.
39.^ Murphy, J. B.; Nance, R. D. (1965). "How do supercontinents assemble?". American
Scientist 92: 324–33. doi:10.1511/2004.4.324. http://scienceweek.com/2004/sa040730-5.htm.
Retrieved 2007-03-05.
40.^ Purves, William Kirkwood; Sadava, David; Orians, Gordon H.; Heller, Craig (2001). Life,
the Science of Biology: The Science of Biology. Macmillan. p. 455. ISBN 0716738732.
41.^ Doolittle, W. Ford; Worm, Boris (February 2000). "Uprooting the tree of life". Scientific
American 282 (6): 90–95. doi:10.1038/nature03582.
42.^ Berkner, L. V.; Marshall, L. C. (1965). "On the Origin and Rise of Oxygen Concentration in
the Earth's Atmosphere". Journal of Atmospheric Sciences 22 (3): 225–261. doi:10.1175/1520-
0469(1965)022<0225:OTOARO>2.0.CO;2. http://adsabs.harvard.edu/abs/1965JAtS...22..225B.
Retrieved 2007-03-05.
43.^ Burton, Kathleen (2002-11-29). "Astrobiologists Find Evidence of Early Life on Land".
NASA. http://www.nasa.gov/centers/ames/news/releases/2000/00_79AR.html. Retrieved 2007-03-
05.
44.^ Kirschvink, J. L. (1992). Schopf, J.W.; Klein, C. and Des Maris, D. ed. Late Proterozoic low-
latitude global glaciation: the Snowball Earth . The Proterozoic Biosphere: A Multidisciplinary Study.
Cambridge University Press. pp. 51–52. ISBN 0521366151.
 45.^ Raup, D. M.; Sepkoski, J. J. (1982). "Mass Extinctions in the Marine Fossil Record".
Science 215 (4539): 1501–1503. doi:10.1126/science.215.4539.1501. PMID 17788674.
http://adsabs.harvard.edu/abs/1982Sci...215.1501R. Retrieved 2007-03-05.
46.^ Gould, Stephan J. (October 1994). "The Evolution of Life on Earth". Scientific American.
http://brembs.net/gould.html. Retrieved 2007-03-05.
47.^ Wilkinson, B. H.; McElroy, B. J. (2007). "The impact of humans on continental erosion and
sedimentation". Bulletin of the Geological Society of America 119 (1–2): 140–156.
doi:10.1130/B25899.1. http://bulletin.geoscienceworld.org/cgi/content/abstract/119/1-2/140.
Retrieved 2007-04-22.
48.^ Staff. "Paleoclimatology – The Study of Ancient Climates". Page Paleontology Science
Center. http://www.lakepowell.net/sciencecenter/paleoclimate.htm. Retrieved 2007-03-02.
49.^ a b c Sackmann, I.-J.; Boothroyd, A. I.; Kraemer, K. E. (1993). "Our Sun. III. Present and
Future" (PDF). Astrophysical Journal 418: 457–468. doi:10.1086/173407.
Bibcode: 1993ApJ...418..457S. http://articles.adsabs.harvard.edu/cgi-bin/nph-iarticle_query?
1993ApJ...418..457S&data_type=PDF_HIGH&whole_paper=YES&type=PRINTER&filetype=.pdf.
Retrieved 2008-07-08.
50.^ Kasting, J.F. (1988). "Runaway and Moist Greenhouse Atmospheres and the Evolution of
Earth and Venus". Icarus 74: 472–494. doi:10.1016/0019-1035(88)90116-9. PMID 11538226.
http://adsabs.harvard.edu/abs/1988Icar...74..472K. Retrieved 2007-03-31.
51.^ a b Ward, Peter D.; Brownlee, Donald (2002). The Life and Death of Planet Earth: How the
New Science of Astrobiology Charts the Ultimate Fate of Our World . New York: Times Books, Henry
Holt and Company. ISBN 0-8050-6781-7.
52.^ Li, King-Fai; Pahlevan, Kaveh; Kirschvink, Joseph L.; Yung, Yuk L. (2009). "Atmospheric
Pressure as a Natural Climate Regulator for a Terrestrial Planet with a Biosphere". Proceedings of
the National Academy of Sciences 1–6 (24): 9576–9579. doi:10.1073/pnas.0809436106.
PMID 19487662. PMC 2701016. http://www.gps.caltech.edu/~kfl/paper/Li_PNAS2009.pdf. Retrieved
2009-07-19.
53.^ Guillemot, H.; Greffoz, V. (March 2002). "Ce que sera la fin du monde" (in French). Science
et Vie N° 1014.
54.^ Bounama, Christine (2001). "The fate of Earth's ocean". Hydrology and Earth System
Sciences (Germany: Potsdam Institute for Climate Impact Research) 5 (4): 569–575.
doi:10.5194/hess-5-569-2001. http://www.hydrol-earth-syst-sci.net/5/569/2001/hess-5-569-2001.pdf.
Retrieved 2009-07-03.
55.^ a b Schröder, K.-P.; Connon Smith, Robert (2008). "Distant future of the Sun and Earth
revisited". Monthly Notices of the Royal Astronomical Society 386: 155. doi:10.1111/j.1365-
2966.2008.13022.x. arXiv:0801.4031.
See also Palmer, Jason (2008-02-22). "Hope dims that Earth will survive Sun's death".
NewScientist.com news service. http://space.newscientist.com/article/dn13369-hope-dims-that-
earth-will-survive-suns-death.html?feedId=online-news_rss20. Retrieved 2008-03-24.
56.^ Stern, David P. (2001-11-25). "Planetary Magnetism". NASA.
http://astrogeology.usgs.gov/HotTopics/index.php?/archives/147-Names-for-the-Columbia-
astronauts-provisionally-approved.html. Retrieved 2007-04-01.
57.^ Tackley, Paul J. (2000-06-16). "Mantle Convection and Plate Tectonics: Toward an
Integrated Physical and Chemical Theory". Science 288 (5473): 2002–2007.
doi:10.1126/science.288.5473.2002. PMID 10856206.
58.^ Milbert, D. G.; Smith, D. A. "Converting GPS Height into NAVD88 Elevation with the
GEOID96 Geoid Height Model". National Geodetic Survey, NOAA.
http://www.ngs.noaa.gov/PUBS_LIB/gislis96.html. Retrieved 2007-03-07.
 59.^ a b Sandwell, D. T.; Smith, W. H. F. (2006-07-07). "Exploring the Ocean Basins with
Satellite Altimeter Data". NOAA/NGDC.
http://www.ngdc.noaa.gov/mgg/bathymetry/predicted/explore.HTML. Retrieved 2007-04-21.
60.^ Mohr, P.J.; Taylor, B.N. (October 2000). "Unit of length (meter)". NIST Reference on
Constants, Units, and Uncertainty. NIST Physics Laboratory.
http://physics.nist.gov/cuu/Units/meter.html. Retrieved 2007-04-23.
61.^ Staff (November 2001). "WPA Tournament Table & Equipment Specifications". World Pool-
Billiards Association. http://www.wpa-pool.com/index.asp?content=rules_spec. Retrieved 2007-03-
10.
62.^ Senne, Joseph H. (2000). "Did Edmund Hillary Climb the Wrong Mountain". Professional
Surveyor 20 (5): 16–21.
63.^ Sharp, David (2005-03-05). "Chimborazo and the old kilogram". The Lancet 365 (9462):
831–832. doi:10.1016/S0140-6736(05)71021-7.
64.^ "Tall Tales about Highest Peaks". Australian Broadcasting Corporation.
http://www.abc.net.au/science/k2/moments/s1086384.htm. Retrieved 2008-12-29.
65.^ Brown, Geoff C.; Mussett, Alan E. (1981). The Inaccessible Earth (2nd ed.). Taylor &
Francis. p. 166. ISBN 0045500282. Note: After Ronov and Yaroshevsky (1969).
66.^ Morgan, J. W.; Anders, E. (1980). "Chemical composition of Earth, Venus, and Mercury".
Proceedings of the National Academy of Science 71 (12): 6973–6977. doi:10.1073/pnas.77.12.6973.
PMID 16592930. PMC 350422. http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=350422.
Retrieved 2007-02-04.
67.^ This article incorporates text from a publication now in the public domain: Chisholm,
Hugh, ed (1911). "Petrology". Encyclopædia Britannica (Eleventh ed.). Cambridge University Press.
 68.^ Tanimoto, Toshiro (1995). Thomas J. Ahrens. ed (PDF). Crustal Structure of the Earth.
Washington, DC: American Geophysical Union. ISBN 0-87590-851-9.
http://www.agu.org/reference/gephys/15_tanimoto.pdf. Retrieved 2007-02-03. [dead link]
69.^ Kerr, Richard A. (2005-09-26). "Earth's Inner Core Is Running a Tad Faster Than the Rest
of the Planet". Science 309 (5739): 1313. doi:10.1126/science.309.5739.1313a. PMID 16123276.
70.^ Jordan, T. H. (1979). "Structural Geology of the Earth's Interior". Proceedings National
Academy of Science 76 (9): 4192–4200. doi:10.1073/pnas.76.9.4192. PMID 16592703. PMC 411539.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?artid=411539. Retrieved 2007-03-24.
71.^ Robertson, Eugene C. (2001-07-26). "The Interior of the Earth". USGS.
http://pubs.usgs.gov/gip/interior/. Retrieved 2007-03-24.
72.^ a b Turcotte, D. L.; Schubert, G. (2002). "4". Geodynamics (2 ed.). Cambridge, England,
UK: Cambridge University Press. pp. 136–137. ISBN 978-0-521-66624-4.
73.^ Sanders, Robert (2003-12-10). "Radioactive potassium may be major heat source in
Earth's core". UC Berkeley News.
http://www.berkeley.edu/news/media/releases/2003/12/10_heat.shtml. Retrieved 2007-02-28.
74.^ Alfè, D.; Gillan, M. J.; Vocadlo, L.; Brodholt, J; Price, G. D. (2002). "The ab initio simulation
of the Earth's core" (PDF). Philosophical Transaction of the Royal Society of London 360 (1795):
1227–1244. http://chianti.geol.ucl.ac.uk/~dario/pubblicazioni/PTRSA2002.pdf. Retrieved 2007-02-28.
75.^ Vlaar, N; =Vankeken, P.; Vandenberg, A. (1994). "Cooling of the earth in the Archaean:
Consequences of pressure-release melting in a hotter mantle". Earth and Planetary Science Letters
121: 1. doi:10.1016/0012-821X(94)90028-0.
76.^ Turcotte, D. L.; Schubert, G. (2002). "4". Geodynamics (2 ed.). Cambridge, England, UK:
Cambridge University Press. p. 137. ISBN 978-0-521-66624-4.
 77.^ a b Sclater, John G; Parsons, Barry; Jaupart, Claude (1981). "Oceans and Continents:
Similarities and Differences in the Mechanisms of Heat Loss". Journal of Geophysical Research 86:
11535. doi:10.1029/JB086iB12p11535.
78.^ Richards, M. A.; Duncan, R. A.; Courtillot, V. E. (1989). "Flood Basalts and Hot-Spot
Tracks: Plume Heads and Tails". Science 246 (4926): 103–107. doi:10.1126/science.246.4926.103.
PMID 17837768. http://adsabs.harvard.edu/abs/1989Sci...246..103R. Retrieved 2007-04-21.
79.^ Brown, W. K.; Wohletz, K. H. (2005). "SFT and the Earth's Tectonic Plates". Los Alamos
National Laboratory. http://www.ees1.lanl.gov/Wohletz/SFT-Tectonics.htm. Retrieved 2007-03-02.
80.^ Kious, W. J.; Tilling, R. I. (1999-05-05). "Understanding plate motions". USGS.
http://pubs.usgs.gov/gip/dynamic/understanding.html. Retrieved 2007-03-02.
81.^ Seligman, Courtney (2008). "The Structure of the Terrestrial Planets". Online Astronomy
eText Table of Contents. cseligman.com. http://cseligman.com/text/planets/innerstructure.htm.
Retrieved 2008-02-28.
82.^ Duennebier, Fred (1999-08-12). "Pacific Plate Motion". University of Hawaii.
http://www.soest.hawaii.edu/GG/ASK/plate-tectonics2.html. Retrieved 2007-03-14.
83.^ Mueller, R.D.; Roest, W.R.; Royer, J.-Y.; Gahagan, L.M.; Sclater, J.G. (2007-03-07). "Age
of the Ocean Floor Poster". NOAA. http://www.ngdc.noaa.gov/mgg/fliers/96mgg04.html. Retrieved
2007-03-14.
84.^ Bowring, Samuel A.; Williams, Ian S. (1999). "Priscoan (4.00–4.03 Ga) orthogneisses from
northwestern Canada". Contributions to Mineralogy and Petrology 134: 3.
doi:10.1007/s004100050465.
85.^ Meschede, M.; Udo Barckhausen, U. (2000-11-20). "Plate Tectonic Evolution of the Cocos-
Nazca Spreading Center". Proceedings of the Ocean Drilling Program. Texas A&M University.
http://www-odp.tamu.edu/publications/170_SR/chap_07/chap_07.htm. Retrieved 2007-04-02.
 86.^ Staff. "GPS Time Series". NASA JPL. http://sideshow.jpl.nasa.gov/mbh/series.html.
Retrieved 2007-04-02.
87.^ a b Pidwirny, Michael (2006). "Fundamentals of Physical Geography (2nd Edition)".
PhysicalGeography.net. http://www.physicalgeography.net/fundamentals/7h.html. Retrieved 2007-
03-19.
88.^ Kring, David A. "Terrestrial Impact Cratering and Its Environmental Effects". Lunar and
Planetary Laboratory. http://www.lpi.usra.edu/science/kring/epo_web/impact_cratering/intro/.
Retrieved 2007-03-22.
89.^ Staff. "Layers of the Earth". Volcano World.
http://volcano.oregonstate.edu/vwdocs/vwlessons/plate_tectonics/part1.html. Retrieved 2007-03-11.
90.^ Jessey, David. "Weathering and Sedimentary Rocks". Cal Poly Pomona.
http://geology.csupomona.edu/drjessey/class/Gsc101/Weathering.html. Retrieved 2007-03-20.
91.^ de Pater, Imke; Lissauer, Jack J. (2010). Planetary Sciences (2nd ed.). Cambridge
University Press. p. 154. ISBN 0521853710.
92.^ Wenk, Hans-Rudolf; Bulakh, Andreĭ Glebovich (2004). Minerals: their constitution and
origin. Cambridge University Press. p. 359. ISBN 0521529581.
93.^ FAO Staff (1995). FAO Production Yearbook 1994 (Volume 48 ed.). Rome, Italy: Food and
Agriculture Organization of the United Nations. ISBN 9250038445.
94.^ Sverdrup, H. U.; Fleming, Richard H. (1942-01-01). The oceans, their physics, chemistry,
and general biology. Scripps Institution of Oceanography Archives.
http://repositories.cdlib.org/sio/arch/oceans/. Retrieved 2008-06-13.
95.^ "7,000 m Class Remotely Operated Vehicle KAIKO 7000". Japan Agency for Marine-Earth
Science and Technology (JAMSTEC).
http://www.jamstec.go.jp/e/about/equipment/ships/kaiko7000.html. Retrieved 2008-06-07.
 96.^ Charette, Matthew A.; Smith, Walter H. F. (June 2010). "The Volume of Earth's Ocean".
Oceanography 23 (2): 112–114.
http://www.tos.org/oceanography/issues/issue_archive/issue_pdfs/23_2/23-2_charette.pdf. Retrieved
2010-06-04.
97.^ Shiklomanov, Igor A.; et al. (1999). "World Water Resources and their use Beginning of the
21st century Prepared in the Framework of IHP UNESCO". State Hydrological Institute, St.
Petersburg. http://webworld.unesco.org/water/ihp/db/shiklomanov/. Retrieved 2006-08-10.
98.^ Kennish, Michael J. (2001). Practical handbook of marine science. Marine science series
(3rd ed.). CRC Press. p. 35. ISBN 0849323916.
99.^ Mullen, Leslie (2002-06-11). "Salt of the Early Earth". NASA Astrobiology Magazine.
http://www.astrobio.net/news/article223.html. Retrieved 2007-03-14.
100.^ Morris, Ron M. "Oceanic Processes". NASA Astrobiology Magazine.
http://seis.natsci.csulb.edu/rmorris/oxy/oxy4.html. Retrieved 2007-03-14.
101.^ Scott, Michon (2006-04-24). "Earth's Big heat Bucket". NASA Earth Observatory.
http://earthobservatory.nasa.gov/Study/HeatBucket/. Retrieved 2007-03-14.
102.^ Sample, Sharron (2005-06-21). "Sea Surface Temperature". NASA.
http://science.hq.nasa.gov/oceans/physical/SST.html. Retrieved 2007-04-21.
103.^ Geerts, B.; Linacre, E. (November 1997). "The height of the tropopause". Resources in
Atmospheric Sciences. University of Wyoming. http://www-
das.uwyo.edu/~geerts/cwx/notes/chap01/tropo.html. Retrieved 2006-08-10.
104.^ a b Staff (2003-10-08). "Earth's Atmosphere". NASA.
http://www.nasa.gov/audience/forstudents/9-12/features/912_liftoff_atm.html. Retrieved 2007-03-21.
 105.^ a b Moran, Joseph M. (2005). "Weather". World Book Online Reference Center.
NASA/World Book, Inc. http://www.nasa.gov/worldbook/weather_worldbook.html. Retrieved 2007-03-
17.
106.^ a b Berger, Wolfgang H. (2002). "The Earth's Climate System". University of California,
San Diego. http://earthguide.ucsd.edu/virtualmuseum/climatechange1/cc1syllabus.shtml. Retrieved
2007-03-24.
107.^ Rahmstorf, Stefan (2003). "The Thermohaline Ocean Circulation". Potsdam Institute for
Climate Impact Research. http://www.pik-potsdam.de/~stefan/thc_fact_sheet.html. Retrieved 2007-
04-21.
108.^ Various (1997-07-21). "The Hydrologic Cycle". University of Illinois.
http://ww2010.atmos.uiuc.edu/(Gh)/guides/mtr/hyd/home.rxml. Retrieved 2007-03-24.
109.^ Sadava, David E.; Heller, H. Craig; Orians, Gordon H. (2006). Life, the Science of Biology
(8th ed.). MacMillan. p. 1114. ISBN 0716776715.
110.^ Staff. "Climate Zones". UK Department for Environment, Food and Rural Affairs.
http://www.ace.mmu.ac.uk/eae/Climate/Older/Climate_Zones.html. Retrieved 2007-03-24.
111.^ Staff (2004). "Stratosphere and Weather; Discovery of the Stratosphere". Science Week.
http://scienceweek.com/2004/rmps-23.htm. Retrieved 2007-03-14.
112.^ de Córdoba, S. Sanz Fernández (2004-06-21). "100 km. Altitude Boundary for
Astronautics". Fédération Aéronautique Internationale. http://www.un.org/members/list.shtml.
Retrieved 2007-04-21.
113.^ Liu, S. C.; Donahue, T. M. (1974). "The Aeronomy of Hydrogen in the Atmosphere of the
Earth". Journal of Atmospheric Sciences 31 (4): 1118–1136. doi:10.1175/1520-
0469(1974)031<1118:TAOHIT>2.0.CO;2. http://adsabs.harvard.edu/abs/1974JAtS...31.1118L.
Retrieved 2007-03-02.
 114.^ Catling, David C.; Zahnle, Kevin J.; McKay, Christopher P. (2001). "Biogenic Methane,
Hydrogen Escape, and the Irreversible Oxidation of Early Earth". Science 293 (5531): 839–843.
doi:10.1126/science.1061976. PMID 11486082.
http://www.sciencemag.org/cgi/content/full/293/5531/839.
115.^ Abedon, Stephen T. (1997-03-31). "History of Earth". Ohio State University.
http://www.mansfield.ohio-state.edu/~sabedon/biol1010.htm. Retrieved 2007-03-19.
116.^ Hunten, D. M.; Donahue, T M (1976). "Hydrogen loss from the terrestrial planets". Annual
review of earth and planetary sciences 4: 265–292. doi:10.1146/annurev.ea.04.050176.001405.
http://adsabs.harvard.edu/abs/1976AREPS...4..265H. Retrieved 2008-11-07.
117.^ Fitzpatrick, Richard (2006-02-16). "MHD dynamo theory". NASA WMAP.
http://farside.ph.utexas.edu/teaching/plasma/lectures/node69.html. Retrieved 2007-02-27.
118.^ Campbell, Wallace Hall (2003). Introduction to Geomagnetic Fields. New York:
Cambridge University Press. p. 57. ISBN 0521822068.
119.^ Stern, David P. (2005-07-08). "Exploration of the Earth's Magnetosphere". NASA.
http://www-spof.gsfc.nasa.gov/Education/wmap.html. Retrieved 2007-03-21.
120.^ "Leap seconds". Time Service Department, USNO.
http://tycho.usno.navy.mil/leapsec.html. Retrieved 2008-09-23.
121.^ Seidelmann, P. Kenneth (1992). Explanatory Supplement to the Astronomical Almanac.
Mill Valley, CA: University Science Books. p. 48. ISBN 0-935702-68-7.
122.^ Staff. "IERS Excess of the duration of the day to 86400s ... since 1623". International
Earth Rotation and Reference Systems Service (IERS). http://hpiers.obspm.fr/eop-
pc/earthor/ut1lod/lod-1623.html. Retrieved 2008-09-23. —Graph at end.
123.^ Staff. "IERS Variations in the duration of the day 1962–2005". International Earth Rotation
and Reference Systems Service (IERS). Archived from the original on 2007-08-13.
http://web.archive.org/web/20070813203913/http://hpiers.obspm.fr/eop-
pc/earthor/ut1lod/figure3.html. Retrieved 2008-09-23.
124.^ Zeilik, M.; Gregory, S. A. (1998). Introductory Astronomy & Astrophysics (4th ed.).
Saunders College Publishing. p. 56. ISBN 0030062284.
125.^ a b Williams, David R. (2006-02-10). "Planetary Fact Sheets". NASA.
http://nssdc.gsfc.nasa.gov/planetary/planetfact.html. Retrieved 2008-09-28. —See the apparent
diameters on the Sun and Moon pages.
126.^ Williams, David R. (2004-09-01). "Moon Fact Sheet". NASA.
http://nssdc.gsfc.nasa.gov/planetary/factsheet/moonfact.html. Retrieved 2007-03-21.
127.^ Vázquez, M.; Montañés Rodríguez, P.; Palle, E. (2006). "The Earth as an Object of
Astrophysical Interest in the Search for Extrasolar Planets" (PDF). Instituto de Astrofísica de
Canarias. http://www.iac.es/folleto/research/preprints/files/PP06024.pdf. Retrieved 2007-03-21.
128.^ Astrophysicist team (2005-12-01). "Earth's location in the Milky Way". NASA.
http://imagine.gsfc.nasa.gov/docs/ask_astro/answers/030827a.html. Retrieved 2008-06-11.
129.^ Bromberg, Irv (2008-05-01). "The Lengths of the Seasons (on Earth)". University of
Toronto. http://www.sym454.org/seasons/. Retrieved 2008-11-08.
130.^ Animation of precession of moon orbit
131.^ Fisher, Rick (1996-02-05). "Earth Rotation and Equatorial Coordinates". National Radio
Astronomy Observatory. http://www.cv.nrao.edu/~rfisher/Ephemerides/earth_rot.html. Retrieved
2007-03-21.
132.^ Aphelion is 103.4% of the distance to perihelion. Due to the inverse square law, the
radiation at perihelion is about 106.9% the energy at aphelion.
133.^ Williams, Jack (2005-12-20). "Earth's tilt creates seasons". USAToday.
http://www.usatoday.com/weather/tg/wseason/wseason.htm. Retrieved 2007-03-17.
 134.^ Espenak, F.; Meeus, J. (2007-02-07). "Secular acceleration of the Moon". NASA.
http://sunearth.gsfc.nasa.gov/eclipse/SEcat5/secular.html. Retrieved 2007-04-20.
135.^ Poropudas, Hannu K. J. (1991-12-16). "Using Coral as a Clock". Skeptic Tank.
http://www.skepticfiles.org/origins/coralclo.htm. Retrieved 2007-04-20.
136.^ Laskar, J.; Robutel, P.; Joutel, F.; Gastineau, M.; Correia, A.C.M.; Levrard, B. (2004). "A
long-term numerical solution for the insolation quantities of the Earth". Astronomy and Astrophysics
428: 261–285. doi:10.1051/0004-6361:20041335.
http://adsabs.harvard.edu/abs/2004A&A...428..261L. Retrieved 2007-03-31.
137.^ Murray, N.; Holman, M (2001). "The role of chaotic resonances in the solar system".
Nature 410 (6830): 773–779. doi:10.1038/35071000. PMID 11298438. http://arxiv.org/abs/astro-
ph/0111602v1. Retrieved 2008-08-05.
138.^ Williams, D.M.; J.F. Kasting (1996). "Habitable planets with high obliquities". Lunar and
Planetary Science 27: 1437–1438. http://adsabs.harvard.edu/abs/1996LPI....27.1437W. Retrieved
2007-03-31.
139.^ Canup, R.; Asphaug, E., RM; Asphaug, E (2001). "Origin of the Moon in a giant impact
near the end of the Earth's formation". Nature 412 (6848): 708–712. doi:10.1038/35089010.
PMID 11507633.
140.^ Whitehouse, David (2002-10-21). "Earth's little brother found". BBC News.
http://news.bbc.co.uk/1/hi/sci/tech/2347663.stm. Retrieved 2007-03-31.
141.^ Staff (September 2003). "Astrobiology Roadmap". NASA, Lockheed Martin.
http://astrobiology.arc.nasa.gov/roadmap/g1.html. Retrieved 2007-03-10.
142.^ Dole, Stephen H. (1970). Habitable Planets for Man (2nd ed.). American Elsevier
Publishing Co. ISBN 0-444-00092-5. http://www.rand.org/pubs/reports/R414/. Retrieved 2007-03-11.
 143.^ Ward, P. D.; Brownlee, D. (2000-01-14). Rare Earth: Why Complex Life is Uncommon in
the Universe (1st ed.). New York: Springer-Verlag. ISBN 0387987010.
144.^ Hillebrand, Helmut (2004). "On the Generality of the Latitudinal Gradient". American
Naturalist 163 (2): 192–211. doi:10.1086/381004. PMID 14970922.
145.^ Staff (2006-11-24). "Mineral Genesis: How do minerals form?". Non-vertebrate
Paleontology Laboratory, Texas Memorial Museum.
http://www.utexas.edu/tmm/npl/mineralogy/mineral_genesis/. Retrieved 2007-04-01.
146.^ Rona, Peter A. (2003). "Resources of the Sea Floor". Science 299 (5607): 673–674.
doi:10.1126/science.1080679. PMID 12560541.
http://www.sciencemag.org/cgi/content/full/299/5607/673?
ijkey=AHVbRrqUsmdHY&keytype=ref&siteid=sci. Retrieved 2007-02-04.
147.^ Staff (2007-02-02). "Evidence is now 'unequivocal' that humans are causing global
warming – UN report". United Nations. http://www.un.org/apps/news/story.asp?
NewsID=21429&Cr=climate&Cr1=change. Retrieved 2007-03-07.
148.^ United States Census Bureau (2008-01-07). "World POP Clock Projection". United States
Census Bureau International Database. http://www.census.gov/ipc/www/popclockworld.html.
Retrieved 2008-01-07.
149.^ Staff. "World Population Prospects: The 2006 Revision". United Nations.
http://www.un.org/esa/population/publications/wpp2006/wpp2006.htm. Retrieved 2007-03-07.
150.^ Staff (2007). "Human Population: Fundamentals of Growth: Growth". Population
Reference Bureau.
http://www.prb.org/Educators/TeachersGuides/HumanPopulation/PopulationGrowth/QuestionAnswer
.aspx. Retrieved 2007-03-31.
 151.^ Peel, M. C.; Finlayson, B. L.; McMahon, T. A. (2007). "Updated world map of the Köppen-
Geiger climate classification". Hydrology and Earth System Sciences Discussions 4: 439–473.
doi:10.5194/hessd-4-439-2007. http://www.hydrol-earth-syst-sci-discuss.net/4/439/2007/hessd-4-
439-2007.html. Retrieved 2007-03-31.
152.^ Staff. "Themes & Issues". Secretariat of the Convention on Biological Diversity.
http://www.biodiv.org/programmes/default.shtml. Retrieved 2007-03-29.
153.^ Staff (2006-08-15). "Canadian Forces Station (CFS) Alert". Information Management
Group. http://www.tscm.com/alert.html. Retrieved 2007-03-31.
154.^ Kennedy, Paul (1989). The Rise and Fall of the Great Powers (1st ed.). Vintage.
ISBN 0679720197.
155.^ "U.N. Charter Index". United Nations. http://www.un.org/aboutun/charter/. Retrieved 2008-
12-23.
156.^ Staff. "International Law". United Nations. http://www.un.org/law/. Retrieved 2007-03-27.
157.^ Kuhn, Betsy (2006). The race for space: the United States and the Soviet Union compete
for the new frontier. Twenty-First Century Books. p. 34. ISBN 0822559846.
158.^ Ellis, Lee (2004). Who's who of NASA Astronauts. Americana Group Publishing.
ISBN 0966796144.
159.^ Shayler, David; Vis, Bert (2005). Russia's Cosmonauts: Inside the Yuri Gagarin Training
Center. Birkhäuser. ISBN 0387218947.
160.^ Wade, Mark (2008-06-30). "Astronaut Statistics". Encyclopedia Astronautica.
http://www.astronautix.com/articles/aststics.htm. Retrieved 2008-12-23.
161.^ "Reference Guide to the International Space Station". NASA. 2007-01-16.
http://www.nasa.gov/mission_pages/station/news/ISS_Reference_Guide.html. Retrieved 2008-12-
23.
 162.^ Cramb, Auslan (2007-10-28). "Nasa's Discovery extends space station". Telegraph.
http://www.telegraph.co.uk/earth/earthnews/3311903/Nasas-Discovery-extends-space-station.html.
163.^ Random House Unabridged Dictionary. Random House. July 2005. ISBN 0-375-42599-3.
164.^ Liungman, Carl G. (2004). "Group 29: Multi-axes symmetric, both soft and straight-lined,
closed signs with crossing lines". Symbols – Encyclopedia of Western Signs and Ideograms. New
York: Ionfox AB. pp. 281–282. ISBN 91-972705-0-4.
165.^ Arnett, Bill (July 16, 2006). "Earth". The Nine Planets, A Multimedia Tour of the Solar
System: one star, eight planets, and more. http://nineplanets.org/earth.html. Retrieved 2010-03-09.
166.^ Dutch, S.I. (2002). "Religion as belief versus religion as fact" (PDF). Journal of
Geoscience Education 50 (2): 137–144. http://nagt.org/files/nagt/jge/abstracts/Dutch_v50n2p137.pdf.
Retrieved 2008-04-28.
167.^ Taner Edis (2003) (PDF). A World Designed by God: Science and Creationism in
Contemporary Islam. Amherst: Prometheus. ISBN 1-59102-064-6.
http://www2.truman.edu/~edis/writings/articles/CFI-2001.pdf. Retrieved 2008-04-28.
168.^ * Ross, M.R. (2005). "Who Believes What? Clearing up Confusion over Intelligent Design
and Young-Earth Creationism" (PDF). Journal of Geoscience Education 53 (3): 319.
http://www.nagt.org/files/nagt/jge/abstracts/Ross_v53n3p319.pdf. Retrieved 2008-04-28.
169.^ Pennock, R. T. (2003). "Creationism and intelligent design". Annu Rev Genomics Hum
Genet 4: 143–63. doi:10.1146/annurev.genom.4.070802.110400. PMID 14527300.
170.^ Science, Evolution, and Creationism. Washington, D.C: National Academies Press. 2008.
ISBN 0-309-10586-2. http://books.nap.edu/openbook.php?record_id=11876&page=R1.
171.^ Colburn, A.; Henriques, Laura (2006). "Clergy views on evolution, creationism, science,
and religion". Journal of Research in Science Teaching 43 (4): 419–442. doi:10.1002/tea.20109.
 172.^ Frye, Roland Mushat (1983). Is God a Creationist? The Religious Case Against Creation-
Science. Scribner's. ISBN 0-68417-993-8.
173.^ Gould, S. J. (1997). "Nonoverlapping magisteria" (PDF). Natural History 106 (2): 16–22.
http://www.jbburnett.com/resources/gould_nonoverlapping.pdf. Retrieved 2008-04-28.
174.^ Russell, Jeffrey B. "The Myth of the Flat Earth". American Scientific Affiliation.
http://www.asa3.org/ASA/topics/history/1997Russell.html. Retrieved 2007-03-14. ; but see also
Cosmas Indicopleustes
175.^ Jacobs, James Q. (1998-02-01). "Archaeogeodesy, a Key to Prehistory".
http://www.jqjacobs.net/astro/aegeo.html. Retrieved 2007-04-21.
176.^ Fuller, R. Buckminster (1963). Operating Manual for Spaceship Earth (First ed.). New
York: E.P. Dutton & Co. ISBN 0-525-47433-1. http://www.futurehi.net/docs/OperatingManual.html.
Retrieved 2007-04-21.
177.^ Lovelock, James E. (1979). Gaia: A New Look at Life on Earth (First ed.). Oxford: Oxford
University Press. ISBN 0-19-286030-5.
178.^ For example: McMichael, Anthony J. (1993). Planetary Overload: Global Environmental
Change and the Health of the Human Species. Cambridge University Press. ISBN 0521457599.

[edit] Further reading

• Comins, Neil F. (2001). Discovering the Essential Universe (Second ed.). W. H. Freeman.
ISBN 0-7167-5804-0. http://adsabs.harvard.edu/abs/2003deu..book.....C. Retrieved 2007-03-17.
[edit] External links
Find more about Earth on Wikipedia's sister projects:

Definitions from Wiktionary

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Quotations from Wikiquote

Source texts from Wikisource

Images and media from Commons

News stories from Wikinews

Learning resources from Wikiversity

• USGS Geomagnetism Program

• NASA Earth Observatory
 • Audio - Cain/Gay (2007) Astronomy Cast Earth
• Earth Profile by NASA's Solar System Exploration
• Climate changes cause Earth's shape to change – NASA
• The Gateway to Astronaut Photography of Earth
• "Global Measured Extremes of Temperature and Precipitation" . National Climatic Data
Center. August 20, 2008. http://www.ncdc.noaa.gov/oa/climate/globalextremes.html. Retrieved 2010-
04-22.

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Retrieved from "http://en.wikipedia.org/wiki/Earth"

Categories: Earth | Geography | Geology | Terrestrial planets
 W000

Terre
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir Terre (homonymie).

Terre
 La Terre vue depuis Apollo 17 en 1972[Note 1].
Caractéristiques orbitales
(Époque J2000.0)

Demi-grand axe 149 597 887,5 km

 (1,0000001124 UA)

152 097 701 km

Aphélie
(1,0167103335 UA)

147 098 074 km

Périhélie
(0,9832898912 UA)

924 375 700 km

Circonférence orbitale
(6,1790699007 UA)

Excentricité 0,01671022

Période de révolution 365,25696 d

Période synodique —d

Vitesse orbitale moyenne 29,783 km/s

Vitesse orbitale maximale 30,287 km/s

Vitesse orbitale minimale 29,291 km/s

Inclinaison sur l'écliptique (par définition) 0°

Nœud ascendant 174,873°

 Argument du périhélie 288,064°

Satellites connus 1, la Lune

Caractéristiques physiques

Rayon équatorial 6 378,137 km

Rayon polaire 6 356,752 km

Rayon moyen
6 371,0 km
volumétrique

Aplatissement 0,0033529

40 075,017 ;
Périmètre équatorial
périmètre méridional "polaire" = 40 007,864 km

Superficie 510 067 420 km2

Volume 1,08321×1012 km3

Masse 5,9736×1024 kg

Masse volumique globale 5,515×103 kg/m3

 Gravité de surface 9,780 m/s2
(0,99732 g)

Vitesse de libération 11,186 km/s

Période de rotation 0,99726949 d

(jour sidéral) (23 h 56 min 4,084 s)

Vitesse de rotation
1 674,364 km/h
(à l’équateur)

Inclinaison de l’axe 23,4388°

Albédo géométrique visuel 0,367

Albédo de Bond 0,306

Irradiance solaire 1 367,6 W/m2

(1 Terre)

Température d'équilibre
254,3 K (-18,7 °C)
du corps noir

Température de surface :

Maximum : 57,8 °C[1]

 Moyenne : 15 °C

Minimum : -89,2 °C[2]

Caractéristiques de l’atmosphère

Pression atmosphérique 101 325 Pa

Masse volumique au sol 1,217 kg/m3

Masse totale 5,1×1018 kg

Hauteur 8,5 km

Masse molaire moyenne 28,97 g/mol

Azote N2 78,084 % volume sec

Oxygène O2 20,946 % volume sec

Argon Ar 0,9340 % volume sec

Dioxyde de carbone CO2 390 ppm volume sec

 Néon Ne 18,18 ppm volume sec

Hélium He 5,24 ppm volume sec

Méthane CH4 1,79 ppm volume sec

Krypton Kr 1,14 ppm volume sec

Hydrogène H2 550 ppb volume sec

Protoxyde d'azote N2O 300 ppb volume sec

Monoxyde de carbone CO 100 ppb volume sec

Xénon Xe 90 ppb volume sec

Ozone O3 0 à 70 ppb volume sec

Dioxyde d'azote NO2 20 ppb volume sec

Iode I 10 ppb volume sec

Vapeur d'eau H2O ~ 0,4 % volume global

~ de 1 à 4 % en surface (valeurs typiques)
 Histoire

• Nature planétaire pressentie par

l'école pythagoricienne (Philolaos de Crotone).
Découverte par
• Attestée à
l'époque hellénistique (Aristarque de Samos, puis Ératosthène).

• Ve siècle av. J.-C.

Découverte le
• IIIe siècle av. J.-C.
La Terre est la troisième planète du Système solaire par ordre de distance croissante au Soleil, et la
quatrième par taille et par masse croissantes. C'est la plus grande et la plus massive des quatre planètes
telluriques, les trois autres étant Mercure, Vénus et Mars.
Elle possède un satellite naturel, la Lune, qui est le cinquième plus gros satellite du Système solaire.
Elle est dotée d'un puissant champ magnétique qui dirige vers les pôles les particules chargées
véhiculées par le vent solaire, y provoquant des aurores polaires et générant des ceintures de radiations
concentriques autour du globe résultant de l'accumulation de ces particules piégées dans le champ
magnétique de la Terre. La magnétosphère agit ainsi comme un bouclier protégeant notre planète du vent
solaire.
Couramment appelée en français Terre, planète Terre, planète bleue ou encore Monde[Note 2], c'est
une planète à manteau actif, dotée d'une atmosphère comportant de l'oxygène, recouverte d'eau liquide.
Sommaire
[masquer]
• 1 Histoire
• 2 Composition et structure
• 2.1 Composition chimique
• 2.2 Structure géologique
• 2.3 Plaques tectoniques
• 3 Atmosphère
• 3.1 Constitution
• 3.2 Structure de l'atmosphère
• 4 Données physiques et astronomiques
• 4.1 Forme de la Terre
• 4.2 Données orbitales
• 4.3 Jour, saisons et année
• 4.4 Accélération de la pesanteur
• 4.5 Satellites de la Terre
• 4.6 Position dans l'Univers
• 5 Jour de la Terre
• 6 Notes
• 7 Références
• 8 Voir aussi
Histoire [modifier]
Article détaillé : Histoire de la Terre.
La Terre ainsi que les autres planètes du système solaire se sont formées il y a 4,467 milliards
d'années à partir d'une nébuleuse solaire, masse de poussières et de gaz en forme de disque détachée du
Soleil en formation. Initialement en fusion, la couche externe de la Terre s'est refroidie pour former une
croûte solide. Plus tard, les impacts d'astéroïdes ont causé de nombreux changements sur l'environnement à
la surface. La Lune s'est formée peu de temps après, sans doute à la suite d'une collision avec un objet de la
taille de Mars (quelquefois appelé Théia). Une partie de cet objet se serait agglomérée avec la Terre, tandis
qu'une autre portion, mêlée avec peut-être 10 % de la masse totale de la Terre, aurait été éjectée dans
l'espace, où elle aurait formé la Lune.
L'activité volcanique a produit une atmosphère primitive. De la vapeur d'eau condensée, mêlée à de
la glace apportée par des comètes, a produit les océans. On suppose qu'une activité chimique intense dans
un milieu hautement énergétique a produit une molécule capable de se reproduire, il y a environ 4 milliards
d'années. La vie elle-même serait apparue 500 000 ans plus tard.
L'apparition de la photosynthèse met ensuite l'énergie solaire au service de la vie. Il en résulte en
effet à la fois une accumulation de dioxygène dans l'atmosphère, favorisant la vie animale, et le
développement d'une couche d'ozone[Note 3] dans la haute atmosphère, protégeant la surface de la planète
de l'agression des rayons ultraviolets. Dans ce nouveau cadre, la vie évolue de plus en plus vite vers des
formes toujours plus complexes.
La surface du globe se transforme continuellement, sur des périodes de plusieurs centaines de
millions d'années. Des continents ou supercontinents se forment puis se divisent. C'est ainsi qu'il y a environ
750 millions d'années, le plus vieux des supercontinents connus, Rodinia, commença à se disloquer. Les
continents entre lesquels il s'était divisé se recombinèrent plus tard pour former Pannotia, il y a 650-540
millions d'années, puis finalement Pangée, au Permien, qui se fragmenta il y a 180 millions d'années.
Depuis les années 1960, de nombreuses hypothèses ont été émises dont une qui affirme qu'une (ou
une série) de grande(s) glaciation(s) eut lieu il y a 750 et 580 millions d'années, pendant le
Néoprotérozoïque, et qui couvrit la planète d'une couche de glace. Cette hypothèse a été nommée Snowball
Earth (« Terre boule de neige »), et est d'un intérêt particulier parce qu'elle précède l'explosion cambrienne,
quand des formes de vies multicellulaires commencèrent à proliférer.
À la suite de l'explosion cambrienne, il y a 535 millions d'années, 5 extinctions massives eurent lieu.
La dernière extinction majeure date de 65 millions d'années, quand une présumée météorite est entrée en
collision avec la Terre, exterminant les dinosaures et d'autres grands reptiles, épargnant de plus petits
animaux comme les mammifères, oiseaux, lézards, etc. Dans les 65 millions d'années qui se sont écoulées
depuis, les mammifères se sont diversifiés, l'espèce humaine s'étant développée depuis deux millions
d'années. Des changements périodiques à long terme de l'orbite de la Terre, causés par l'influence
gravitationnelle des autres astres, sont probablement une des causes des glaciations qui ont couvert une
bonne partie de la planète. À l'issue de la dernière glaciation, le développement de l'agriculture et, ensuite,
des civilisations, permit aux humains de modifier la surface de la Terre dans une courte période de temps,
comme aucune autre espèce avant lui sur terre, affectant la nature tout comme les autres formes de vies.

Composition et structure [modifier]

La Terre est une planète tellurique, c'est-à-dire une planète essentiellement rocheuse à noyau
métallique, contrairement aux géantes gazeuses, telles que Jupiter, essentiellement constituées de gaz
légers (hydrogène et hélium).
 Il s'agit de la plus grande des quatre planètes telluriques du système solaire, que ce soit en termes
de taille ou de masse. De ces quatre planètes, la Terre a aussi la masse volumique globale la plus élevée, la
plus forte gravité de surface et le plus puissant champ magnétique global. Cependant, plusieurs planètes
telluriques plus grandes que la Terre ont été découvertes en dehors du système solaire, parmi lesquelles
l'exoplanète Gliese 581 c, qui possède un diamètre 50 % supérieur à celui de la Terre. Plusieurs missions
sont en cours, ou prévues, afin de découvrir de nouvelles planètes similaires à la Terre, appelées exoterres.
La surface externe de la Terre est divisée en plusieurs segments rigides, ou plaques tectoniques, qui
se déplacent lentement sur la surface sur une durée de plusieurs millions d'années. Environ 71 % de la
surface est couverte d'océans d'eau salée, les 29 % restants consistant en continents et îles. L'eau liquide,
nécessaire à la vie telle que nous la connaissons, est très abondante sur Terre, et aucune autre planète n'a
encore été découverte avec des étendues d'eau liquide (lacs, mers, océans) à sa surface.

Composition chimique [modifier]

La masse de la Terre est d'approximativement 5,98×1024 kg. Elle est composée principalement de
fer (32,1 %[3]), d'oxygène (30,1 %), de silicium (15,1 %), de magnésium (19,9 %), de soufre (2,9 %), de
nickel (1,8 %), de calcium (1,5 %) et d'aluminium (1,4 %), le 1,2 % restant consistant en de légères traces
d'autres éléments. Les éléments les plus denses ayant tendance à se concentrer au centre de la Terre
(phénomène de différenciation planétaire), on pense que le cœur de la Terre est composé majoritairement
de fer (88,8 %), avec une plus petite quantité de nickel (5,8 %), de soufre (4,5 %) et moins de 1 % d'autres
éléments.
Le géochimiste F. W. Clarke a calculé que 47 % (en poids) de la croûte terrestre est faite d'oxygène,
présent principalement sous forme d'oxydes, dont les principaux sont les oxydes de silicium, aluminium, fer,
calcium, magnésium, potassium et sodium. La silice est le constituant majeur de la croûte, sous forme de
pyroxénoïdes, les minéraux les plus communs des roches magmatiques et métamorphiques. Après une
synthèse basée sur l'analyse de 1 672 types de roches, Clarke a obtenu les pourcentages présentés dans le
tableau ci-dessous.
Pourcentage
Oxyde
(pondéral)

Silice (SiO2) 59,71

Oxyde d'aluminium (Al2O3) 15,41

Oxyde de calcium (CaO) 4,90

Oxyde de magnésium (MgO) 4,36

Oxyde de sodium (Na2O) 3,55

Oxyde de fer(II) (FeO) 3,52

Oxyde de potassium (K2O) 2,80

Oxyde de fer(III) (Fe2O3) 2,63

 Eau (H2O) 1,52

Dioxyde de titane (TiO2) 0,60

Pentoxyde de phosphore (P2O5) 0,22

Total 99,22

Structure géologique [modifier]

 Structure de la Terre.
1. croûte continentale,
2. croûte océanique,
3. manteau supérieur,
4. manteau inférieur,
5. noyau externe,
6. noyau interne,
A : Discontinuité de Mohorovicic,
B : Discontinuité de Gutenberg,
C : Discontinuité de Lehmann.
Article détaillé : Structure interne de la Terre.
La Terre est constituée de plusieurs couches internes identifiables à peu près concentriques : la
croûte terrestre (océanique ou continentale), le manteau supérieur, le manteau inférieur, et le noyau externe
et interne. La lithosphère est constituée de la croûte et de la zone superficielle du manteau supérieur.
L'asthénosphère est la zone plus profonde du manteau supérieur (en dessous de la lithosphère).
La croûte terrestre est relativement jeune par rapport à la Terre elle-même. Pendant la période
relativement courte d'environ 500 millions d'années pendant laquelle l'érosion et les processus tectoniques
ont détruit, puis recréé, la plupart des couches superficielles de la Terre, la presque totalité des traces de
l'histoire géologique de sa surface (cratères d'impact, par exemple) ont disparu.
Plus de 99 % de la surface terrestre aurait moins de 2 milliards d'années.
 La structure interne de la Terre est connue au moyen de l'étude de la propagation des ondes
sismiques entre une source et différents points de la surface terrestre.
La vitesse d'une onde sismique change en effet assez brutalement au passage entre deux couches
de composition ou phase minérale différentes. Ces limites ont parfois reçu des noms particuliers, tels que la
discontinuité de Mohorovicic, la discontinuité de Lehmann ou la discontinuité de Gutenberg.
La constitution de la Terre s'explique par son mode de formation, par accrétion de météorites, qui a
produit une stratification en phase fluide par masse volumique décroissante depuis les couches internes vers
les couches externes.
La plus grande partie de la chaleur interne de la Terre (87 %) est produite par la radioactivité des
roches qui constituent la croûte terrestre : radioactivité naturelle produite par la désintégration de l'uranium,
du thorium et du potassium.

Plaques tectoniques [modifier]

Selon la théorie de la tectonique des plaques, la partie supérieure de l'intérieur de la Terre est
composée de deux couches : la lithosphère, comprenant la croûte, et la partie solide du manteau. Au-
dessous de la lithosphère se trouve l'asthénosphère, qui forme le cœur du manteau. L'asthénosphère
ressemble à du liquide extrêmement chaud et visqueux.
La lithosphère flotte essentiellement sur l'asthénosphère et est brisée en pièces qui sont appelées
plaques tectoniques. Ces plaques sont des segments rigides qui bougent en relation avec les autres de trois
façons : en convergence, en divergence, et par transcurrence. C'est ainsi que sont créés les tremblements
de terre, l'activité volcanique ainsi que les montagnes.
 Certaines plaques ont une plus petite superficie comme la plaque indienne, la plaque arabique, la
plaque caraïbe et la plaque de Nazca à l'ouest de la côte de l'Amérique du Sud. La plaque australienne s'est
fusionnée quelque peu à la plaque indienne il y a 50 à 55 millions d'années. Les plaques les plus rapides
dans leur mouvement sont les plaques océaniques, se déplaçant d'environ 70 Millimètresmm/an. À l'opposé,
la plaque la plus lente est la plaque eurasienne, progressant d'environ 21 Millimètresmm/an.
Les principales plaques tectoniques sont :
Aire
Nom de la
Carte totale, en (106 Couvre
plaque
km2)

Plaque africaine 61,3 Afrique

Carte des plaques tectoniques terrestres. Les
flèches indiquent les mouvements relatifs de chaque Plaque
60,9 Antarctique
plaque. antarctique

Plaque
47,2 Australie
australienne

Plaque Asie et
67,8
eurasienne l'Europe

Plaque nord- 75,9 Amérique

 du Nord et Nord-Est
américaine
de la Sibérie

Plaque sud- Amérique

43,6
américaine du Sud

Océan
Plaque pacifique 103,3
Pacifique
Atmosphère [modifier]
 Schéma des couches de l'atmosphère.
La Terre est entourée d'une enveloppe gazeuse qu'elle retient par attraction gravitationnelle :
l'atmosphère. L'atmosphère de la Terre est intermédiaire entre celle, très épaisse, de Vénus, et celle, très
ténue, de Mars : sa pression au niveau de la mer est en moyenne de 101 325 Pa, soit 1 atm par définition.
Outre une proportion variable de vapeur d'eau comprise entre 0 et 4 %, elle est constituée de 78,09 %
d'azote, 20,95 % d'oxygène,0,93 % d'argon et 0,039 %9 % de dioxyde de carbone, ainsi que de divers autres
gaz. Ce taux élevé d'oxygène est unique dans le Système solaire, et résulte de l'activité photosynthétique
des organismes chlorophylliens : la Terre est, en effet, le seul astre connu pour abriter la vie, conséquence
probable du fait que c'est également le seul astre connu pour avoir des conditions de température et de
pression permettant l'existence d'eau liquide en surface.
Cette atmosphère donne à la planète un reflet bleuté depuis l'espace, d'où son surnom de « planète
bleue ». La constitution et la densité de l'atmosphère sont telles que la lumière incidente du Soleil et la
lumière réfléchie par les continents et les mers sont diffractées ; donnant sa couleur au ciel, et par réflexion,
aux étendues d'eau.

Constitution [modifier]
Cette enveloppe, dont la masse globale est de l'ordre de 5×10 18 kg (un millionième de la masse de
la Terre), est contenue à 99 % dans les 30 premiers kilomètres (50 % dans les 5 premiers kilomètres).
La basse atmosphère (du niveau de la mer jusqu'à environ 45 km) est composée de gaz
« permanents », gaz dont les proportions restent constantes, et de gaz de concentration variable avec
l'altitude.
 • Le diazote, le dioxygène et l'argon constituent, en volume, 99,997 % des gaz permanents
(voir tableau ci-dessus) ; le brassage vertical de l'air permet de conserver une répartition constante à
tous les niveaux, même pour les gaz les plus légers, tels que l'hélium ou l'hydrogène.
• Les gaz à concentration variable sont essentiellement la vapeur d'eau H2O ; et dans une
moindre mesure le dioxyde de carbone CO2, le dioxyde de soufre SO2 et l'ozone O3.

L'atmosphère terrestre peut être considérée, à un instant donné, comme un mélange

thermodynamique d'air sec et de vapeur d'eau.
Les particules liquides, solides, ou mixtes, en suspension dans l'atmosphère constituent l'aérosol
atmosphérique.
Ces particules jouent un rôle primordial dans les phénomènes de condensation (nuages) et de
formation de cristaux de glace, ainsi qu'à différents processus physico-chimiques dans l'atmosphère. Leur
concentration varie de plusieurs puissances de 10 (de plusieurs ordres de grandeurs) en fonction du lieu et
du temps ; en concentration élevée, elles constituent un facteur de pollution. Les particules se classent en :
• particules d'Aitken : 1 nm < d < 0,1 µm
• grosses particules : 0,1 µm < d < 5 µm
• particules géantes : 5 µm < d < 50 µm environ
L'atmosphère atténue de façon importante le rayonnement solaire reçu au sol ; suivant l'importance
de la couverture nuageuse, le sol reçoit de 68 % à 28 % (ou moins) du rayonnement solaire parvenant à
l'atmosphère, un flux solaire initial de 1 370 W/m2.
Structure de l'atmosphère [modifier]
La composition chimique de l'atmosphère, sa température, ou les phénomènes qui y sont observés
présentent des discontinuités marquées lorsque l'altitude augmente. Ces discontinuités correspondent à des
couches homogènes dont les propriétés évoluent de façon continue ; ce sont (par altitude croissante) :
• la troposphère
• la stratosphère
• la mésosphère
• la thermosphère
• l'exosphère
Les limites de ces couches (d'altitude variable) ont reçu des désignations particulières : tropopause,
stratopause, mésopause et thermopause.

Données physiques et astronomiques [modifier]

Forme de la Terre [modifier]

La forme de la Terre est modélisée par un ellipsoïde, légèrement aplati aux pôles, et plus
précisément par le géoïde. Le diamètre approximatif de référence est de 12 742 km.
La rotation de la Terre crée un léger bourrelet équatorial, de sorte que le diamètre à l’équateur est
43 km plus long que le diamètre polaire (du pôle Nord au pôle Sud). Les plus grandes dénivellations du sol
de la Terre sont l'Everest (8 848 m au-dessus du niveau de la mer) et la fosse des Mariannes (11 500 m
sous le niveau de la mer). Par contre, à cause de l’aplatissement, l’objet le plus éloigné du cœur de la Terre
est en fait le volcan Chimborazo en Équateur.

Comparaison de quelques caractéristiques physiques des quatre planètes telluriques

In
Planète Rayon équatorial Masse Gravité

2 439,7 km (0,383 e23/3.3023,302×102

Mercure[4] 3,701 m/s² (0,377 g) ~
Terre) 3 kg (0,055 Terre)

e24/4.86854,8685×1
Vénus[5] 6 051,8 km (0,95 Terre) 8,87 m/s² (0,904 g) 1
024 kg (0,815 Terre)

e24/5.97365,9736×1 9,780 m/s² (0,99732

Terre[6] 6 378,14 km 2
024 kg g)

3 402,45 km (0,533 e23/6.41856,4185×1

Mars[7] 3,69 m/s² (0,376 g) 2
Terre) 023 kg (0,107 Terre)
 Photomontage comparatif des tailles des planètes telluriques (de gauche à droite) : Mercure, Vénus, la Terre et Ma

La conception sphérique de la Terre remonte à l'antiquité grecque, vers le Ve siècle av. J.-C., et plus
spécifiquement aux pythagoriciens. On retrouve cette conception chez Parménide, Platon ou Aristote. Elle
s'appuie sur le fait que, lors des éclipses de Lune, l'ombre de la Terre est sphérique, et sur le fait que les
constellations varient lorsqu'on se déplace du Nord au Sud[8]. Au IIIe siècle av. J.-C., Ératosthène donna une
estimation du rayon terrestre que nous supposons[Note 5] très proche de la réalité, ainsi que Posidonios, au
Ie siècle av. J.-C.. Vers la même époque, Cléomède définit les notions d'équateur, de tropiques, d'arctique et
d'antarctique. Reprenant ces notions, le géographe Ptolémée fournit au IIe siècle des informations
géographiques qui furent utilisées jusqu'à la Renaissance.
 La civilisation arabo-musulmane conserva la connaissance d'une Terre sphérique et au IXe siècle, le
calife Al-Mamun, à Bagdad, fit procéder à une mesure d'une partie d'un méridien, conduisant à une bonne
approximation de la circonférence de la Terre. Plus explicite encore est qu'une figure comme Abou Hanîfa,
fondateur de l'une des quatre écoles de jurisprudence musulmane, avait foi dans la sphéricité de la Terre[9].
Dans le monde chrétien, cette idée fut parfois remise en cause, par exemple au VIe siècle par Cosmas
Indicopleustès. En effet, les Pères de l'Église ne pouvaient concilier la vision du monde sphérique d'Aristote
constitué de deux zones polaires et deux zones tempérées, séparées par une zone torride infranchissable,
avec l'universalité du message du Christ, ce message ne pouvant parvenir à d'hypothétiques[Note 6]
habitants des antipodes. Jusqu'au XIIe siècle, on s'attacha donc à représenter le monde sous forme
symbolique, mais des philosophes ou des religieux tels Isidore de Séville, Bède le Vénérable, Jean Scot
Erigène, Gerbert d'Aurillac, Thomas d'Aquin, Albert le Grand ou Roger Bacon avaient très bien intégré la
représentation sphérique. Vers 1150, le livre De imagine mundi, dont l'auteur présumé est Honorius d'Autun,
décrit l'univers de façon plus mythologique que scientifique, mais dans lequel la Terre est une sphère
d'environ 35 000 km de circonférence. Charlemagne est d'ailleurs représenté sur quelques enluminures
comme tenant à la main une représentation d'un petit globe terrestre surmonté d'une croix.
 Position et taille du continent asiatique selon Christophe Colomb.
Les récits de voyages de missionnaires, de Marco Polo et de l'explorateur Jean de Mandeville (avec
son Livre des merveilles du monde) diffusaient dans la société l'image d'une terre sphérique, qui pouvait
théoriquement faire l'objet d'une « circumnavigation ». L’Imago mundi du cardinal Pierre d'Ailly retenait cette
représentation sphérique. On sait que Christophe Colomb a été influencé par le Livre des merveilles du
monde de Mandeville, et qu'il possédait un exemplaire de l’Imago mundi abondamment annoté et commenté
par ses soins. En sous-estimant grandement le rayon terrestre et en imaginant un continent asiatique trois
fois plus étendu vers l'Est qu'il ne l'est en réalité, Colomb a pu envisager de façon raisonnable la possibilité
de rejoindre les Indes par l'Ouest. Une connaissance plus précise des distances aurait découragé toute
tentative de traversée de l'Océan avec les moyens de l'époque.
 Les voyages des Portugais dès le début du XVe siècle pour rejoindre les Indes en contournant
l'Afrique, la redécouverte des textes grecs à la Renaissance, en particulier la Géographie de Ptolémée, leur
diffusion au moyen de l'imprimerie ont également largement contribué à propager les représentations
modernes de la Terre, avec le Nord vers le haut des cartes, les méridiens, les parallèles, l'équateur et les
deux tropiques. Le plus ancien globe terrestre connu est fabriqué par Martin Behaim vers la fin du
XVe siècle, peu avant que Vasco de Gama, Christophe Colomb ou Magellan entreprennent leurs voyages.
On y voit l'Europe, l'Afrique et l'Asie, mais bien entendu, ni les Amériques, ni l'Océanie.
Mercator a, en dessinant ses cartes, mentionné et dessiné un énorme continent austral : Terra
incognita australis (terre australe (du sud) inconnue). Cette « terre australe inconnue » a été dessinée au
Sud car Mercator pensait, à la suite des Grecs, que sans ce poids la Terre n'était pas équilibrée. Les
réflexions et travaux en géographie (relevés cartographiques, projection de Mercator) au XVIe siècle ont
permis de faire évoluer la connaissance de la Terre.
C'est au XVIIIe siècle que l'aplatissement des pôles est reconnu, avec les expéditions menées au
Pérou et en Laponie.

Données orbitales [modifier]

Le demi-grand axe de son orbite définit l'unité astronomique, qui vaut 149 597 870 700 ± 3 m selon
la valeur actuellement la plus précise[10]. Son éloignement au Soleil est compris entre 0,9833 et 1,0167 UA.

Jour, saisons et année [modifier]

Article connexe : Année (astronomie).
 La Terre tourne autour d'elle même en un jour sidéral de 23h56', et autour du soleil en une année
d'environ 365 jours. Son axe de rotation est incliné par rapport à l'écliptique, ce qui produit l'alternance de
quatre saisons sur la surface du globe.
La périodicité des saisons se mesure, elle, au moyen de l'année tropique, légèrement plus courte
que l'année sidérale de la valeur annuelle de la précession de l'axe de rotation. La période du passage aux
points de l'ellipse de l'orbite comme le périhélie, est l'année anomalistique, légèrement plus longue que
l'année sidérale de la valeur annuelle de la récession du périhélie.
Enfin, dans les calculs astronomiques, on utilise l'année julienne valant exactement 365,25 jours.

Accélération de la pesanteur [modifier]

Pour consulter un article plus général, voir : Pesanteur.
L'accélération de la pesanteur (ou « champ de pesanteur ») varie légèrement à la surface de la Terre
pour trois raisons :
• Elle dépend de l'altitude, l'accélération étant inversement proportionnelle au carré de la
distance entre le centre de gravité de la Terre et le point où il est mesuré.
• La Terre n'est pas parfaitement sphérique, mais un peu aplatie aux pôles, la gravitation est
plus grande aux pôles, pour la même raison.
• La Terre tourne sur elle-même, ce qui fait qu'un objet à l'équateur est un tout petit peu plus
léger (voir Force centrifuge).
D'autres facteurs peuvent influer de façon minime sur le champ de pesanteur local (Voir
Gravimétrie) :
 • La composition du sous-sol (roches, grottes…)
L'accélération de la pesanteur peut se calculer comme suit :
g=9,780318 [m/s2] × (1 + 5,3024×10–3 × sin2(L) + 5,9×10–6 × sin2(2×L) – 3,15×10–7 × h)
où :
• L=la latitude
• h=l'altitude en mètre.
Au niveau de la mer, h=0 m :
• à l'équateur (L=0°) : g=9,7803 m/s²
• à la latitude (L=45°) : g=9,8063 m/s²
• aux pôles (L=90°) : g=9,8322 m/s²
La force de pesanteur englobe donc à la fois la gravité et la force centrifuge, qui elle provient du
mouvement de rotation. Il reste une force provenant de la rotation de la Terre qui ne peut être incluse dans la
pesanteur : la force de Coriolis, qui elle dépend de la vitesse de l'objet sur Terre. La pesanteur a été définie
pour être indépendante du mouvement de l'objet sur Terre.

Satellites de la Terre [modifier]

Article connexe : Lune.
 Représentation à l'échelle de la taille et de la distance de la Terre et de la Lune. (1 px = 500 km)
La Lune est un satellite naturel, situé à environ 380 500 km de la Terre. Relativement grand, son
diamètre est environ le quart de celui de la Terre. Au sein du système solaire, c'est l'un des plus grands
satellites naturels (après Ganymède, Titan, Callisto et Io), le plus grand d'une planète non gazeuse. Elle est
relativement proche de la taille de la planète Mercure. Les satellites naturels orbitant autour des autres
planètes sont communément appelés « lunes » en référence à la Lune de la Terre.
L'attraction gravitationnelle entre la Terre et la Lune cause les marées sur Terre. Le même effet a
lieu sur la Lune, faisant en sorte que sa période de rotation est identique au temps qu'il lui faut pour orbiter
autour de la Terre, présentant ainsi toujours la même face vers notre planète. En orbitant autour de la Terre,
différentes parties du côté visible de la Lune sont illuminées par le Soleil, causant les phases lunaires.
À cause du couple des marées, la Lune s'éloigne de la Terre à un rythme d'environ 38 mm par an,
produisant aussi l'allongement du jour terrestre de 23 microsecondes par an. Sur plusieurs millions d'années,
l'effet cumulé de ces petites modifications produit d'importants changements. Durant la période du Dévonien,
à approximativement 410 millions d'années d'aujourd'hui, il y avait 400 jours dans une année, chaque jour
durant 21,8 heures.
Vue de la Terre, la Lune est assez éloignée pour avoir la même taille apparente que le Soleil. La
taille angulaire des deux corps est quasiment égale car même si le diamètre du Soleil est 400 fois plus grand
que celui de la Lune, celle-ci est 400 fois plus rapprochée de la Terre que ce dernier. Ceci permet encore
tout juste les éclipses totales sur Terre.
La théorie acceptée sur les origines de la Lune est celle d'un impact géant entre un planètoïde de la
taille de Mars, appelé Théia, et la Terre nouvellement formée. Cette hypothèse explique en partie le fait que
la composition de la Lune ressemble particulièrement à celle de la croûte terrestre.
La Terre a aussi deux satellites co-orbitaux, l'astéroïde (3753) Cruithne et 2002 AA29, ainsi que des
quasi-satellites, 2003 YN107 (jusqu'en 2006), 2001 GO2 et (164207) 2004 GU9.

Position dans l'Univers [modifier]

On sait aujourd'hui que la Terre tourne sur elle-même et autour du Soleil. Mais cette formulation
sous-entend un certain nombre de principes liés au développement de l'astronomie et de la physique, et qui
ont mis plusieurs siècles avant de s'affirmer.
La position de la Terre dans l'Univers fut la source de longs débats opposant durant des siècles
philosophes, savants et religieux de tous bords. Pendant longtemps, la Terre fut considérée comme au
centre de l'Univers, conception défendue par Aristote ou Ptolémée. Cette théorie, le géocentrisme, affirmait
que tous les objets célestes, Soleil, Lune, planètes et étoiles, (astres) orbitaient autour de la Terre.
L'héliocentrisme présente une Terre en orbite autour du Soleil, avec la Lune, satellite naturel de la Terre.
Défendue par Aristarque de Samos, cette théorie fut oubliée jusqu'à ce que Nicolas Copernic la redécouvre
et la complète dans son traité publié en 1543 : De revolutionibus orbium coelestium libri VI ). Tycho Brahe
proposa un système dans lequel le Soleil tournait autour de la Terre, et les autres planètes autour du Soleil.
L'idée de Copernic fut soutenue par Johannes Kepler et Galilée. Les ellipses képlériennes y firent beaucoup
en permettant des tables d'une précision jamais égalée, et en expliquant les variations de latitude des
planètes par rapport au plan de l'écliptique. À la fin du XVIIème, Isaac Newton introduisit sa mécanique qui
explique le mouvement de la Terre par les forces s'appliquant sur elle, en se plaçant dans un espace qu'il
supposait absolu. On disposait alors d'un système mécanique expliquant les mouvements des planètes,
conformes à la théorie héliocentrique.
Dans la modélisation actuelle de la mécanique newtonienne, on ne retient plus l'idée d'espace
absolu. La position des objets est définie par rapport à un référentiel, notion déjà présente chez Galilée,
Huygens et Newton, et la mécanique newtonienne repose sur l'hypothèse de l'existence d'une classe de
référentiels privilégiés, les référentiels galiléens. Dans un référentiel galiléen, les mouvements des planètes
s'expliquent par les seules forces de gravitation.
Dans le langage courant, quand rien n'est précisé, on se réfère à un référentiel galiléen. Cela est dû
au statut privilégié des référentiels galiléens en mécanique. C'est pour cela que la phrase « la Terre tourne »
n'est pas considérée comme imprécise en physique.
Il faut remarquer qu'en vertu du principe de relativité, y compris dans la version de Galilée, les effets
induits par le mouvement de la Terre sont extrêmement faibles pour ceux qui s'y trouvent. C'est pour cela
que les Anciens avaient pu supposer la Terre immobile.
Plus généralement les astrophysiciens considèrent aujourd'hui que non seulement l'Univers est
dépourvu de centre, mais même de tout point privilégié. Cette conjecture est appelée principe de Copernic,
puisque le nom de Copernic est associé à la fin de la vision de la Terre comme centre de l'univers.
 Le drapeau non-officiel du Jour de la Terre.

Jour de la Terre [modifier]

Le Jour de la Terre a lieu le 22 avril, depuis 1970. Il marque la création de la mouvance écologiste.

Notes [modifier]
1. ↑ La Bille bleue, photo prise par l'équipage d'Apollo 17 le 7 décembre 1972. L'année 2009
marque le 50eanniversaire de la première photographie couleur à avoir été envoyée de l'espace le
1er décembre(1959 en science).
2. ↑ La Terre est la seule planète tournant autour du Soleil qui ne soit pas dotée d'un nom
universel pour toutes les langues issues de la mythologie grecque ou romaine. En français toutefois,
« Terre » s'apparente à Terra, déesse romaine de la terre, Gaïa en grec.
 3. ↑ L'ozone est une des variante moléculaire de l'oxygène O 3.
4. ↑ La rotation de Vénus étant rétrograde, l’inclinaison de son axe est supérieure à 90°. On
pourrait dire que son axe est incliné de « -2,64° ».
5. ↑ Il utilise en effet comme base de son calcul la distance d'Alexandrie à Syène en stades, la
longueur du « stade » étant approximative à l'époque et la distance d'Alexandrie à Syène ne pouvant
alors s'apprécier que par une durée de marche de chameaux (Ian Stewart et Jack Cohen, The
Science of Discworld)
6. ↑ Augustin d'Hippone émet par exemple des doutes sur la possibilité que soient habitées les
antipodes, ce qui montre que le concept d'antipodes était bien admis.

Références [modifier]
1. ↑ Température relevée à El Azizia, Libye, le 13 septembre 1922 : (en) Global Measured
Extremes of Temperature and Precipitation [archive], National Climatic Data Center. Mis en ligne le
20 août 2008
2. ↑ Température relevée le 21 juillet 1983 à Vostok, Antarctique : (ru) Budretsky, A.B.,
« New absolute minimum of air temperature », dans Bulletin of the Soviet Antarctic Expedition ,
Gidrometeoizdat, Leningrad, no 105, 1984 [ texte intégral [archive] ]
3. ↑ pourcentage pondéral
4. ↑ (en) David R. Williams, « Mercury Fact Sheet [archive] », novembre 2007, NASA,
National Space Science Data Center.
5. ↑ (en) David R. Williams, « Venus Fact Sheet [archive] », avril 2005, NASA, National
Space Science Data Center.
 6. ↑ (en) David R. Williams, « Earth Fact Sheet [archive] », avril 2007, NASA, National
Space Science Data Center.
7. ↑ (en) David R. Williams, « Mars Fact Sheet [archive] », novembre 2007, NASA, National
Space Science Data Center.
8. ↑ cf par exemple Vitruve, De architectura, Livre IX, ch. VII, trad. Coignard (1673) : « J'ai
parlé ici des constellations dont les figures ont été formées dans le Ciel par l'esprit divin qui est
auteur de la Nature, ainsi que le philosophe Démocrite les a désignées ; j'entends seulement celles
qui se lèvent et qui se couchent en notre horizon. Car tout de même que celles qui sont au
septentrion, et qui, faisant leur cours autour du pôle septentrional, ne se couchent point et ne
passent jamais sous la terre, ainsi il y en a d'autres sous la terre qui tournent aussi autour du pôle
méridional, demeurant toujours cachées sans se lever jamais sur la terre ; ce qui fait qu'on ne sait
point quelle est leur figure ; comme il se prouve par l'étoile appelée Canopus, que nous ne
connaissons que par le rapport des marchands qui ont voyagé dans les extrémités de l'Egypte, et
jusqu'au terres qui sont au bout du monde. »
9. ↑ Muhammad Hamidullah. L'Islam et son impulsion scientifique originelle, Tiers-Monde,
1982, vol. 23, n° 92, pp. 789: «... Enfin, l'imam Abou Hanifah (m. 767) savait que la terre était
sphérique ... »
10.↑ (en) IAU WG on NSFA – 10 août 2009 [archive] « Current Best Estimates »

Voir aussi [modifier]

Sur les autres projets Wikimédia :
• Terre sur Wikimedia Commons (ressources multimédia)
 • Terre sur le Wiktionnaire (dictionnaire universel)
• Terre sur Wikibooks (livres pédagogiques)

Articles connexes [modifier]

• Conceptions actuelles
• Monde
• Liste des pays du monde
• Liste des pays du monde par continent
• Géographie
• Cartographie
• Projection cartographique
• Sciences de la Terre et de l'Univers
• Écologie
• Développement durable
• Pollution
• Géologie
• Géodésie
• Géophysique
• Hypothèse Gaïa
• Structure interne du globe terrestre
• Cycle biogéochimique
• Astronomie
 • Planète
• Un point bleu pâle
• Conceptions anciennes
• Ératosthène
• Figure de la Terre dans l'Antiquité
• Figure de la Terre au Moyen Âge
• Figure de la Terre à la Renaissance
• Différents modèles
• le sphéroïde de Clairaut
• le modèle ellipsoïdal
• Le modèle incluant la notion de gravitation universelle

Liens externes [modifier]

• (fr) GéoManips, les mouvements de la Terre, site CNRS/sagascience
• (fr) Observer et interagir en 3D avec la Terre: Voir le jour et la nuit polaires
• (fr) Caractéristiques de la Terre sur le site de l'IMCCE
• (fr) Le-systeme-solaire.net
• (fr) astrosurf.com
• (fr) Informations cartes et statistiques
• (fr) Cours de géologie en ligne
• (en) Moteur de recherche de coordonnées géographiques
 • (en) Logiciel Google Earth
• (en) Vidéo comparant la taille de la Terre à d'autres astres
[Enrouler]
v·d·m
Système solaire

Étoile Soleil

Mercure • Vénus • Terre • Mars • Jupiter • Saturne • Uranus

Planètes
• Neptune

Planètes naines Cérès • Pluton • Haumea • Makemake • Éris

Sedna • Quaoar • (225088) 2007 OR10 • Charon • (84522)

Objets massifs de la ceinture de Kuiper
2002 TC302 • Orcus • Varuna • 2007 UK126 • 2005 QU182

Mercurienne • Vénusienne • Terrestres : Lune • Martiennes •

Joviennes : Io · Europe · Ganymède · Callisto • Saturniennes : Titan
Lunes et lunes astéroïdales
• Uraniennes • Neptuniennes : Triton • Plutonniennes •
Haumeainnes • Érisienne

Anneaux Joviens• Saturniens • Uraniens • Neptuniens • Plutoniens •

 de Ganymède • de Callisto • d'Europe • de Rhéa

Astéroïdes (liste) : Pallas · Junon · Vesta • Comètes :

Petits corps
1P/Halley · 2P/Encke • Damocloïde • Météoroïdes
Vulcanoïde • Ceinture d’astéroïdes • Centaure • Ceinture de
Principales zones Kuiper • Disque d’objets épars • Ceinture intermédiaire • Nuage de
Hills • Nuage d’Oort

Héliosphère • Héliopause • Héliogaine • Formation et

Autres évolution du système solaire • Milieu interplanétaire • Planètes
hypothétiques• C/1992 J1

Liste des objets du système solaire classés par : taille • masse • distance au Soleil

• Portail de la géodésie et de la géophysique

• Portail des sciences de la Terre et de l’Univers

• Portail de l’astronomie

• Portail de la géographie

• Portail de la biologie
Ce document provient de « http://fr.wikipedia.org/wiki/Terre ».

Catégories : Terre | Planète tellurique | [+]

 w000

Abiogenesis
From Wikipedia, the free encyclopedia

Jump to: navigation, search

"Primordial soup" redirects here. For the board game, see Primordial Soup (board game).
"Origin of life" redirects here. For views on the origins of life outside the natural sciences, see
Creation myth.
 Pre-Cambrian stromatolites in the Siyeh Formation, Glacier National Park. In 2002, William Schopf of
UCLA published a paper in the scientific journal Nature arguing that geological formations such as this
possess 3.5 Ga (billion years old) fossilized cyanobacteria microbes. If true, they would be evidence of the
earliest known life on earth.
In natural science, abiogenesis (pronounced /ˌeɪbaɪ.ɵˈdʒɛnɨsɪs/, AY-bye-oh-JEN-ə-siss) or biopoesis
is the study of how life on Earth could have arisen from inanimate matter. It should not be confused with
evolution, which is the study of how groups of already living things change over time, or with cosmogony,
which covers how the universe might have arisen. Most amino acids, often called "the building blocks of life",
can form via natural chemical reactions unrelated to life, as demonstrated in the Miller–Urey experiment and
similar experiments, which involved simulating some of the conditions of the early Earth, in a scientific
laboratory.[1] In all living things, these amino acids are organized into proteins, and the construction of these
proteins is mediated by nucleic acids. Which of these organic molecules first arose and how they formed the
first life is the focus of abiogenesis.
In any theory of abiogenesis, two aspects of life have to be accounted for: replication, and
metabolism. The question of which came first gave rise to different types of theories. In the beginning,
metabolism-first theories (Oparin coacervate) were proposed, and only later thinking gave rise to modern,
replication-first approach.
In modern, still somewhat limited understanding, the first living things on Earth are thought to be
single cell prokaryotes (which lack a cell nucleus), perhaps evolved from protobionts (organic molecules
surrounded by a membrane-like structure).[2] The oldest ancient fossil microbe-like objects are dated to be
3.5 Ga (billion years old), approximately one billion years after the formation of the Earth itself.[3][4] By
2.4 Ga, the ratio of stable isotopes of carbon, iron and sulfur shows the action of living things on inorganic
minerals and sediments[5][6] and molecular biomarkers indicate photosynthesis, demonstrating that life on
Earth was widespread by this time.[7][8]
The sequence of chemical events that led to the first nucleic acids is not known. Several hypotheses
about early life have been proposed, most notably the iron-sulfur world theory (metabolism without genetics)
and the RNA world hypothesis (RNA life-forms).
Contents
[hide]
• 1 Conceptual history
• 1.1 Spontaneous generation
• 1.2 Pasteur and Darwin
• 1.3 "Primordial soup" theory
• 2 Early conditions
• 3 Current models
• 3.1 Origin of organic molecules
• 3.1.1 "Soup" theory today: Miller's
experiment and subsequent work
• 3.1.1.1 Regarding the reducing
atmosphere
• 3.1.1.2 Regarding monomer
formation
• 3.1.1.3 Regarding monomer
accumulation
• 3.1.1.4 Regarding further
transformation
• 3.1.2 The deep sea vent theory
• 3.1.3 Fox's experiments
• 3.1.4 Eigen's hypothesis
[edit] Conceptual history
[edit] Spontaneous generation
Main article: Spontaneous generation
Until the early 19th century, people generally believed in the ongoing spontaneous generation of
certain forms of life from non-living matter. This was paired with heterogenesis, the belief that one form of life
derives from a different form (e.g. bees from flowers).[9] Classical notions of abiogenesis, now more
precisely known as spontaneous generation, held that certain complex, living organisms are generated by
decaying organic substances. According to Aristotle it was a readily observable truth that aphids arise from
the dew which falls on plants, fleas from putrid matter, mice from dirty hay, crocodiles from rotting logs at the
bottom of bodies of water, and so on.[10]
In the 17th century, such assumptions started to be questioned; for example, in 1646, Sir Thomas
Browne published his Pseudodoxia Epidemica (subtitled Enquiries into Very many Received Tenets, and
Commonly Presumed Truths), which was an attack on false beliefs and "vulgar errors." His conclusions were
not widely accepted. For example, his contemporary, Alexander Ross wrote: "To question this (i.e.,
spontaneous generation) is to question reason, sense and experience. If he doubts of this let him go to
Egypt, and there he will find the fields swarming with mice, begot of the mud of Nylus, to the great calamity of
the inhabitants."[11]
In 1665, Robert Hooke published the first drawings of a microorganism. Hooke was followed in 1676
by Anton van Leeuwenhoek, who drew and described microorganisms that are now thought to have been
protozoa and bacteria.[12] Many felt the existence of microorganisms was evidence in support of
spontaneous generation, since microorganisms seemed too simplistic for sexual reproduction, and asexual
reproduction through cell division had not yet been observed.
The first solid evidence against spontaneous generation came in 1668 from Francesco Redi, who
proved that no maggots appeared in meat when flies were prevented from laying eggs. It was gradually
shown that, at least in the case of all the higher and readily visible organisms, the previous sentiment
regarding spontaneous generation was false. The alternative seemed to be biogenesis: that every living thing
came from a pre-existing living thing (omne vivum ex ovo, Latin for "every living thing from an egg").
In 1768, Lazzaro Spallanzani demonstrated that microbes were present in the air, and could be killed
by boiling. In 1861, Louis Pasteur performed a series of experiments which demonstrated that organisms
such as bacteria and fungi do not spontaneously appear in sterile, nutrient-rich media.
[edit] Pasteur and Darwin

Charles Darwin in 1879.

By the middle of the 19th century, the theory of biogenesis had accumulated so much evidential
support, due to the work of Louis Pasteur and others, that the alternative theory of spontaneous generation
had been effectively disproven. Pasteur himself remarked, after a definitive finding in 1864, "Never will the
doctrine of spontaneous generation recover from the mortal blow struck by this simple experiment."[13] The
collapse of spontaneous generation, however, left a vacuum of scientific thought on the question of how life
had first arisen.
 In a letter to Joseph Dalton Hooker on February 1, 1871,[14] Charles Darwin addressed the question,
suggesting that the original spark of life may have begun in a "warm little pond, with all sorts of ammonia and
phosphoric salts, lights, heat, electricity, etc. present, so that a protein compound was chemically formed
ready to undergo still more complex changes". He went on to explain that "at the present day such matter
would be instantly devoured or absorbed, which would not have been the case before living creatures were
formed."[15] In other words, the presence of life itself makes the search for the origin of life dependent on the
sterile conditions of the laboratory.
[edit] "Primordial soup" theory

Alexander Oparin (right) at the laboratory.

Further information: Miller–Urey experiment
No new notable research or theory on the subject appeared until 1924, when Alexander Oparin
reasoned that atmospheric oxygen prevents the synthesis of certain organic compounds that are necessary
building blocks for the evolution of life. In his The Origin of Life,[16][17] Oparin proposed that the
"spontaneous generation of life" that had been attacked by Louis Pasteur, did in fact occur once, but was
now impossible because the conditions found in the early earth had changed, and the presence of living
organisms would immediately consume any spontaneously generated organism. Oparin argued that a
"primeval soup" of organic molecules could be created in an oxygen-less atmosphere through the action of
sunlight. These would combine in ever-more complex fashions until they formed coacervate droplets. These
droplets would "grow" by fusion with other droplets, and "reproduce" through fission into daughter droplets,
and so have a primitive metabolism in which those factors which promote "cell integrity" survive, and those
that do not become extinct. Many modern theories of the origin of life still take Oparin's ideas as a starting
point.
Around the same time, J. B. S. Haldane suggested that the Earth's pre-biotic oceans–very different
from their modern counterparts–would have formed a "hot dilute soup" in which organic compounds could
have formed. This idea was called biopoiesis or biopoesis, the process of living matter evolving from self-
replicating but nonliving molecules.[18][19]

[edit] Early conditions

Main article: Timeline of evolution
Morse and MacKenzie have suggested that oceans may have appeared first in the Hadean eon, as
soon as two hundred million years (200 Ma) after the Earth was formed, in a hot 100 °C (212 °F) reducing
environment, and that the pH of about 5.8 rose rapidly towards neutral.[20] This has been supported by
Wilde[3] who has pushed the date of the zircon crystals found in the metamorphosed quartzite of Mount
Narryer in Western Australia, previously thought to be 4.1–4.2 Ga, to 4.404 Ga. This means that oceans and
continental crust existed within 150 Ma of Earth's formation.
 Despite this, the Hadean environment was one highly hazardous to life. Frequent collisions with large
objects, up to 500 kilometres (310 mi) in diameter, would have been sufficient to vaporise the ocean within a
few months of impact, with hot steam mixed with rock vapour leading to high altitude clouds completely
covering the planet. After a few months the height of these clouds would have begun to decrease but the
cloud base would still have been elevated for about the next thousand years. After that, it would have begun
to rain at low altitude. For another two thousand years rains would slowly have drawn down the height of the
clouds, returning the oceans to their original depth only 3,000 years after the impact event.[21]
Between 3.8 and 4.1 Ga, changes in the orbits of the gaseous giant planets may have caused a late
heavy bombardment that pockmarked the moon and other inner planets (Mercury, Mars, and presumably
Earth and Venus). This would likely have sterilized the planet had life appeared before that time.
By examining the time interval between such devastating environmental events, the time interval
when life might first have come into existence can be found for different early environments. The study by
Maher and Stevenson shows that if the deep marine hydrothermal setting provides a suitable site for the
origin of life, abiogenesis could have happened as early as 4.0 to 4.2 Ga, whereas if it occurred at the
surface of the earth abiogenesis could only have occurred between 3.7 and 4.0 Ga.[22]
Other research suggests a colder start to life. Work by Leslie Orgel and colleagues on the synthesis
of purines has shown that freezing temperatures are advantageous, due to the concentrating effect for key
precursors such as HCN.[23] Research by Stanley Miller and colleagues suggested that while adenine and
guanine require freezing conditions for synthesis, cytosine and uracil may require boiling temperatures.[24]
Based on this research, Miller suggested a beginning of life involving freezing conditions and exploding
meteorites.[25] An article in Discover Magazine points to research by the Miller group indicating the formation
of seven different amino acids and 11 types of nucleobases in ice when ammonia and cyanide were left in a
freezer from 1972–1997.[26][27] This article also describes research by Christof Biebricher showing the
formation of RNA molecules 400 bases long under freezing conditions using an RNA template, a single-
strand chain of RNA that guides the formation of a new strand of RNA. As that new RNA strand grows, it
adheres to the template.[28] The explanation given for the unusual speed of these reactions at such a low
temperature is eutectic freezing. As an ice crystal forms, it stays pure: only molecules of water join the
growing crystal, while impurities like salt or cyanide are excluded. These impurities become crowded in
microscopic pockets of liquid within the ice, and this crowding causes the molecules to collide more often.
Evidence of the early appearance of life comes from the Isua supercrustal belt in Western Greenland
and from similar formations in the nearby Akilia Islands. Carbon entering into rock formations has a ratio of
Carbon-13 (13C) to Carbon-12 (12C) of about −5.5 (in units of δ13C), where because of a preferential biotic
uptake of 12C, biomass has a δ13C of between −20 and −30. These isotopic fingerprints are preserved in the
sediments, and Mojzis has used this technique to suggest that life existed on the planet already by 3.85
billion years ago.[29] Lazcano and Miller (1994) suggest that the rapidity of the evolution of life is dictated by
the rate of recirculating water through mid-ocean submarine vents. Complete recirculation takes 10 million
years, thus any organic compounds produced by then would be altered or destroyed by temperatures
exceeding 300 °C (572 °F). They estimate that the development of a 100 kilobase genome of a DNA/protein
primitive heterotroph into a 7000 gene filamentous cyanobacterium would have required only 7 Ma.[30]

[edit] Current models

There is no truly "standard model" of the origin of life. Most currently accepted models draw at least
some elements from the framework laid out by the Oparin-Haldane hypothesis. Under that umbrella,
however, are a wide array of disparate discoveries and conjectures such as the following, listed in a rough
order of postulated emergence:
 1. Some theorists suggest that the atmosphere of the early Earth may have been chemically
reducing in nature, composed primarily of methane (CH4), ammonia (NH3), water (H2O), hydrogen
sulfide (H2S), carbon dioxide (CO2) or carbon monoxide (CO), and phosphate (PO43-), with
molecular oxygen (O2) and ozone (O3) either rare or absent.
2. In such a reducing atmosphere, electrical activity can catalyze the creation of certain basic
small molecules (monomers) of life, such as amino acids. This was demonstrated in the Miller–Urey
experiment by Stanley L. Miller and Harold C. Urey in 1953.
3. Phospholipids (of an appropriate length) can spontaneously form lipid bilayers, a basic
component of the cell membrane.
4. A fundamental question is about the nature of the first self-replicating molecule. Since
replication is accomplished in modern cells through the cooperative action of proteins and nucleic
acids, the major schools of thought about how the process originated can be broadly classified as
"proteins first" and "nucleic acids first".
5. The principal thrust of the "nucleic acids first" argument is as follows:
1. The polymerization of nucleotides into random RNA molecules might have resulted
in self-replicating ribozymes (RNA world hypothesis)
2. Selection pressures for catalytic efficiency and diversity might have resulted in
ribozymes which catalyse peptidyl transfer (hence formation of small proteins), since
oligopeptides complex with RNA to form better catalysts. The first ribosome might have been
created by such a process, resulting in more prevalent protein synthesis.
3. Synthesized proteins might then outcompete ribozymes in catalytic ability, and
therefore become the dominant biopolymer, relegating nucleic acids to their modern use,
predominantly as a carrier of genomic information.
 As of 2010, no one has yet synthesized a "protocell" using basic components which would have the
necessary properties of life (the so-called "bottom-up-approach"). Without such a proof-of-principle,
explanations have tended to be short on specifics. However, some researchers are working in this field,
notably Steen Rasmussen at Los Alamos National Laboratory and Jack Szostak at Harvard University.
Others have argued that a "top-down approach" is more feasible. One such approach, attempted by Craig
Venter and others at The Institute for Genomic Research, involves engineering existing prokaryotic cells with
progressively fewer genes, attempting to discern at which point the most minimal requirements for life were
reached. The biologist John Desmond Bernal coined the term Biopoesis for this process[citation needed],
and suggested that there were a number of clearly defined "stages" that could be recognised in explaining
the origin of life.
• Stage 1: The origin of biological monomers
• Stage 2: The origin of biological polymers
• Stage 3: The evolution from molecules to cell
Bernal suggested that evolution may have commenced early, some time between Stage 1 and 2[31].

[edit] Origin of organic molecules

There are two possible sources of organic molecules on the early Earth:
1. Terrestrial origins–organic synthesis driven by impact shocks or by other energy sources
(such as ultraviolet light or electrical discharges) (eg.Miller's experiments)
2. Extraterrestrial origins–delivery by objects (e.g. carbonaceous chondrites) or gravitational
attraction of organic molecules or primitive life-forms from space
 Recently, estimates of these sources suggest that the heavy bombardment before 3.5 Ga within the
early atmosphere made available quantities of organics comparable to those produced by other energy
sources.[32]

[edit] "Soup" theory today: Miller's experiment and subsequent work

Biochemist Robert Shapiro has summarized the "Primordial Soup" theory of Oparin and Haldane in
its "mature form" as follows:[33]
1. The early Earth had a chemically reducing atmosphere, as discussed above.
2. This atmosphere, exposed to energy in various forms, produced simple organic compounds
("monomers").
3. These compounds accumulated in a "soup", which may have been concentrated at various
locations (Shorelines, oceanic vents etc.).
4. By further transformation, more complex organic polymers— and ultimately life— developed in
the soup.

[edit] Regarding the reducing atmosphere

Whether the mixture of gases used in the Miller–Urey experiment truly reflects the atmospheric
content of early Earth is a controversial topic. Other less reducing gases produce a lower yield and variety. It
was once thought that appreciable amounts of molecular oxygen were present in the prebiotic
atmosphere[citation needed], which would have essentially prevented the formation of organic molecules;
however, the current scientific consensus is that such was not the case. (See Oxygen catastrophe).
[edit] Regarding monomer formation
Main article: Miller experiment
One of the most important pieces of experimental support for the "soup" theory came in 1953. A
graduate student, Stanley Miller, and his professor, Harold Urey, performed an experiment that demonstrated
how organic molecules could have spontaneously formed from inorganic precursors, under conditions like
those posited by the Oparin-Haldane Hypothesis. The now-famous "Miller–Urey experiment" used a highly
reduced mixture of gases–methane, ammonia and hydrogen–to form basic organic monomers, such as amino
acids.[34] This provided direct experimental support for the second point of the "soup" theory as described
above, and it is around the remaining two points of the theory that much of the debate now centers.
Apart from the Miller–Urey experiment, described above, the next most important step in research on
prebiotic organic synthesis was the demonstration by John Oró that the nucleic acid purine base, adenine,
was formed by heating aqueous ammonium cyanide solutions.[35] In support of abiogenesis in eutectic ice
(see above), more recent work demonstrated the formation of s-triazines (alternative nucleobases),
pyrimidines (including cytosine and uracil), and adenine from urea solutions subjected to freeze-thaw cycles
under a reductive atmosphere (with spark discharges as an energy source).[36]

[edit] Regarding monomer accumulation

The "soup" theory relies on the assumption proposed by Darwin (see above) that in an environment
with no pre-existing life, organic molecules may have accumulated and provided an environment for chemical
evolution.

[edit] Regarding further transformation

 The spontaneous formation of complex polymers from abiotically generated monomers under the
conditions posited by the "soup" theory is not at all a straightforward process. Besides the necessary basic
organic monomers, compounds that would have prohibited the formation of polymers were formed in high
concentration during the Miller–Urey and Oró experiments. The Miller experiment, for example, produces
many substances that would undergo cross-reactions with the amino acids or terminate the peptide chain.
More fundamentally, it can be argued that the most crucial challenge unanswered by this theory is
how the relatively simple organic building blocks polymerise and form more complex structures, interacting in
consistent ways to form a protocell. For example, in an aqueous environment hydrolysis of
oligomers/polymers into their constituent monomers would be favored over the condensation of individual
monomers into polymers.

[edit] The deep sea vent theory

The deep sea vent, or hydrothermal vent, theory for the origin of life on Earth posits that life may
have begun at submarine hydrothermal vents, where hydrogen-rich fluids emerge from below the sea floor
and interface with carbon dioxide-rich ocean water. Sustained chemical energy in such systems is derived
from redox reactions, in which electron donors, such as molecular hydrogen, react with electron acceptors,
such as carbon dioxide (see iron-sulfur world theory).

[edit] Fox's experiments

In the 1950s and 1960s, Sidney W. Fox studied the spontaneous formation of peptide structures
under conditions that might plausibly have existed early in Earth's history. He demonstrated that amino acids
could spontaneously form small peptides. These amino acids and small peptides could be encouraged to
form closed spherical membranes, called protenoid microspheres, which show many of the basic
characteristics of 'life'.[37]

[edit] Eigen's hypothesis

In the early 1970s the problem of the origin of life was approached by Manfred Eigen and Peter
Schuster of the Max Planck Institute for Biophysical Chemistry. They examined the transient stages between
the molecular chaos and a self-replicating hypercycle in a prebiotic soup.[38]
In a hypercycle, the information storing system (possibly RNA) produces an enzyme, which catalyzes
the formation of another information system, in sequence until the product of the last aids in the formation of
the first information system. Mathematically treated, hypercycles could create quasispecies, which through
natural selection entered into a form of Darwinian evolution. A boost to hypercycle theory was the discovery
that RNA, in certain circumstances, forms itself into ribozymes, capable of catalyzing their own chemical
reactions.[39] However, these reactions are limited to self-excisions (in which a longer RNA molecule
becomes shorter), and much rarer small additions that are incapable of coding for any useful protein. The
hypercycle theory is further degraded since the hypothetical RNA would require the existence of complex
biochemicals such as nucleotides which are not formed under the conditions proposed by the Miller–Urey
experiment.

[edit] Hoffmann's contributions

Geoffrey W. Hoffmann, a student of Eigen, contributed to the concept of life involving both replication
and metabolism emerging from catalytic noise. His contributions included showing that an early sloppy
translation machinery can be stable against an error catastrophe of the type that had been envisaged as
problematical by Leslie Orgel ("Orgel's paradox")[40][41] and calculations regarding the occurrence of a set
of required catalytic activities together with the exclusion of catalytic activities that would be disruptive. This is
called the stochastic theory of the origin of life.[42]

[edit] Wächtershäuser's hypothesis

Main article: iron-sulfur world theory
Deep-sea black smoker
 Another possible answer to this polymerization conundrum was provided in 1980s by the German
chemist Günter Wächtershäuser, in his iron-sulfur world theory. In this theory, he postulated the evolution of
(bio)chemical pathways as fundamentals of the evolution of life. Moreover, he presented a consistent system
of tracing today's biochemistry back to ancestral reactions that provide alternative pathways to the synthesis
of organic building blocks from simple gaseous compounds.
In contrast to the classical Miller experiments, which depend on external sources of energy (such as
simulated lightning or UV irradiation), "Wächtershäuser systems" come with a built-in source of energy,
sulfides of iron and other minerals (e.g. pyrite). The energy released from redox reactions of these metal
sulfides is not only available for the synthesis of organic molecules, but also for the formation of oligomers
and polymers. It is therefore hypothesized that such systems may be able to evolve into autocatalytic sets of
self-replicating, metabolically active entities that would predate the life forms known today.
The experiment produced a relatively small yield of dipeptides (0.4% to 12.4%) and a smaller yield of
tripeptides (0.10%) but the authors also noted that: "under these same conditions dipeptides hydrolysed
rapidly."[43]

[edit] Radioactive beach hypothesis

Zachary Adam at the University of Washington, Seattle, claims that stronger tidal processes from a
much closer moon may have concentrated grains of uranium and other radioactive elements at the high
water mark on primordial beaches where they may have been responsible for generating life's building
blocks.[44] According to computer models reported in Astrobiology,[45] a deposit of such radioactive
materials could show the same self-sustaining nuclear reaction as that found in the Oklo uranium ore seam in
Gabon. Such radioactive beach sand provides sufficient energy to generate organic molecules, such as
amino acids and sugars from acetonitrile in water. Radioactive monazite also releases soluble phosphate
into regions between sand-grains, making it biologically "accessible". Thus amino acids, sugars and soluble
phosphates can all be simultaneously produced, according to Adam. Radioactive actinides, then in greater
concentrations, could have formed part of organo-metallic complexes. These complexes could have been
important early catalysts to living processes.
John Parnell of the University of Aberdeen suggests that such a process could provide part of the
"crucible of life" on any early wet rocky planet, so long as the planet is large enough to have generated a
system of plate tectonics which brings radioactive minerals to the surface. As the early Earth is believed to
have had many smaller "platelets" it would provide a suitable environment for such processes.[46]

[edit] Thermodynamic Origin of Life: Ultraviolet and Temperature-Assisted

Replication (UVTAR) Model
Karo Michaelian of the National Autonomous University of Mexico (UNAM) points out that any model
for the origin of life must take into account the fact that life is an irreversible thermodynamic process which
arises and persists to produce entropy. Entropy production is not incidental to the process of life, but rather
the fundamental reason for its existence. Present day life augments the entropy production of Earth by
catalysing the water cycle through evapotranspiration.[47] Michaelian argues that if the thermodynamic
function of life today is to produce entropy through coupling with the water cycle, then this probably was its
function at its very beginnings. It turns out that both RNA and DNA when in water solution are very strong
absorbers and extremely rapid dissipaters of ultraviolet light within the 200 nm - 300 nm wavelength range,
just that high energy part of the sun's spectrum that could have penetrated the dense prebiotic atmosphere.
Cnossen et al.[48] have shown that the amount of UV light reaching the Earth's surface in the Archean could
have been up to 31 orders of magnitude larger than it is today at 260 nm where RNA and DNA absorb most
strongly. Absorption and dissipation of UV light by these organic molecules at the Archean ocean surface
would have increased significantly the temperature of the surface skin layer leading to enhanced evaporation
and thus augmenting the primitive water cycle. Since absorption and dissipation of high energy photons is an
entropy producing process, Michaelian argues that non-equilbrium abiogenic synthesis of RNA and DNA
utilizing UV light [49] would have been thermodynamically favored.
A simple mechanism to explain the replication of RNA and DNA without the use of enzymes can also
be given within the same thermodynamic framework by assuming that life arose when the temperature of the
primitive seas had cooled to somewhat below the denaturing temperature of RNA or DNA (based on the ratio
of 18O/16O found in cherts of the Barberton greenstone belt of South Africa of about 3.5 to 3.2 Ga., surface
temperatures are predicted to have been around 70±15 °C [50], similar to RNA or DNA denaturing
temperatures). During the night, the surface water temperature would be below the denaturing temperature
and single strand RNA/DNA could act as a template for the formation of double strand RNA/DNA. During the
daylight hours, RNA and DNA would absorb UV light and convert this directly to heating of the ocean surface,
raising the local temperature enough to allow for denaturing of RNA and DNA. The copying process would be
repeated during the cool period overnight [51] . Such a temperature assisted mechanism of replication bears
similarity to Polymerase Chain Reaction (PCR), a routine laboratory procedure to multiply DNA segments.
Michaelian suggests that traditional origin of life research, expecting to describe the emergence of life from
near-equilibrium conditions, is erroneous and that non-equilibrium conditions must be considered, in
particular, the importance of entropy production to the emergence of life.
Since denaturation would be most probable in the late afternoon when the Archean sea surface
temperature would be highest, and since late afternoon sunlight is somewhat circularly polarized, the
homochirality of the organic molecules of life can also be explained within the proposed thermodynamic
framework.[52]
 [edit] Models to explain homochirality
Main article: Homochirality
Some process in chemical evolution must account for the origin of homochirality, i.e. all building
blocks in living organisms having the same "handedness" (amino acids being left-handed, nucleic acid
sugars (ribose and deoxyribose) being right-handed, and chiral phosphoglycerides). Chiral molecules can be
synthesized, but in the absence of a chiral source or a chiral catalyst, they are formed in a 50/50 mixture of
both enantiomers. This is called a racemic mixture. Clark has suggested that homochirality may have started
in space, as the studies of the amino acids on the Murchison meteorite showed L-alanine to be more than
twice as frequent as its D form, and L-glutamic acid was more than 3 times prevalent than its D counterpart. It
is suggested that polarised light has the power to destroy one enantiomer within the proto-planetary disk.
Noyes[53] showed that beta decay caused the breakdown of D-leucine, in a racemic mixture, and that the
presence of 14C, present in larger amounts in organic chemicals in the early Earth environment, could have
been the cause. Robert M. Hazen reports upon experiments conducted in which various chiral crystal
surfaces act as sites for possible concentration and assembly of chiral monomer units into macromolecules.
[54] Once established, chirality would be selected for.[55] Work with organic compounds found on meteorites
tends to suggest that chirality is a characteristic of abiogenic synthesis, as amino acids show a left-handed
bias, whereas sugars show a predominantly right-handed bias.[56]

[edit] Self-organization and replication

Main article: Self-organization
While features of self-organization and self-replication are often considered the hallmark of living
systems, there are many instances of abiotic molecules exhibiting such characteristics under proper
conditions. For example Martin and Russel show that physical compartmentation by cell membranes from the
environment and self-organization of self-contained redox reactions are the most conserved attributes of
living things, and they argue therefore that inorganic matter with such attributes would be life's most likely last
common ancestor.[57]
Virus self-assembly within host cells has implications for the study of the origin of life,[58] as it lends
further credence to the hypothesis that life could have started as self-assembling organic molecules.[59] [60]

[edit] From organic molecules to protocells

The question "How do simple organic molecules form a protocell?" is largely unanswered but there
are many hypotheses. Some of these postulate the early appearance of nucleic acids ("genes-first") whereas
others postulate the evolution of biochemical reactions and pathways first ("metabolism-first"). Recently,
trends are emerging to create hybrid models that combine aspects of both.

[edit] "Genes first" models: the RNA world

Main article: RNA world hypothesis
The RNA world hypothesis describes an early Earth with self-replicating and catalytic RNA but no
DNA or proteins. This has spurred scientists to try to determine if relatively short RNA molecules could have
spontaneously formed that were capable of catalyzing their own continuing replication.[61] A number of
hypotheses of modes of formation have been put forward.[62] Early cell membranes could have formed
spontaneously from proteinoids, protein-like molecules that are produced when amino acid solutions are
heated–when present at the correct concentration in aqueous solution, these form microspheres which are
observed to behave similarly to membrane-enclosed compartments. Other possibilities include systems of
chemical reactions taking place within clay substrates or on the surface of pyrite rocks. Factors supportive of
an important role for RNA in early life include its ability to act both to store information and catalyse chemical
reactions (as a ribozyme); its many important roles as an intermediate in the expression and maintenance of
the genetic information (in the form of DNA) in modern organisms; and the ease of chemical synthesis of at
least the components of the molecule under conditions approximating the early Earth. Relatively short RNA
molecules which can duplicate others have been artificially produced in the lab.[63] Such replicase RNA,
which functions as both code and catalyst provides a template upon which copying can occur. Jack Szostak
has shown that certain catalytic RNAs can, indeed, join smaller RNA sequences together, creating the
potential, in the right conditions for self-replication. If these were present, Darwinian selection would favour
the proliferation of such self-catalysing structures, to which further functionalities could be added.[64] Lincoln
and Joyce identified an RNA enzyme capable of self sustained replication.[65]
Researchers have pointed out difficulties for the abiotic synthesis of nucleotides from cytosine and
uracil.[66] Cytosine has a half-life of 19 days at 100 °C (212 °F) and 17,000 years in freezing water.[67]
Larralde et al., say that "the generally accepted prebiotic synthesis of ribose, the formose reaction, yields
numerous sugars without any selectivity."[68] and they conclude that their "results suggest that the backbone
of the first genetic material could not have contained ribose or other sugars because of their instability." The
ester linkage of ribose and phosphoric acid in RNA is known to be prone to hydrolysis.[69]
A slightly different version of the RNA-world hypothesis is that a different type of nucleic acid, such
as PNA, TNA or GNA, was the first one to emerge as a self-reproducing molecule, to be replaced by RNA
only later.[70][71] Pyrimidine ribonucleosides and their respective nucleotides have been prebiotically
synthesised by a sequence of reactions which by-pass the free sugars, and are assembled in a stepwise
fashion by going against the dogma that nitrogenous and oxygenous chemistries should be avoided. In a
series of publications, The Sutherland Group at the School of Chemistry, University of Manchester have
demonstrated high yielding routes to cytidine and uridine ribonucleotides built from small 2 and 3 carbon
fragments such as glycolaldehyde, glyceraldehyde or glyceraldehyde-3-phosphate, cyanamide and
cyanoacetylene. One of the steps in this sequence allows the isolation of enantiopure ribose aminooxazoline
if the enantiomeric excess of glyceraldehyde is 60 % or greater.[72] This can be viewed as a prebiotic
purification step, where the said compound spontaneously crystallised out from a mixture of the other
pentose aminooxazolines. Ribose aminooxazoline can then react with cyanoacetylene in a mild and highly
efficient manner to give the alpha cytidine ribonucleotide. Photoanomerization with UV light allows for
inversion about the 1' anomeric centre to give the correct beta stereochemistry.[73] In 2009 they showed that
the same simple building blocks allow access, via phosphate controlled nucleobase elaboration, to 2',3'-
cyclic pyrimidine nucleotides directly, which are known to be able to polymerise into RNA. This paper also
highlights the possibility for the photo-sanitization of the pyrimidine-2',3'-cyclic phosphates.[49] James
Ferris's studies have shown that clay minerals of montmorillonite will catalyze the formation of RNA in
aqueous solution, by joining activated mono RNA nucleotides to join together to form longer chains.[74]
Although these chains have random sequences, the possibility that one sequence began to non-randomly
increase its frequency by increasing the speed of its catalysis is possible to "kick start" biochemical evolution.

[edit] "Metabolism first" models

Several models reject the idea of the self-replication of a "naked-gene" and postulate the emergence
of a primitive metabolism which could provide an environment for the later emergence of RNA replication.

[edit] Iron-sulfur world

One of the earliest incarnations of this idea was put forward in 1924 with Alexander Oparin's notion of
primitive self-replicating vesicles which predated the discovery of the structure of DNA. More recent variants
in the 1980s and 1990s include Günter Wächtershäuser's iron-sulfur world theory and models introduced by
Christian de Duve based on the chemistry of thioesters. More abstract and theoretical arguments for the
plausibility of the emergence of metabolism without the presence of genes include a mathematical model
introduced by Freeman Dyson in the early 1980s and Stuart Kauffman's notion of collectively autocatalytic
sets, discussed later in that decade.
However, the idea that a closed metabolic cycle, such as the reductive citric acid cycle, could form
spontaneously (proposed by Günter Wächtershäuser) remains debated. In an article entitled "Self-Organizing
Biochemical Cycles",[75] the late Leslie Orgel summarized his analysis of the proposal by stating, "There is
at present no reason to expect that multistep cycles such as the reductive citric acid cycle will self-organize
on the surface of FeS/FeS2 or some other mineral." It is possible that another type of metabolic pathway was
used at the beginning of life. For example, instead of the reductive citric acid cycle, the "open" acetyl-CoA
pathway (another one of the five recognised ways of carbon dioxide fixation in nature today) would be
compatible with the idea of self-organisation on a metal sulfide surface. The key enzyme of this pathway,
carbon monoxide dehydrogenase/acetyl-CoA synthase harbours mixed nickel-iron-sulfur clusters in its
reaction centers and catalyses the formation of acetyl-CoA (which may be regarded as a modern form of
acetyl-thiol) in a single step.

[edit] Thermosynthesis world

Today’s bioenergetic process of fermentation is related to the just mentioned citric acid cycle or the
Acetyl-CoA pathway that have been connected to the primordial iron-sulfur world. In a different approach,
today’s bioenergetic process of chemiosmosis, which plays an essential role in cellular respiration and
photosynthesis, is considered as more fundamental than fermentation: in Anthonie Muller’s “thermosynthesis
world” the ATP Synthase enzyme that sustains chemiosmosis is proposed as today’s enzyme that is the
closest connected to the first metabolic process.[76][77]
First life needed an energy source to bring about the condensation reaction that yielded the peptide
bonds of proteins and the phosphodiester bonds of RNA. In a generalization and thermal variation of the
binding change mechanism of today’s ATP Synthase, the “First Protein” would have bound substrates
(peptides, phosphate, nucleosides, RNA ‘monomers’) and condensed them to a reaction product that
remained bound until it after a temperature change was released upon a thermal unfolding.
The energy source of the thermosynthesis world was thermal cycling, the result of suspension of the
protocell in a convection current, as is plausible in a volcanic hot spring; the convection accounts for the self-
organization and dissipative structure required in any origin of life model. The still ubiquitous role of thermal
cycling in germination and cell division is considered a relic of primordial thermosynthesis.
By phosphorylating cell membrane lipids, this ‘First Protein’ gave a selective advantage to the lipid
protocell that contained the protein. In the beginning this First Protein also synthesized a library with many
proteins, of which only a minute fraction had thermosynthesis capabilities. Just as proposed by Dyson [78] for
the first proteins, the First Protein propagated functionally: it made daughters with similar capabilities, but it
did not copy itself. Functioning daughters consisted of different amino acid sequences.
Over a long time, RNA sequences were selected among the at first randomly synthesized RNAs by
the criterion of speed and efficiency increase of First Protein synthesis, for instance by the creation of RNA
that functioned as messenger RNA[79], Transfer RNA[80] and ribosomal RNA, or, even more generally, all
the components of the RNA World were also generated and selected. The thermosynthesis world therefore in
theory accounts for the origin of the genetic machinery.
 Whereas the iron-sulfur world identifies a circular pathway as the most simple—and therefore
assumes the existence of enzymes—the thermosynthesis world does not even invoke a pathway, and does
not assume the existence of regular enzymes: ATP Synthase’s binding change mechanism resembles a
physical adsorption process that yields free energy[81], rather than a regular enzyme’s mechanism, which
decreases the free energy. The RNA World also implies the existence of several enzymes. But even the
emergence of a single enzyme by chance is implausible.[82] The thermosynthesis world is therefore more
simple, and thus more plausible, than the iron-sulfur and RNA worlds.

[edit] Possible role of bubbles

Waves breaking on the shore create a delicate foam composed of bubbles. Winds sweeping across
the ocean have a tendency to drive things to shore, much like driftwood collecting on the beach. It is possible
that organic molecules were concentrated on the shorelines in much the same way. Shallow coastal waters
also tend to be warmer, further concentrating the molecules through evaporation. While bubbles composed
mostly of water burst quickly, water containing amphiphiles forms much more stable bubbles, lending more
time to the particular bubble to perform these crucial reactions.
Amphiphiles are oily compounds containing a hydrophilic head on one or both ends of a hydrophobic
molecule. Some amphiphiles have the tendency to spontaneously form membranes in water. A spherically
closed membrane contains water and is a hypothetical precursor to the modern cell membrane. If a protein
would increase the integrity of its parent bubble, that bubble had an advantage, and was placed at the top of
the natural selection waiting list. Primitive reproduction can be envisioned when the bubbles burst, releasing
the results of the 'experiment' into the surrounding medium. Once enough of the 'right stuff' was released into
the medium, the development of the first prokaryotes, eukaryotes, and multicellular organisms could be
achieved.[83]
 Similarly, bubbles formed entirely out of protein-like molecules, called microspheres, will form
spontaneously under the right conditions. But they are not a likely precursor to the modern cell membrane, as
cell membranes are composed primarily of lipid compounds rather than amino-acid compounds (for types of
membrane spheres associated with abiogenesis, see protobionts, micelle, coacervate).
A recent model by Fernando and Rowe[84] suggests that the enclosure of an autocatalytic non-
enzymatic metabolism within protocells may have been one way of avoiding the side-reaction problem that is
typical of metabolism first models.

[edit] Other models

[edit] Autocatalysis
In 1993 Stuart Kauffman proposed that life initially arose as autocatalytic chemical networks.[85]
British ethologist Richard Dawkins wrote about autocatalysis as a potential explanation for the origin
of life in his 2004 book The Ancestor's Tale. Autocatalysts are substances which catalyze the production of
themselves, and therefore have the property of being a simple molecular replicator. In his book, Dawkins
cites experiments performed by Julius Rebek and his colleagues at the Scripps Research Institute in
California in which they combined amino adenosine and pentafluorophenyl ester with the autocatalyst amino
adenosine triacid ester (AATE). One system from the experiment contained variants of AATE which
catalysed the synthesis of themselves. This experiment demonstrated the possibility that autocatalysts could
exhibit competition within a population of entities with heredity, which could be interpreted as a rudimentary
form of natural selection.
[edit] Clay theory
A model for the origin of life based on clay was forwarded by A. Graham Cairns-Smith of the
University of Glasgow in 1985 and explored as a plausible illustration by several other scientists, including
Richard Dawkins[86]. Clay theory postulates that complex organic molecules arose gradually on a pre-
existing, non-organic replication platform—silicate crystals in solution. Complexity in companion molecules
developed as a function of selection pressures on types of clay crystal is then exapted to serve the replication
of organic molecules independently of their silicate "launch stage".
Cairns-Smith is a staunch critic of other models of chemical evolution.[87] However, he admits that
like many models of the origin of life, his own also has its shortcomings (Horgan 1991).
In 2007, Kahr and colleagues reported their experiments to examine the idea that crystals can act as
a source of transferable information, using crystals of potassium hydrogen phthalate. "Mother" crystals with
imperfections were cleaved and used as seeds to grow "daughter" crystals from solution. They then
examined the distribution of imperfections in the crystal system and found that the imperfections in the
mother crystals were indeed reproduced in the daughters. The daughter crystals had many additional
imperfections. For a gene-like behavior the additional imperfections should be much less than the parent
ones, thus Kahr concludes that the crystals "were not faithful enough to store and transfer information from
one generation to the next".[88][89]

[edit] Gold's "Deep-hot biosphere" model

In the 1970s, Thomas Gold proposed the theory that life first developed not on the surface of the
Earth, but several kilometers below the surface. The discovery in the late 1990s of nanobes (filamental
structures that are smaller than bacteria, but that may contain DNA) in deep rocks [90] might be seen as
lending support to Gold's theory.
It is now reasonably well established that microbial life is plentiful at shallow depths in the Earth, up
to 5 kilometres (3.1 mi) below the surface,[90] in the form of extremophile archaea, rather than the better-
known eubacteria (which live in more accessible conditions). It is claimed that discovery of microbial life
below the surface of another body in our solar system would lend significant credence to this theory. Thomas
Gold also asserted that a trickle of food from a deep, unreachable, source is needed for survival because life
arising in a puddle of organic material is likely to consume all of its food and become extinct. Gold's theory is
that flow of food is due to out-gassing of primordial methane from the Earth's mantle; more conventional
explanations of the food supply of deep microbes (away from sedimentary carbon compounds) is that the
organisms subsist on hydrogen released by an interaction between water and (reduced) iron compounds in
rocks.

[edit] "Primitive" extraterrestrial life

An alternative to Earthly abiogenesis is the hypothesis that primitive life may have originally formed
extraterrestrially, either in space or on a nearby planet (Mars). (Note that exogenesis is related to, but not the
same as, the notion of panspermia). A supporter of this theory was Francis Crick.
Organic compounds are relatively common in space, especially in the outer solar system where
volatiles are not evaporated by solar heating.[91] Comets are encrusted by outer layers of dark material,
thought to be a tar-like substance composed of complex organic material formed from simple carbon
compounds after reactions initiated mostly by irradiation by ultraviolet light. It is supposed that a rain of
material from comets could have brought significant quantities of such complex organic molecules to Earth.
 An alternative but related hypothesis, proposed to explain the presence of life on Earth so soon after
the planet had cooled down, with apparently very little time for prebiotic evolution, is that life formed first on
early Mars. Due to its smaller size Mars cooled before Earth (a difference of hundreds of millions of years),
allowing prebiotic processes there while Earth was still too hot. Life was then transported to the cooled Earth
when crustal material was blasted off Mars by asteroid and comet impacts. Mars continued to cool faster and
eventually became hostile to the continued evolution or even existence of life (it lost its atmosphere due to
low volcanism); Earth is following the same fate as Mars, but at a slower rate.
Neither hypothesis actually answers the question of how life first originated, but merely shifts it to
another planet or a comet. However, the advantage of an extraterrestrial origin of primitive life is that life is
not required to have evolved on each planet it occurs on, but rather in a single location, and then spread
about the galaxy to other star systems via cometary and/or meteorite impact. Evidence to support the
plausibility of the concept is scant, but it finds support in recent study of Martian meteorites found in
Antarctica and in studies of extremophile microbes.[92] Additional support comes from a recent discovery of
a bacterial ecosytem whose energy source is radioactivity.[93]
A 2001 experiment led by Jason Dworkin[94] subjected a frozen mixture of water, methanol,
ammonia and carbon monoxide to UV radiation, mimicking conditions found in an extraterrestrial
environment. This combination yielded large amounts of organic material that self-organised to form bubbles
or micelles when immersed in water. Dworkin considered these bubbles to resemble cell membranes that
enclose and concentrate the chemistry of life, separating their interior from the outside world.
The bubbles produced in these experiments were between 10 to 40 micrometres (0.00039 to 0.0016
in), or about the size of red blood cells. Remarkably, the bubbles fluoresced, or glowed, when exposed to UV
light. Absorbing UV and converting it into visible light in this way was considered one possible way of
providing energy to a primitive cell. If such bubbles played a role in the origin of life, the fluorescence could
have been a precursor to primitive photosynthesis. Such fluorescence also provides the benefit of acting as a
sunscreen, diffusing any damage that otherwise would be inflicted by UV radiation. Such a protective function
would have been vital for life on the early Earth, since the ozone layer, which blocks out the sun's most
destructive UV rays, did not form until after photosynthetic life began to produce oxygen.[56]

[edit] Extraterrestrial amino acids

Another idea is that amino acids which were formed extra-terrestrially arrived on Earth via comets. In
2009 it was announced by NASA that scientists have identified one of the fundamental chemical buildings
blocks of life in a comet for the first time: glycine, an amino acid, was detected in the material ejected from
Comet Wild-2 in 2004 and grabbed by NASA's Stardust probe. Tiny grains, just a few thousandths of a
millimetre in size, were collected from the comet and returned to Earth in 2006 in a sealed capsule, and
distributed among the world's leading astro-biology labs. NASA said in a statement that it took some time for
the investigating team, led by Dr Jamie Elsila, to convince itself that the glycine signature found in Stardust's
sample bay was genuine and not just Earthly contamination. Glycine has been detected in meteorites before
and there are also observations in interstellar gas clouds claimed for telescopes, but the Stardust find is
described as a first in cometary material. It is known that prior to the emergence of life on Earth, the early
solar system's planets were regularly bombarded by comets. Dr. Carl Pilcher, who leads NASA's
Astrobiology Institute commented that "The discovery of glycine in a comet supports the idea that the
fundamental building blocks of life are prevalent in space, and strengthens the argument that life in the
Universe may be common rather than rare."[95]
[edit] Lipid World
This theory postulates that the first self-replicating object was lipid-like.[96] It is known that
phospholipids form bilayers in water while under agitation– the same structure as in cell membranes. These
molecules were not present on early Earth, however other amphiphilic long chain molecules also form
membranes. Furthermore, these bodies may expand (by insertion of additional lipids), and under excessive
expansion may undergo spontaneous splitting which preserves the same size and composition of lipids in the
two progenies. The main idea in this theory is that the molecular composition of the lipid bodies is the
preliminary way for information storage, and evolution led to the appearance of polymer entities such as RNA
or DNA that may store information favorably. Still, no biochemical mechanism has been offered to support
the Lipid World theory.

[edit] Polyphosphates
The problem with most scenarios of abiogenesis is that the thermodynamic equilibrium of amino acid
versus peptides is in the direction of separate amino acids. What has been missing is some force that drives
polymerization. The resolution of this problem may well be in the properties of polyphosphates.[97][98]
Polyphosphates are formed by polymerization of ordinary monophosphate ions PO 4−3. Several mechanisms
for such polymerization have been suggested. Polyphosphates cause polymerization of amino acids into
peptides[citation needed]. They are also logical precursors in the synthesis of such key biochemical
compounds as ATP. A key issue seems to be that calcium reacts with soluble phosphate to form insoluble
calcium phosphate (apatite), so some plausible mechanism must be found to keep calcium ions from causing
precipitation of phosphate. There has been much work on this topic over the years, but an interesting new
idea is that meteorites may have introduced reactive phosphorus species on the early Earth.[99]
[edit] PAH world hypothesis
Main article: PAH world hypothesis
Other sources of complex molecules have been postulated, including extraterrestrial stellar or
interstellar origin. For example, from spectral analyses, organic molecules are known to be present in comets
and meteorites. In 2004, a team detected traces of polycyclic aromatic hydrocarbons (PAH's) in a nebula.
[100] Those are the most complex molecules so far found in space. The use of PAH's has also been
proposed as a precursor to the RNA world in the PAH world hypothesis.[101] The Spitzer Space Telescope
has recently detected a star, HH 46-IR, which is forming by a process similar to that by which the sun formed.
In the disk of material surrounding the star, there is a very large range of molecules, including cyanide
compounds, hydrocarbons, and carbon monoxide. PAHs have also been found all over the surface of galaxy
M81, which is 12 million light years away from the Earth, confirming their widespread distribution in space.
[102]

[edit] Multiple genesis

Different forms of life may have appeared quasi-simultaneously in the early history of Earth.[103] The
other forms may be extinct, leaving distinctive fossils through their different biochemistry (e.g., using arsenic
instead of phosphorus), survive as extremophiles, or simply be unnoticed through their being analogous to
organisms of the current life tree. Hartman[104] for example combines a number of theories together, by
proposing that:
The first organisms were self-replicating iron-rich clays which fixed carbon dioxide into oxalic
and other dicarboxylic acids. This system of replicating clays and their metabolic phenotype
then evolved into the sulfide rich region of the hotspring acquiring the ability to fix nitrogen.
 Finally phosphate was incorporated into the evolving system which allowed the synthesis of
nucleotides and phospholipids. If biosynthesis recapitulates biopoesis, then the synthesis of
amino acids preceded the synthesis of the purine and pyrimidine bases. Furthermore the
polymerization of the amino acid thioesters into polypeptides preceded the directed
polymerization of amino acid esters by polynucleotides.

Lynn Margulis's endosymbiotic theory suggests that multiple forms of bacteria entered into symbiotic
relationship to form the eukaryotic cell. The horizontal transfer of genetic material between bacteria promotes
such symbiotic relationships, and thus many separate organisms may have contributed to building what has
been recognised as the Last Universal Common Ancestor (LUCA) of modern organisms. James Lovelock's
Gaia theory, proposes that such bacterial symbiosis establishes the environment as a system produced by
and supportive of life. His arguments strongly weaken the case for life having evolved elsewhere in the solar
system.

[edit] See also

• Astrochemistry
• Autocatalytic reactions and order creation
• Biogenesis
• Common descent
• Drake equation
• Entropy and life
• History of Earth
• List of independent discoveries
 • List of publications in biology
• Mediocrity principle
• Origin of the world's oceans
• Mimivirus
• Planetary habitability
• Rare Earth hypothesis
• Shadow biosphere
• Thermosynthesis
• Zeolite
• John Needham

[edit] References
1. ^ Miller-Urey Experiment: Amino Acids & The Origins of Life on Earth
2. ^ Zimmer C (August 2009). "Origins. On the origin of eukaryotes". Science 325 (5941): 666–
8. doi:10.1126/science.325_666. PMID 19661396.
3. ^ a b Wilde SA, Valley JW, Peck WH, Graham CM (January 2001). "Evidence from detrital
zircons for the existence of continental crust and oceans on the Earth 4.4 Gyr ago". Nature 409
(6817): 175–8. doi:10.1038/35051550. PMID 11196637.
4. ^ Schopf JW, Kudryavtsev AB, Agresti DG, Wdowiak TJ, Czaja AD (March 2002). "Laser--
Raman imagery of Earth's earliest fossils". Nature 416 (6876): 73–6. doi:10.1038/416073a.
PMID 11882894.
 5. ^ Hayes, John M.; Waldbauer, Jacob R. (2006). "The carbon cycle and associated redox
processes through time". Phil. Trans. R. Soc. B 361 (1470): 931–950. doi:10.1098/rstb.2006.1840.
PMID 16754608.
6. ^ Archer, Corey; Vance, Derek (2006). "Coupled Fe and S isotope evidence for Archean
microbial Fe(III) and sulfate reduction". Geology 34 (3): 153–156. doi:10.1130/G22067.1.
7. ^ Cavalier-Smith, Thomas; Brasier, Martin; Embley, T. Martin (2006). "Introduction: how and
when did microbes change the world?". Phil. Trans. R. Soc. B 361 (1470): 845–50.
doi:10.1098/rstb.2006.1847. PMID 16754602.
8. ^ Summons, Roger E.; et al. (2006). "Steroids, triterpenoids and molecular oxygen". Phil.
Trans. R. Soc. B 361 (1470): 951–68. doi:10.1098/rstb.2006.1837. PMID 16754609.
9. ^ Philip P. Wiener, ed (1973). "Spontaneous Generation". Dictionary of the History of Ideas.
New York: Charles Scribner's Sons. http://xtf.lib.virginia.edu/xtf/view?
docId=DicHist/uvaBook/tei/DicHist1.xml;chunk.id=dv4-39. Retrieved 2009-01-24.
10.^ Lennox, James (2001). Aristotle's Philosophy of Biology: Studies in the Origins of Life
Science. New York, NY: Cambridge Press. pp. 229–258. ISBN 978-0521659765.
11.^ Balme, D. M. (1962). "Development of Biology in Aristotle and Theophrastus: Theory of
Spontaneous Generation". Phronesis: a journal for Ancient Philosophy 7 (1–2): 91–104.
doi:10.1163/156852862X00052.
12.^ Dobell, C. (1960). Antony Van Leeuwenhoek and his little animals. New York: Dover
Publications. ISBN 0486605949.
13.^ Oparin, Aleksandr I. (1953). Origin of Life. Dover Publications, New York. pp. 196.
ISBN 0486602133.
14.^ First life on Earth windmillministries.org, Retrieved on 2008-01-18
 15.^ "It is often said that all the conditions for the first production of a living organism are now
present, which could ever have been present. But if (and oh! what a big if!) we could conceive in
some warm little pond, with all sorts of ammonia and phosphoric salts, light, heat, electricity, &c.,
present, that a proteine compound was chemically formed ready to undergo still more complex
changes, at the present day such matter would be instantly devoured or absorbed, which would not
have been the case before living creatures were formed." written in 1871, published in Darwin,
Francis, ed. 1887. The life and letters of Charles Darwin, including an autobiographical chapter.
London: John Murray. Volume 3. p. 18
16.^ Oparin, A. I. (1924) Proiskhozhozhdenie zhizny, Moscow (Translated by Ann Synge in
Bernal (1967), The Origin of Life, Weidenfeld and Nicolson, London, pages 199–234.
17.^ Oparin, A. I. (1952). The Origin of Life. New York: Dover. ISBN 0486495221.
18.^ Bernal, J.D. (1969). Origins of Life. London: Wiedenfeld and Nicholson.
19.^ Bryson, Bill (2004). A short history of nearly everything. London: Black Swan. pp. 300–2.
ISBN 0-552-99704-8.
20.^ Morse, J. W.; MacKenzie, F. T. (1998). "Hadean Ocean Carbonate chemistry". Aquatic
Geochemistry 4: 301–19. doi:10.1023/A:1009632230875.
21.^ Sleep, Norman H.; et al. (1989). "Annihilation of ecosystems by large asteroid impacts on
early Earth". Nature 342 (6246): 139–142. doi:10.1038/342139a0. PMID 11536616.
22.^ Maher, Kevin A.; Stevenson, David J. (1988). "Impact frustration of the origin of life".
Nature 331 (6157): 612–4. doi:10.1038/331612a0. PMID 11536595.
23.^ Orgel, Leslie E. (2004). "Prebiotic adenine revisited: Eutectics and photochemistry". Origins
of Life and Evolution of Biospheres 34: 361–9. doi:10.1023/B:ORIG.0000029882.52156.c2.
24.^ Robertson, Michael P.; Miller, Stanley L. (1995). "An efficient prebiotic synthesis of cytosine
and uracil". Nature 375 (6534): 772–774. doi:10.1038/375772a0. PMID 7596408.
 25.^ Bada, J. L.; Bigham, C.; Miller, S. L. (1994). "Impact Melting of Frozen Oceans on the Early
Earth: Implications for the Origin of Life" (abstract). Proc. Natl. Acad. Sci. U.S.A. 91 (4): 1248–50.
doi:10.1073/pnas.91.4.1248. PMID 11539550. PMC 43134.
http://www.pnas.org/cgi/content/abstract/91/4/1248.
26.^ "Did Life Evolve in Ice?". DISCOVER Magazine. http://discovermagazine.com/2008/feb/did-
life-evolve-in-ice/article_view?b_start:int=0&-C=. Retrieved 2008-07-03.
27.^ Levy, M.; Miller, S. L.; Brinton, K.; Bada, J. L. (June 2000). "Prebiotic synthesis of adenine
and amino acids under Europa-like conditions". Icarus 145 (2): 609–13. doi:10.1006/icar.2000.6365.
PMID 11543508.
28.^ Trinks, Hauke; Schröder, Wolfgang; Biebricher, Christof (October 2005). "Ice And The
Origin Of Life". Origins of Life and Evolution of the Biosphere 35 (5): 429–45. doi:10.1007/s11084-
005-5009-1. PMID 16231207.
http://www.ingentaconnect.com/content/klu/orig/2005/00000035/00000005/00005009#aff_1.
Retrieved 2008-02-11.
29.^ Mojzis, S. J.; et al. (1996). "Evidence for life on earth before 3,800 million years ago".
Nature 384 (6604): 55–9. doi:10.1038/384055a0. PMID 8900275.
30.^ Lazcano, A.; Miller, S. L. (1994). "How long did it take for life to begin and evolve to
cyanobacteria?". Journal of Molecular Evolution 39 (6): 546–54. doi:10.1007/BF00160399.
PMID 11536653.
31.^ Bernal, John Desmond (1949). "The Physical Basis of Life". Proceedings of the Physical
Society. Section A, 1949 62: 537–538. doi:10.1088/0370-1298/62/9/301.
32.^ Chyba, Christopher; Sagan, Carl (1992). "Endogenous production, exogenous delivery and
impact-shock synthesis of organic molecules: an inventory for the origins of life". Nature 355 (6356):
125–32. doi:10.1038/355125a0. PMID 11538392.
 33.^ Shapiro, Robert (1987). Origins: A Skeptic's Guide to the Creation of Life on Earth . Bantam
Books. p. 110. ISBN 0671459392.
34.^ Miller, Stanley L. (1953). "A Production of Amino Acids Under Possible Primitive Earth
Conditions". Science 117 (3046): 528–9. doi:10.1126/science.117.3046.528. PMID 13056598.
35.^ Oró, J. (1961). "Mechanism of synthesis of adenine from hydrogen cyanide under possible
primitive Earth conditions". Nature 191: 1193–4. doi:10.1038/1911193a0. PMID 13731264.
36.^ Menor-Salván C, Ruiz-Bermejo DM, Guzmán MI, Osuna-Esteban S, Veintemillas-
Verdaguer S (2007). "Synthesis of pyrimidines and triazines in ice: implications for the prebiotic
chemistry of nucleobases.". Chemistry 15 (17) (17): 4411–8. doi:10.1002/chem.200802656.
PMID 19288488.
37.^ Experiments on origin of organic molecules Nitro.biosci.arizona.edu, Retrieved on 2008-01-
13
38.^ Schuster, P.; Eigen, M. (1979). The hypercycle, a principle of natural self-organization .
Berlin: Springer-Verlag. ISBN 0-387-09293-5.
39.^ origin of life thebioreview.com Retrieved on 2008-01-14
40.^ Hoffmann, G. W. (1974). "On the Origin of the Genetic Code and the Stability of the
Translation Apparatus". J. Mol. Biol. 86: pp. 349–362.
41.^ Orgel, L. (1963). "The Maintenance of the Accuracy of Protein Synthesis and its Relevance
to Ageing". Proc. Nat. Acad. Sci. USA 49: pp. 517–521.
42.^ Hoffmann, G. W. (1975). "The Stochastic Theory of the Origin of Life". Annual Review of
Physical Chemistry 26: pp. 123–144.
43.^ Huber, C.; Wächterhäuser, G. (1998). "Peptides by activation of amino acids with CO on
(Ni,Fe)S surfaces: implications for the origin of life". Science 281 (5377): 670–2.
doi:10.1126/science.281.5377.670. PMID 9685253.
 44.^ Dartnell, Lewis (2008-01-12). "Life's a beach on planet Earth". New Scientist.
45.^ Adam, Zachary (2007). "Actinides and Life's Origins". Astrobiology 7 (6): 852–72.
doi:10.1089/ast.2006.0066. PMID 18163867.
46.^ Parnell, John (2004). "Mineral Radioactivity in Sands as a Mechanism for Fixation of
Organic Carbon on the Early Earth" (PDF). Origins of Life and Evolution of Biospheres 34 (6): 533–
547. doi:10.1023/B:ORIG.0000043132.23966.a1.
http://www.springerlink.com/content/mp42778372jv6054/fulltext.pdf.
47.^ Michaelian, Karo (2009). "Thermodynamic Function of Lfe" (PDF). ArXiv.
http://arxiv.org/abs/0907.0040.
48.^ Cnossen et al., I.; Sanz-Forcada, Jorge; Favata, Fabio; Witasse, Olivier; Zegers, Tanja;
Arnold, Neil F. (2007). "The habitat of early life: Solar X-ray and UV radiation at Earth’s surface 4-3.5
billion years ago" (PDF). J. Geophys. Research 112: E02008. doi:10.1029/2006JE002784.
49.^ a b Powner MW, Gerland B, Sutherland JD (May 2009). "Synthesis of activated pyrimidine
ribonucleotides in prebiotically plausible conditions". Nature 459 (7244): 239–42.
doi:10.1038/nature08013. PMID 19444213.
50.^ Lowe, D. R. and Tice, M. M, Donald R.; Tice, Michael M. (2004). "Geologic evidence for
Archean atmospheric and climatic evolution: Fluctuating levels of CO2, CH4, and O2 with an
overriding tectonic control" (PDF). Geology 32: 493–496. doi:10.1130/G20342.1.
51.^ Michaelian, Karo (2010). "Thermodynamic Origin of Lfe" (PDF). Earth Syst. Dynam.
Discuss. 1: 1–39. doi:10.5194/esdd-1-1-2010. http://www.earth-syst-dynam-
discuss.net/1/1/2010/esdd-1-1-2010.html.
52.^ Michaelian, Karo (2009). "Thermodynamic Origin of Lfe" (PDF). ArXiv.
http://arxiv.org/abs/0907.0042.
 53.^ Noyes HP, Bonner WA, Tomlin JA (April 1977). "On the origin of biological chirality via
natural beta-decay". Orig. Life 8 (1): 21–3. doi:10.1007/BF00930935. PMID 896189.
54.^ Hazen, Robert M. (2005). Genesis: the scientific quest for life's origin. Washington, D.C:
Joseph Henry Press. ISBN 0-309-09432-1.
55.^ Clark, S. (1999). "Polarised starlight and the handedness of Life". American Scientist 97:
336–43. doi:10.1511/1999.4.336.
56.^ a b Mullen L (September 5, 2005). "Building Life from Star-Stuff". Astrobiology Magazine.
http://www.astrobio.net/news/article1702.html.
57.^ Martin, William; Russel, Michael J. (2003). "On the origins of cells: a hypothesis for the
evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from
prokaryotes to nucleated cells". Phil. Trans. R. Soc. B 358 (1429): 59–85.
doi:10.1098/rstb.2002.1183. PMID 12594918.
58.^ Koonin EV, Senkevich TG, Dolja VV (2006). "The ancient Virus World and evolution of
cells". Biol. Direct 1: 29. doi:10.1186/1745-6150-1-29. PMID 16984643. PMC 1594570.
http://www.biology-direct.com/content/1//29. Retrieved 2008-10-20.
59.^ Vlassov AV, Kazakov SA, Johnston BH, Landweber LF (August 2005). "The RNA world on
ice: a new scenario for the emergence of RNA information". J. Mol. Evol. 61 (2): 264–73.
doi:10.1007/s00239-004-0362-7. PMID 16044244.
60.^ Nussinov, Mark D.; Vladimir A. Otroshchenkob and Salvatore Santoli (1997). "Emerging
Concepts of Self-organization and the Living State". Biosystems 42 (2–3): 111–118.
doi:10.1016/S0303-2647(96)01699-1. PMID 9184757. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6T2K-3W0FTTV-
G&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&_docanchor=&view=c&_acct=C000050221&_
version=1&_urlVersion=0&_userid=10&md5=2ee9045613a39f99b996e96f342023d3. Retrieved
2009-08-02.
61.^ Ma W, Yu C, Zhang W, Hu J (November 2007). "Nucleotide synthetase ribozymes may
have emerged first in the RNA world". RNA (New York, N.Y.) 13 (11): 2012–9.
doi:10.1261/rna.658507. PMID 17878321.
62.^ Villarreal LP, Witzany G (February 2010). "Viruses are essential agents within the roots and
stem of the tree of life". J Theor Biol 262 (4): 698–710. doi:10.1016/j.jtbi.2009.10.014.
PMID 19833132.
63.^ Johnston, W. K.; et al. (2001). "RNA-Catalyzed RNA Polymerization: Accurate and General
RNA-Templated Primer Extension". Science 292 (5520): 1319–1325. doi:10.1126/science.1060786.
PMID 11358999.
64.^ Szostak, Jack W. (June 4, 2008). "The Origins of Function in Biological Nucleic Acids,
Proteins, and Membranes". HHMI. http://www.hhmi.org/research/investigators/szostak.html.
Retrieved 2008-11-29.
65.^ Lincoln, Tracey A.; Joyce, Gerald F. (January 8, 2009). "Self-Sustained Replication of an
RNA Enzyme". Science (New York: American Association for the Advancement of Science) 323
(5918): 1229–32. doi:10.1126/science.1167856. ISSN 1095-9203. PMID 19131595. PMC 2652413.
http://www.sciencemag.org/cgi/content/abstract/1167856. Retrieved 2009-01-13.
66.^ Orgel, L. (1994). "The origin of life on earth". Scientific American 271 (4): 81.
67.^ Levy, Matthew; Miller, Stanley L. (1998). "The stability of the RNA bases: Implications for
the origin of life". PNAS 95 (14): 7933–7938. doi:10.1073/pnas.95.14.7933. PMID 9653118.
68.^ Larralde, R.; Robertson, M. P.; Miller, S. L. (1995). "Rates of Decomposition of Ribose and
Other Sugars: Implications for Chemical Evolution". PNAS 92 (18): 8158–8160.
doi:10.1073/pnas.92.18.8158. PMID 7667262.
 69.^ Lindahl, Tomas (1993). "Instability and decay of the primary structure of DNA". Nature 362
(6422): 709–715. doi:10.1038/362709a0. PMID 8469282.
70.^ Orgel, Leslie (2000). "A Simpler Nucleic Acid". Science 290 (5495): 1306–1307.
doi:10.1126/science.290.5495.1306. PMID 11185405.
71.^ Nelson, K. E.; Levy, M.; Miller, S. L. (2000). "Peptide nucleic acids rather than RNA may
have been the first genetic molecule" (abstract). PNAS 97 (8): 3868–3871.
doi:10.1073/pnas.97.8.3868. PMID 10760258. PMC 18108.
http://www.pnas.org/cgi/content/abstract/97/8/3868.
72.^ Anastasi C, Crowe MA, Powner MW, Sutherland JD (2006). "Direct Assembly of
Nucleoside Precursors from Two- and Three-Carbon Units". Angewandte Chemie International
Edition 45 (37): 6176–9. doi:10.1002/anie.200601267. PMID 16917794.
http://www3.interscience.wiley.com/journal/112752038/abstract.
73.^ Powner MW, Sutherland JD (2008). "Potentially Prebiotic Synthesis of Pyrimidine β-D-
Ribonucleotides by Photoanomerization/Hydrolysis of α-D-Cytidine-2-Phosphate". ChemBioChem 9
(15): 2386–7. doi:10.1002/cbic.200800391. PMID 18798212.
http://www3.interscience.wiley.com/journal/121410594/abstract.
74.^ Huang W, Ferris JP (July 2006). "One-step, regioselective synthesis of up to 50-mers of
RNA oligomers by montmorillonite catalysis". J. Am. Chem. Soc. 128 (27): 8914–9.
doi:10.1021/ja061782k. PMID 16819887.
75.^ Orgel LE (November 2000). "Self-organizing biochemical cycles". Proc. Natl. Acad. Sci.
U.S.A. 97 (23): 12503–7. doi:10.1073/pnas.220406697. PMID 11058157. PMC 18793.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=11058157.
76.^ Muller, A.W.J. (1985). "Thermosynthesis by biomembranes: energy gain from cyclic
temperature changes". Journal of Theoretical Biology 115: 319–321.
 77.^ Muller, A.W.J. (1995). "Were the first organisms heat engines? A new model for biogenesis
and the early evolution of biological energy conversion". Progress in Biophysics and Molecular
Biology 63 (2): 193–231. doi:10.1016/0079-6107(95)00004-7. PMID 7542789.
78.^ Freeman Dyson (1985). Origins of Life. Cambridge: Cambridge University Press.
ISBN 0521626684.
79.^ Muller, A.W.J. (2005). "Thermosynthesis as energy source for the RNA World: a model for
the bioenergetics of the origin of life". Biosystems 82 (1): 93–102.
doi:10.1016/j.biosystems.2005.06.003. PMID 16024164.
80.^ Sun, F.J. and Caetano-Anolles, G. (2008). "The origin and evolution of tRNA inferred from
phylogenetic analysis of structure". Journal of Molecular Evolution 66 (1): 21–35. doi:10.1007/s00239-
007-9050-8. PMID 18058157.
81.^ Muller, A.W.J. ; Schulze-Makuch, D. (2006). "Sorption heat engines: simple inanimate
negative entropy generators". Physica A 362: 369–381. doi:10.1016/j.physa.2005.12.003.
82.^ Orgel, L. (1987). "Evolution of the genetic apparatus: a review". Cold Spring Harbor
Symposia on Quantitative Biology 52: 9–16. doi:10.1101/sqb.1987.052.01.004 (inactive 2010-01-05).
PMID 2456886.
83.^ Panno, Joseph (2005). The cell: evolution of the first organism. New York: Facts on File.
ISBN 0-8160-4946-7.
84.^ publications
85.^ Stuart Kauffman (1993). The Origins of Order: Self-Organization and Selection in Evolution
(Chapter 7). Oxford University Press. ISBN 978-0-19-507951-7
86.^ Dawkins, Richard (1996) [1986]. The Blind Watchmaker. New York: W. W. Norton &
Company, Inc. pp. 148–161. ISBN 0-393-31570-3.
 87.^ Cairns-Smith, A. G. (1982). Genetic takeover and the mineral origins of life . Cambridge,
UK: Cambridge University Press. ISBN 0-521-23312-7.
88.^ Bullard T, Freudenthal J, Avagyan S, Kahr B (2007). "Test of Cairns-Smith's crystals-as-
genes hypothesis". Faraday Discuss. 136: 231–45. doi:10.1039/b616612c.
http://www.rsc.org/publishing/journals/FD/article.asp?doi=b616612c.
89.^ Caroline Moore (16 July 2007). "Crystals as genes?". Chemical Science.
http://www.rsc.org/Publishing/ChemScience/Volume/2007/08/Crystals_as_genes.asp.
90.^ a b Nanobes–Intro microscopy-uk.org, Retrieved on 2008-01-14
91.^ Chang, Kenneth (2009-08-18). "From a Distant Comet, a Clue to Life". Space & Cosmos
(New York Times): p. A18. http://www.nytimes.com/2009/08/19/science/space/19comet.html?hpw.
Retrieved 2009-08-19.
92.^ Tough Earth bug may be from Mars - 25 September 2002 - New Scientist
93.^ Lin, Li-Hung; Pei-Ling Wang, Douglas Rumble, Johanna Lippmann-Pipke, Erik Boice, Lisa
M. Pratt, Barbara Sherwood Lollar, Eoin L. Brodie, Terry C. Hazen, Gary L. Andersen, Todd Z.
DeSantis, Duane P. Moser, Dave Kershaw, T. C. Onstott (October 2006). "Long-Term Sustainability
of a High-Energy, Low-Diversity Crustal Biome". Science 314 (5798): 479–482.
doi:10.1126/science.1127376. 5798. PMID 17053150.
94.^ Jason P. Dworkin; David W. Deamer, Scott A. Sandford, Louis J. Allamandola (January 30,
2001). "Self-assembling amphiphilic molecules: Synthesis in simulated interstellar/precometary ices".
Proc. Nat. Acad. Sciences 98 (3): 815–819. doi:10.1073/pnas.98.3.815. PMID 11158552.
PMC 14665. http://www.pnas.org/content/98/3/815.abstract.
95.^ "'Life chemical' detected in comet". BBC News. August 18, 2009.
http://news.bbc.co.uk/2/hi/science/nature/8208307.stm.
96.^ Origin of Life at the Weizmann Institute
 97.^ Brown MR, Kornberg A (November 2004). "Inorganic polyphosphate in the origin and
survival of species". Proc. Natl. Acad. Sci. U.S.A. 101 (46): 16085–7. doi:10.1073/pnas.0406909101.
PMID 15520374. PMC 528972. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=15520374.
98.^ The Origin Of Life
99.^ Pasek MA (January 2008). "Rethinking early Earth phosphorus geochemistry". Proc. Natl.
Acad. Sci. U.S.A. 105 (3): 853–8. doi:10.1073/pnas.0708205105. PMID 18195373. PMC 2242691.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=18195373.
100.^ Witt AN, Vijh UP, Gordon KD (2003). "Discovery of Blue Fluorescence by Polycyclic
Aromatic Hydrocarbon Molecules in the Red Rectangle". Bulletin of the American Astronomical
Society 35: 1381.
http://web.archive.org/web/20031219175322/http://www.aas.org/publications/baas/v35n5/aas203/18
9.htm.
101.^ Battersby, S. (2004). Space molecules point to organic origins. Retrieved January 11,
2004 from http://www.newscientist.com/article/dn4552-space-molecules-point-to-organic-origins.html
102.^ Astrobiology Magazine [1] Accessed 26 April 2008
103.^ Are Aliens Among Us? In pursuit of evidence that life arose on Earth more than once,
scientists are searching for microbes that are radically different from all known organisms Scientific
American. 19 November 2007
104.^ Hartman, Hyman (1998) "Photosynthesis and the Origin of Life" (Origins of Life and
Evolution of Biospheres, Volume 28, Numbers 4–6 / October, 1998)
[edit] Further reading
• Arrhenius, Gustaf; et al. (1997). "Entropy and Charge in Molecular Evolution—the Case of
Phosphate". Journal of Theoretical Biology 187 (4): 503–522. doi:10.1006/jtbi.1996.0385.
PMID 9299295.
• Buehler, Lukas K. (2000–2005) The physico-chemical basis of life,
http://www.whatislife.com/about.html accessed 27 October 2005.
• Davies, Paul (1998). The Fifth Miracle. Penguin Science, London. ISBN 0-140-28226-2.
• De Duve, Christian (January 1996). Vital Dust: The Origin and Evolution of Life on Earth .
Basic Books. ISBN 0-465-09045-1.
• Fernando CT, Rowe, J (2007). "Natural selection in chemical evolution". Journal of
Theoretical Biology 247 (1): 152–67. doi:10.1016/j.jtbi.2007.01.028. PMID 17399743.
• Hartman, Hyman (1998). "Photosynthesis and the Origin of Life". Origins of Life and
Evolution of Biospheres 28 (4–6): 515–521. doi:10.1023/A:1006548904157.
• Harris, Henry (2002). Things come to life. Spontaneous generation revisited . Oxford: Oxford
University Press. ISBN 0198515383.
• Hazen, Robert M. (December 2005). Genesis: The Scientific Quest for Life's Origins. Joseph
Henry Press. ISBN 0-309-09432-1. http://newton.nap.edu/books/0309094321/html.
• Gribbon, John (1998). The Case of the Missing Neutrino's and other Curious Phenomena of
the Universe. Penguin Science, London. ISBN 0-140-28734-5.
• Horgan, J (1991). "In the beginning". Scientific American 264: 100–109. (Cited on p. 108).
• Huber, C. and Wächterhäuser, G., (1998). "Peptides by activation of amino acids with CO on
(Ni,Fe)S surfaces: implications for the origin of life". Science 281 (5377): 670–672.
doi:10.1126/science.281.5377.670. PMID 9685253. (Cited on p. 108).
 • Knoll, Andrew H. (2003). Life on a Young Planet: The First Three Billion Years of Evolution
on Earth. Princeton University Press. ISBN 0691009783.
• Luisi, Pier Luigi (2006). The Emergence of Life: From Chemical Origins to Synthetic Biology .
Cambridge University Press. ISBN 0521821177.
• Martin, W. and Russell M.J. (2002). "On the origins of cells: a hypothesis for the evolutionary
transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from prokaryotes to
nucleated cells". Philosophical Transactions of the Royal Society: Biological sciences 358 (1429): 59–
85. doi:10.1098/rstb.2002.1183. PMID 12594918.
• Maynard Smith, John; Szathmary, Eors (2000-03-16). The Origins of Life: From the Birth of
Life to the Origin of Language. Oxford Paperbacks. ISBN 0-19-286209-X.
• Morowitz, Harold J. (1992) "Beginnings of Cellular Life: Metabolism Recapitulates
Biogenesis". Yale University Press. ISBN 0-300-05483-1
• NASA Astrobiology Institute: Earth's Early Environment and Life
• NASA Specialized Center of Research and Training in Exobiology: Gustaf O. Arrhenius
• Pitsch, Stefan; Krishnamurthy, Ramanarayanan; Arrhenius, Gustaf (2000). "Concentration of
Simple Aldehydes by Sulfite-Containing Double-Layer Hydroxide Minerals: Implications for
Biopoesis" (abstract). Helvetica Chimica Acta 83 (9): 2398 2411. doi:10.1002/1522-
2675(20000906)83:9<2398::AID-HLCA2398>3.0.CO;2-5. http://www3.interscience.wiley.com/cgi-
bin/abstract/73501648/ABSTRACT.
• Russell MJ, Hall AJ, Cairns-Smith AG, Braterman PS (1988). "Submarine hot springs and the
origin of life". Nature 336: 117. doi:10.1038/336117a0.
• Dedicated issue of Philosophical Transactions B on Major Steps in Cell Evolution freely
available.
 • Dedicated issue of Philosophical Transactions B on the Emergence of Life on the Early Earth
freely available.

[edit] External links

• The Deep Hot Biosphere Theory - Thomas Gold, article from PNAS Proc. Natl. Acad. Sci.
USA Vol. 89, pp. 6045-6049, July 1992 Microbiology
• The Origin of Life Video by John Maynard Smith
• "Harvard Team Creates the World's 1st Synthesized Cells"
• Martin M Hanczyc and Jack W Szostak. Replicating vesicles as models of primitive cell
growth and division. Current Opinion in Chemical Biology 2004, 8:660–664.PDF (192 KB)
• Martin A. Nowak and Hisashi Ohtsuki.Prevolutionary dynamics and the origin of evolution.
Proceedings of the National Academy of Sciences 2008
• "Exploring Life's Origins: a Virtual Exhibit"
• "SELF-REPLICATION: Even peptides do it" by Stuart A. Kauffman (web archive version as
original page no longer accessible)
• Origins of Life website including papers, resources, by Dr. Michael Russell at the U. of
Glasgow
• Possible Connections Between Interstellar Chemistry and the Origin of Life on the Earth
• Scientists Find Clues That Life Began in Deep Space—NASA Astrobiology Institute
• Self-organizing biochemical cycles—by Leslie Orgel
• How Life Began: New Research Suggests Simple Approach
 • Primordial Soup's On: Scientists Repeat Evolution's Most Famous Experiment –an article in
Scientific American. March 28, 2007
• Illustrations from Evolution (textbook)
• An abiogenesis primer for laymen

[hide]
v•d•e
Origin of life

Hypercycle · Protobiont · Universal common descent · Last universal ancestor · RNA world
hypothesis · Iron-sulfur world theory · PAH world hypothesis · Miller–Urey experiment · Panspermia

Retrieved from "http://en.wikipedia.org/wiki/Abiogenesis"

Categories: Evolutionarily significant biological phenomena | Evolutionary biology | Origin of life

 W000

Origine de la vie
Un article de Wikipédia, l'encyclopédie libre.
(Redirigé depuis Origines de la vie)
Aller à : Navigation, rechercher
Cet article est consacré aux origines de la vie d'un point de vue scientifique. Les aspects mythiques
et religieux sont traités dans l'article Cosmogonie. La précédente théorie scientifique de l'origine de la vie est
traitée dans l'article génération spontanée.
Les origines de la vie sur Terre demeurent incertaines. Cependant, de nombreuses théories
scientifiques existent pour expliquer l'apparition de la vie, telle que nous la connaissons aujourd'hui, dont on
pense qu'elle remonte à environ 3,5 à 3,8 milliards d'années.
Cet article traite des événements antérieurs à l'apparition des trois grandes lignées du vivant.
 Stromatolites du précambrien, dans la formation de Siyeh. En 2002, William Schopf a affirmé que
ces formations étaient âgées de 3,5 milliards d'années, elles seraient alors les plus anciennes traces de vie
sur Terre.
Stromatolites dans la Baie Shark (Australie).
Sommaire
[masquer]
• 1 L'apparition de la vie
• 1.1 L'origine des molécules organiques
• 1.1.1 Une explication : la condensation sur
surfaces minérales
• 1.1.2 L'expérience Urey-Miller et l'origine
des molécules organiques
• 1.1.2.1 L'exploitation de l'idée de
Miller
• 1.1.3 L'expérience de Powner-Sutherland et
l'origine des nucléotides à base pyrimidique
• 1.1.4 Asymétrie des biomolécules
• 1.1.4.1 Théories sur l'origine de
l'homochiralité
• 1.1.4.2 Cristaux et énantiomères
• 1.1.4.3 Asymétricité et
thermodynamique
• 1.1.4.4 Formation d'énantiomères
dans l'espace
• 1.2 Des molécules organiques aux protocellules
• 1.2.1 Compartiments isolés
L'apparition de la vie [modifier]
John Maynard Smith et Eörs Szathmáry définissent huit transitions majeures de l'évolution, dont trois
concernent l'apparition de la vie :
1. transition depuis des molécules autoréplicantes vers une population de molécules dans un
compartiment isolé ;
2. passage de réplicateurs indépendants aux chromosomes ;
3. transition d'un monde à ARN — où ce dernier joue le rôle de génome et d'enzyme — à un
monde à ADN et protéines.
John Desmond Bernal, quant à lui, suggère les trois étapes suivantes :
1. apparition de monomères organiques,
2. transitions vers des polymères organiques,
3. évolution depuis des molécules vers la cellule.
De fait, il n'existe pas de modèle « standard » pour décrire l'origine de la vie. Cependant le modèle le
plus couramment accepté est fondé sur l'enchaînement supposé des événements suivants :
1. Des conditions prébiotiques plausibles entraînent la création de molécules organiques
simples qui sont les briques de base du vivant.
2. Des phospholipides forment spontanément des doubles couches qui sont la structure de
base des membranes cellulaires.
3. Les mécanismes qui produisent aléatoirement des molécules d'ARN (acide ribonucléique),
en mesure d'agir comme des ARN-enzymes capables, dans certaines conditions très particulières,
 de se dupliquer. C'est une première forme de génome, et nous sommes alors en présence de
protocellules.
4. Les ARN-enzymes sont progressivement remplacées par des protéines-enzymes, grâce à
l'apparition des ribozymes, ceux-ci étant capables de réaliser la synthèse des protéines.
5. L'ADN apparaît et remplace l'ARN dans le rôle de support du génome, dans le même temps
les ribozymes sont complétés par des protéines, formant les ribosomes. C'est l'apparition de
l'organisation actuelle des organismes vivants.
Les étapes 2 et 3 sont parfois inversées, l'isolement en compartiment étant alors présenté après
l'apparition des ARN autoréplicants.
L'origine des molécules organiques [modifier]

Une explication : la condensation sur surfaces minérales [modifier]

Cristaux de pyrite
L'assemblage de petites molécules (comme les acides aminés) en macromolécules (comme les
protéines) nécessite l'élimination de molécules d'eau. Or, la thermodynamique indique qu'il est défavorable
de réaliser une telle condensation dans l'eau elle-même. Il est possible pour résoudre cette contradiction de
faire appel à des surfaces minérales, comme les micas, les argiles ou les pyrites. L'adsorption des petites
molécules sur ces surfaces les concentre et les modifie chimiquement, ce qui peut rendre la formation de
macromolécules plus favorable.
 L'argile, par exemple, se trouve très abondamment sur Terre et est constituée d'un empilement de
couches fines. Entre les différentes couches de l'argile peuvent se glisser certaines petites molécules
organiques, ce qui permet une adsorption importante. L'argile est aussi un catalyseur très efficace pour de
nombreuses réactions organiques, et aurait donc pu permettre la polymérisation des acides aminés et/ou
des acides nucléiques. Le chimiste anglais Cairns-Smith a développé cette hypothèse dans Seven clues to
the origin of life en 1985 (traduction française : L'énigme de la vie, 1990).

L'expérience Urey-Miller et l'origine des molécules organiques [modifier]

Article détaillé : Expérience de Miller-Urey.
En 1953, Stanley Miller, accompagné de Harold Urey, a voulu reproduire les conditions de la Terre
primitive. Ils ont enfermé dans un ballon des gaz (méthane CH 4, ammoniac NH3, hydrogène H2 et eau H2O)
et soumis le mélange à des décharges électriques pendant sept jours.
Ils ont obtenu des molécules organiques, les briques du vivant, et notamment de l'urée (CON2H4), du
formaldéhyde (H2CO), de l'acide cyanhydrique (HCN), des bases et des acides aminés (AA), certains
composés étant présents à plus de 2 %.
Miller et Urey ont utilisé une atmosphère réductrice, méthane CH4, NH3, H2, H2O et non pas une
atmosphère oxydante. Depuis l'expérience a été refaite plusieurs fois, en variant la composition de
l'atmosphère et la source d'énergie (utilisation du rayonnement ultraviolet notamment). Cependant, une
atmosphère moins réductrice (dioxyde de carbone CO2, azote N2, eau H2O) qui provient du volcanisme
donne de très mauvais rendements.
L'exploitation de l'idée de Miller [modifier]
Suite aux expériences de Miller, il a fallu déterminer les réactions chimiques qui se sont produites
dans l'enceinte (le ballon dans lequel il avait enfermé les différents gaz). Ainsi est née la chimie organique
dans l'eau.
Ces réactions nécessitent de fortes concentrations, des domaines de température et de pH très
étroits qui font que ces mécanismes sont très peu probables : une mare en voie d'assèchement pourrait
peut-être expliquer les fortes concentrations.
L'expérience fut à l'époque très critiquée à cause de cela. De plus, sa fiabilité a été remise en cause
car les molécules organiques obtenues pourraient a priori provenir d'une contamination extérieure. (?) Une
contamination extérieure est toutefois improbable pour les raisons évoquées ci-après.
En outre, la présence d'hydrogène (libre) dans l'atmosphère primitive est possible, mais seulement
en concentration réduite (de l'ordre du 1/100 de % ; ~100ppm ?), compte tenu de la masse de la Terre et de
la force de gravitation qui en résulte retenant très mal l'hydrogène vis à vis du vent solaire. Une des sources
d'hydrogène (libre) primitive aurait pu être les gaz volcaniques rejetant l'hydrogène issu de la réaction de
l'eau des roches avec des composés réducteurs.

L'expérience de Powner-Sutherland et l'origine des nucléotides à base pyrimidique

[modifier]
En 2009, Sutherland et son équipe[1] sont parvenu à établir une voie de synthèse des nucléotides à
base pyrimidique, l'acide uridylique et l'acide cytidylique. Les chercheurs butaient sur cette voie de synthèse
pré-biotique depuis 40 ans en raison de la difficulté de trouver la bonne façon de lier le ribose à la base
azotée. La clé de cette voie est de passer par un précurseur commun au ribose et la base azotée. Ce
précurseur, le 2-aminooxazol, est obtenu à partir de molécules organiques élémentaires : le glycéraldéhyde,
le cyanimide, le cyanoacétaldéhyde, le cyanoacétylène et le phosphate inorganique. Le mélange réactionnel
alimenté par de l'azote gazeux est soumis à un cycle de chauffage-refroidissement afin de simuler le cycle
d'évaporation d'une mare par le rayonnement solaire et l'alimentation par la pluie. Après une semaine le 2-
aminooxazol s'accumule dans le réacteur. Le précurseur se transforme ensuite en ribose et cytosine liés
ensemble. Le phosphate est ensuite ajouté au milieu réactionnel en présence d'UV durant trois jours
(l'absence d'ozone dans les conditions pré-biotiques engendrait un bombardement intense d'UV). L'acide
cytidylique se synthétise sous l'effet du rayonnement UV et quelques nucléotides portant une cytosine se
transforme en acide uridylique. Pour les deux autres nucléotides, l'équipe de Sutherland travaillent sur un
précurseur commun aux acides nucléiques à base purique.

Asymétrie des biomolécules [modifier]

 L'Alanine, un acide aminé existant sous une forme lévogyre (L-Ala) ou dextrogyre (D-Ala).
Les molécules « chirales » sont des molécules pouvant exister sous deux formes possibles :
lévogyre (gauche) et dextrogyre (droite), de la même façon que la main gauche et la main droite sont l'image
symétrique l'une de l'autre. On appelle énantiomères ces différentes formes.
Une analyse poussée des molécules obtenues dans l'expérience de Miller montre que l'on obtient un
mélange racémique de molécules (autant de formes droites que de gauches), alors que l'on sait depuis le
milieu du XIXe siècle (notamment avec des travaux de Pasteur en 1847) que les acides aminés naturels
n'existent pratiquement que sous une de leurs deux formes énantiomères (on parle alors d'homochiralité) : la
forme lévogyre.
On a cependant retrouvé des traces d'acides aminés dextrogyres sous forme libre, dans des
peptides ou même des protéines. Ces formes, peu fréquentes, auraient par ailleurs des fonctions
physiologiques. Dans le monde vivant, on constate cependant que les sucres présents dans l'ADN sont
uniquement de type dextrogyre, ou que les agents de saveur ont un goût différent selon leur forme.

Théories sur l'origine de l'homochiralité [modifier]

Il existe deux grandes catégories de théories expliquant l'homochiralité : les théories biotiques et les
théories abiotiques.
Dans la première, on postule que la vie serait apparue à partir d'un mélange d'énantiomères, et que
l'homochiralité ne serait apparue que progressivement au cours de l'évolution. Selon Laurent Nahon, ce type
de théorie n'est cependant plus beaucoup soutenu, du fait que l'on a découvert que les protéines ne peuvent
se replier correctement si les acides aminés qui les composent ne sont pas chiraux. On estimerait donc que
l'homochiralité serait plutôt antérieure à l'apparition de la vie, ce sont les théories abiotiques.

Cristaux et énantiomères [modifier]

Les propriétés des cristaux permettent d'imaginer un scénario : certains cristaux inorganiques
exposent des faces possédant une chiralité intrinsèque, comme la calcite ou le quartz. Ces faces pourraient
adsorber préférentiellement une des deux formes énantiomères et donc la concentrer sélectivement aux
dépens de l'autre[2]. Cependant, rien n'indique que les cristaux aient pu jouer un rôle dans la formation de
molécules organiques.

Asymétricité et thermodynamique [modifier]

Il a été découvert que la matière est intrinsèquement asymétrique. « Lorsque l'on place des atomes
de cobalt dans une géométrie asymétrique, c'est-à-dire dans des champs magnétiques, les électrons
produits de la désintégration de neutron se déplacent toujours dans la direction opposée à leur spin (aligné
sur le champ). Les électrons sont donc intrinsèquement gauches. » Cette expérience de Tsung Dao Lee et
Chen Ning Yang qui reçurent le prix Nobel en 1957 peut être reproduite avec n'importe quel atome. Un gaz
de vapeur de césium par exemple dans un champ électromagnétique a un pouvoir rotatoire. C'est ce sur
quoi a travaillé Marie-Anne Bouchiat, directrice de recherche CNRS à l'ENS. Ce phénomène est facilement
observable sur les atomes lourds car la force mise en jeu est la force d'interaction faible entre le noyau et
l'électron.
Ainsi il a été calculé que les acides aminés naturels sont thermodynamiquement plus stables que
leur image dans un miroir.
Formation d'énantiomères dans l'espace [modifier]

Une météorite
Dans les années 1970, des acides aminés ont été découverts dans la météorite de Murchison, or, ils
étaient présents majoritairement sous leur forme lévogyre. On a ainsi découvert 70 acides aminés différents,
dont 3 seulement font partie des 20 acides aminés naturels. Le pourcentage exact de molécules lévogyres
est cependant beaucoup discuté, du fait d'éventuelles contaminations, et varie entre 50 % et -5 % suivant les
équipes de recherche.
L'idée que l'homochiralité aurait pour origine des molécules venues de l'espace s'est donc
développée.
 Les énantiomères absorbent différemment la lumière lorsque celle-ci est polarisée «
circulairement » droite ou gauche. Or, ces molécules se dégradent après absorption, conduisant donc à un
excès d'une forme énantiomérique.

La nébuleuse d'Orion
On a découvert, en 1997, que la nébuleuse d'Orion produit de la lumière polarisée circulaire à 17 %
dans l'infrarouge (IR). L'infrarouge n'a pas assez d'énergie pour casser des liaisons covalentes, mais on peut
supposer que les ultraviolets (UV) sont également polarisés circulairement.
Une expérience menée en 2005 a montré qu'un mélange racémique d’un acide aminé simple irradié
par un rayonnement UV conduit à un mélange homochiral. Dans cette expérience, la leucine est utilisée à
l'état solide, reproduisant les conditions spatiales. Après une irradiation par un rayonnement synchrotron
polarisé circulairement droit (proche de la longueur d'onde observée dans l'espace, dans l'UV lointain),
l'expérience permet d'obtenir un excès de l'énantiomère lévogyre de 2,6 %.
 Or, il suffit d'un excès de 1 % dans des réactions qui s'entretiennent pour conduire à un mélange
homochiral de 100 %. D'après Laurent Nahon, aucune expérience liée à des théories concurrentes n'est
parvenue à un tel excès.

Des molécules organiques aux protocellules [modifier]

Schéma d'une cellule animale.

Aujourd'hui, de nombreux modèles résolvent le problème de l'apparition des molécules organiques.
Les scientifiques arrivent à produire de nombreuses petites molécules biologiques (acides aminés, sucres,
bases nucléiques) dans des conditions prébiotiques en laboratoires.
Les expériences de Miller et les modèles qui en sont dérivés ne fournissent pas d'explication sur les
étapes suivantes qui incluent la transition des monomères aux biopolymères, puis aux protocellules et
finalement aux cellules vivantes ayant un métabolisme de base. Aussi les scientifiques ont exploré d'autres
voies de recherche.
 Compartiments isolés [modifier]
L'apparition de compartiments isolés par une membrane pose de sérieux problèmes. Les
membranes des cellules vivantes sont composées de lipides, or on connaît aujourd'hui des acides gras à
longue chaîne qui peuvent spontanément former des petites membranes sphériques. Bien que l'on puisse
produire de tels compartiments en laboratoires, ces acides gras restent synthétisés par des enzymes. Le
processus permettant de former de tels compartiments en l'absence de ces enzymes demeure inconnu.

Protocellules [modifier]
Un compartiment isolé par une membrane ne forme cependant pas une protocellule. Selon Maynard
Smith, deux conditions sont nécessaires pour former une véritable protocellule :
1. Les molécules capables de répliquer la forme de base (les réplicateurs) doivent se lier entre
elles en un « chromosome », formant ainsi une unité structurelle, garantissant aux réplicateurs de
former un tout cohérent après la réplication ;
2. la membrane doit posséder des mécanismes d'échange avec le milieu extérieur, autres que
les systèmes à protéines actuels.
Schéma général d'un chromosome eucaryote. Schéma d'une membrane semi-perméable.
 L'apparition du génome [modifier]

Les modèles « gènes d'abord » [modifier]

Une molécule d'ADN.

Dans ce modèle, l'apparition du génome a précédé l'apparition du métabolisme. Des molécules
d'ADN ou d'ARN auraient ainsi existé seules, s'autorépliquant à partir des molécules présentes dans leur
environnement. Les « individus » sont donc représentés par les molécules d'acides nucléiques elles-mêmes.
 Les modèles « métabolisme d'abord » [modifier]
Plusieurs modèles rejettent l'idée de l'autoréplication d'un gène « nu » et font l'hypothèse de
l'apparition d'un métabolisme primitif qui aurait précédé l'émergence de la réplication de l'ARN. Une des
premières versions de cette hypothèse fut présentée en 1924 par Alexander Oparin avec son idée de
vésicules primitives capables de se répliquer, à une époque où on ne connaissait pas encore la structure de
l'ADN.
D'autres variantes sont apparues dans les années 1980 et 1990 comme la théorie de Günter
Wächtershäuser sur un monde sulfuro-ferreux, ou les modèles de Christian de Duve basés sur la chimie des
thioesters.
D'autres arguments plus abstraits ont aussi été présentés. On peut citer les modèles mathématiques
de Freeman Dyson au début des années 1980 sur la probabilité de l'émergence d'un métabolisme sans
présence de gènes, ou encore les travaux de Stuart Kauffman sur les ensembles globalement
autocatalytiques (voir génération spontanée pour une présentation des idées de Kauffman sur l'origine de la
vie).

Métabolisme et génome : un monde à ARN [modifier]

Article détaillé : Hypothèse du monde à ARN.
L'hypothèse du monde à acide ribonucléique (ARN) est que l'ARN était la principale — et sans doute
la seule — forme de vie avant l'émergence de la première cellule à ADN. C'est Walter Gilbert qui a utilisé pour
la première fois le terme « monde à ARN » (« RNA world » en anglais) en 1986.
 L'hypothèse d'un monde à ARN a aujourd'hui la faveur des scientifiques et est fondée sur plusieurs
éléments. Notamment sur le fait que l'ARN est en théorie capable aussi bien d'assurer des tâches
métaboliques que d'être le support d'une information génétique.

Stockage et réplication [modifier]

Une protéine liée à un brin d'ADN.

L'ARN a la capacité de stocker une information, en utilisant un code génétique similaire à celui de
l'ADN. L'ARN peut également se comporter comme un ribozyme (de la contraction de ribose et enzyme) et
catalyser certaines réactions, tout comme les protéines. Du point de vue de la reproduction, cette molécule
possède donc deux fonctions primordiales : le stockage de l'information et la catalyse nécessaire à
l'autoréplication.
L'ADN peut aussi se recopier lui-même, mais seulement avec l'assistance de protéines. Les
protéines sont de très bons catalyseurs mais elles sont incapables de stocker l'information requise pour leur
propre réplication. L'ARN est lui capable à la fois de catalyse et d'autoréplication. Ainsi, le ribosome est un
ribozyme, dans le sens où le responsable de la synthèse des protéines n'est pas une protéine (comme c'est
le cas dans la grande majorité des catalyses d'une cellule vivante) mais l'ARN ribosomique lui-même. Ces
ribozymes peuvent se replier dans l'espace, faisant apparaître un site actif pour une catalyse, à l'instar des
protéines.
L'ADN, formant une double hélice rigide, ne peut se replier pour jouer un rôle de catalyseur.

Efficacité des protéines [modifier]

 La structure « quaternaire » d'une protéine.
Les protéines sont des catalyseurs très efficaces, bien plus que les ribozymes. De même, il existe 20
acides aminés dans le monde vivant, mais seulement quatre nucléotides, les protéines sont donc bien plus
diversifiées que les ARN.
D'un point de vue évolutif, il est donc peu probable qu'une protéine-enzyme ait été remplacée par
une ARN-enzyme. À l'inverse, si les ARN sont bien apparus avant les protéines, il est plausible qu'ils aient
été remplacés par des protéines, plus efficaces.
 Cet argument est étayé par le fait que l'ARN joue un rôle dans la synthèse des protéines, via son rôle
fondamental dans les ribosomes actuels. L'ARN aurait donc en quelque sorte conduit à l'apparition des
protéines.
Les protéines utilisées dans la structure du ribosome seraient donc venues plus tard : les premières
protéines auraient été sélectionnées selon leur capacité d'amélioration du fonctionnement des ribozymes,
pour finalement se substituer à eux.

Une grande distribution phylogénétique [modifier]

Les ARN sont présents dans les trois lignées du monde vivant (archées, procaryotes, eucaryotes).
Ils accomplissent dans chacun un grand nombre de tâches différentes, les plus connus sont l'ARN messager
(ARNm, véhiculant l'information génétique de l'ADN vers les ribosomes), l'ARN de transfert (ARNt, faisant le
lien entre acide nucléique et acide aminé) et l'ARN ribosomique (ARNr, composant structuraux et
fonctionnels des ribosomes). À côté de ceux-ci, on peut trouver un grand nombre d'ARN impliqué dans des
fonctions tels que des catalyses, des régulations de l'expression de gènes, des contrôles, des défenses
antivirales, des extinctions de gènes, des inhibitions de synthèses de protéines, des restaurations
génomiques, etc. C'est le cas des ARN interférents (ARNs), dont certains chercheurs qualifient le
mécanisme d'« universel ». Les ARNtm des procaryotes, ont également plusieurs fonctions : ils jouent à la
fois les rôles d'ARN de transfert et d'ARN messager.
Malgré cette grande diversité structurelle et fonctionnelle, la répartition des ARN permet de retrouver
le découpage du vivant. Ainsi, les petits ARN nucléolaire ne sont partagés que par les archées et les
eucaryotes, l'ARN de la télomérase n'est lui présent que chez les eucaryotes alors que les procaryotes sont
les seules à posséder des ARNtm. De la même façon, les trois grands types d'ARN (ARNt, ARNm et ARNr)
sont présents dans les trois lignées.
Le cas de l'ARN de transfert [modifier]
 La structure de l'ARN de transfert.
Le rôle de l'ARNt est de transporter un acide aminé vers le ribosome, où s'effectuera la liaison avec
un autre acide aminé, pour former un polypeptide (donnant ainsi une protéine). Il existe plusieurs ARNt,
possédant chacun trois nucléotides : l'anticodon. L'anticodon correspond à un codon, porté par l'ARNm qui
définit l'ordre d'assemblage des acides aminés par le ribosome.
La particularité de l'ARNt est qu'il est, malgré sa petite taille, en partie constitué de nombreux
nucléotides que l'on ne rencontre pas ailleurs. Ces nucléotides « exotiques » auraient ainsi une origine
prébiotique, vestiges d'un monde à ARN. On retrouve ainsi ces composants dans l'ensemble des trois
domaines du vivant.

Virus et ARNt [modifier]

Il est relativement fréquent d'observer des virus à ARN ou des viroïdes portant des motifs similaires à
de l'ARNt. Ainsi, dans le viroïde PSTV (Potato Spindle Tuber Viroïd) et l'ARNt de la tyrosine, on trouve les
mêmes motifs structuraux en « feuille de trèfle ». Pour Marie-Christine Maurel, « ces derniers jouent un rôle
fondamental dans le vivant et leur ancienneté ne fait pas de doute ».
Autre structure surprenante : chez le virus TYMV (Turnip Yellow Mosaic Virus), l'amorce de la
traduction du génome du virus en protéine se fait par le biais d'une structure de type ARNt qui amorce sa
propre traduction et qui fixe un acide aminé.
 Structure du virus PSTV.

Évolution dirigée d'ARN [modifier]

En 1990, Larry Gold et Jack Szostak ont mis au point une méthode visant à diriger l'évolution d'ARN,
afin de sélectionner ceux montrant une activité catalytique. Ils ont depuis réussi à obtenir des ribozymes
capables de lier des nucléotides entre eux, de lier des acides aminés à des ARNs, d'effectuer des réactions
d'oxydo-réductions, de se lier à des composants de la membrane, etc.
Il est donc en théorie possible, sur ce modèle, que l'ARN seul suffise à établir un métabolisme
primitif. Toutefois, il reste encore à découvrir un ARN capable de se répliquer lui-même.

ARN et hérédité [modifier]

L'ARN joue un rôle dans la transmission de l'activité des gènes : un tel mécanisme (qualifié
d'épigénétique) n'est pas lié à l'ADN, et serait une preuve de la capacité de l'ARN à participer à l'hérédité.

À l'origine de l'ADN dans la cellule [modifier]

Du point de vue de la biologie cellulaire, l'ADN est produit par modification d'un ARN : les
désoxyribonucléotides (précurseurs de l'ADN) sont en effet produits à partir des ribonucléotides (précurseurs
des ARN). De plus, le groupement thymine (identifié comme T dans le code génétique), est construit à partir
d'un groupement uracile (U). Or, bien que l'uracile soit spécifique à l'ARN et la thymine à l'ADN, le
groupement U est, au moment de la synthèse, déjà fixé sur un désoxyribonucléotide.
Uracile. Thymine.
De l'ARN à l'ADN [modifier]

Comparaison entre une molécule d'ARN (à gauche) et une molécule d'ADN (à droite).
Dans l'hypothèse du monde à ARN, apparaissent tout d'abord des viroïdes ressemblant à des ARN
auto-catalytiques, présents dans des compartiments isolés (qu'ils soient membranaires ou cristallins). Puis
des protocellules, capables de métabolisme archaïque, sont soumises à une évolution darwinienne, évoluant
ainsi vers des cellules à ARN, capables de présenter une activité variée et complexe.
 Ribozymes ou protéines ? [modifier]
Bien que l'ARN soit donc à l'origine de l'ADN dans le métabolisme cellulaire, cette réaction est très
difficile à réaliser. De fait, dans les trois lignées, elle est catalysée par des protéines spécialisées : les
ribonucléotides-réductases. De plus, cette réaction est très coûteuse en énergie, du fait de la réduction du
ribose, et elle produit des radicaux libres, très réactifs, sur la protéine. L'ARN étant une molécule fragile, il
paraît improbable qu'elle puisse supporter des radicaux libres sans l'intervention de protéines.
Ainsi, l'origine de l'ADN trouve vraisemblablement sa source après l'apparition des protéines,
indispensables à chaque étape de sa synthèse à partir de précurseurs de type ARN, au sein de la cellule.
 Intérêt de l'ADN [modifier]

Structure chimique de l'ADN.

L'ADN présente un certain nombre d'avantages sur l'ARN, en termes de conservation de l'intégrité
de l'information génétique.
Tout d'abord il se casse moins facilement, car le désoxyribose de l'ADN contient un atome d'oxygène
de moins que le ribose de l'ARN. Or l'oxygène peut facilement interagir sur les liaisons entre nucléotides,
posant alors un problème de stabilité.
 Ensuite, l'ADN permet la réparation d'un problème récurrent : la transformation du groupement
thymine (T) en uracile (U). En effet, l'uracile, uniquement présent dans l'ARN, est une anomalie dans l'ADN.

Avantage sélectif de l'ADN : l'hypothèse du virus [modifier]

Structure de base d'un virus.

Les avantages en termes de stabilité de l'ADN pourraient ne pas suffire à expliquer son adoption.
Ainsi, Patrick Forterre avance l'hypothèse qu'un avantage sélectif supplémentaire peut être dû aux conflits
entre virus et cellules vivantes.
 Dans ce modèle, le premier organisme à ADN serait un virus. L'ADN conférerait au virus le pouvoir
de résister à des enzymes dégradant les génomes à ARN, arme de défense probable des cellules. On
retrouve le même principe chez des virus actuels, qui altèrent leur ADN pour résister à des enzymes
produites par des bactéries infectées.
Actuellement, on peut observer que les enzymes nécessaires à la traduction de l'ARN vers l'ADN
sont très présentes chez les rétrovirus, dont le génome est porté par de l'ARN. De la même façon, de
nombreux virus codent leurs propres enzymes de synthèse de l'ADN.
Cette hypothèse est également corroborée par la découverte de virus à ADN, dont celui-ci contient,
non pas des groupements thymines, mais des groupements uraciles. Du point de vue évolutif, il y aurait donc
eu d'abord apparition des désoxyribonucléotides, puis de l'ADN à uracile (ADN-U), puis d'ADN à thymine
(ADN-T), qui se serait progressivement imposé. D'après Patrick Forterre, il est même probable que l'ADN-T
ait été « inventé deux fois », chez des virus différents.
Les virus à ARN seraient ici des reliques du monde à ARN, les virus à ADN-U seraient alors des
reliques du monde ayant précédé celui à ADN-T.
 Les virus, premiers organismes à ADN [modifier]

Schéma de la réplication de l'ADN chez les eucaryotes.

Les virus à ADN pourraient être plus anciens que la première cellule à ADN : la première cellule à
ADN l'aurait donc emprunté à un ou plusieurs virus, sous la pression d'une course aux armements (théorie
de la reine rouge).
Didier Raoult et Jean-Michel Claverie ont ainsi découvert le mimivirus : un virus géant à ADN (son
génome étant deux fois plus long que le plus petit génome bactérien connu). La particularité de ce virus est
qu'il peut produire des protéines impliquées dans la traduction de l'ARN en protéines (comme des enzymes
chargeant des acides aminés sur des ARNt), il pourrait donc avoir pour ancêtres des virus plus anciens que
la première cellule à ADN.
Eugène Koonin et ses collègues ont mis en avant, en comparant des génomes séquencés, que la
plupart des enzymes impliqués dans la réplication de l'ADN sont différents entre les eubactéries et les
eucaryotes (accompagnés des archées). Ils en concluent que l'ADN aurait été inventé indépendamment
dans la lignée des eubactéries et celle conduisant aux eucaryotes et aux archées.[réf. nécessaire]
De même, les enzymes de réplications des virus à ADN sont très différentes d'un virus à l'autre, ainsi
que par rapport aux enzymes cellulaires jouant le même rôle.
Ces indices laissent penser que les enzymes liées à l'ADN sont apparues au cours d'un « premier
âge » du monde à ADN, où existaient cellules à ARN et virus à ARN et à ADN.

Passage de l'ADN dans les cellules [modifier]

La nature du génome du plus ancien ancêtre commun à tous les êtres vivants (que les scientifiques
prénomment LUCA) reste inconnue : faisait-il encore partie du monde à ARN, ou avait-il déjà un génome à
ADN ? Quoi qu'il en soit, LUCA est le fruit d'une longue évolution. Le génome des premières cellules, qui ont
précédé LUCA, était sans doute constitué par des molécules d'ARN et non pas d'ADN. Contrairement à
l'ADN, l'ARN peut en effet jouer à la fois le rôle d'enzyme et de matériel génétique. Stanley Miller et Christian
de Duve pensent que l'apparition de l'ARN a été elle-même un évènement tardif, en effet, cette molécule ne
semble pas pouvoir être synthétisée par les méthodes simples de la chimie prébiotique. L'ARN aurait donc
été précédé par des molécules dont nous ne connaîtrons sans doute jamais la nature exacte.
 Ce qui amène les scientifiques à penser que l'ARN a précédé l'ADN résulte de ce constat : les trois
grandes lignées du vivant ne partagent que le système de synthèse des protéines, alors qu'elles diffèrent sur
le système de réplication de l'ADN.
Il existe alors plusieurs hypothèses pour le passage de l'ADN (d'origine viral) dans les cellules
vivantes : soit ce passage s'est produit une seule fois, soit il a pu avoir lieu plusieurs fois, indépendamment
dans les différentes lignées. Dans le premier cas, les premiers gènes des enzymes de réplications auraient
donc été remplacés par la suite par ceux d'un autre virus, formant ainsi les trois lignées.
L'hypothèse de Patrick Forterre est que les trois lignées du vivant trouvent leurs sources dans le
remplacement du génome à ARN par le génome à ADN de trois virus différents. On retrouve notamment ici
l'hypothèse d'une origine virale du noyau des eucaryotes.
Les travaux de Carl Woese semblent appuyer cette hypothèse, en démontrant que la vitesse
d'évolution des protéines semble avoir chuté au moment de l'apparition des trois lignées. Cette diminution
serait due au passage de l'ARN à l'ADN, les génomes à ADN étant plus stables, et donc moins sensibles aux
mutations.
Dans ce scénario, l'apparition de trois lignées uniquement s'explique par le fait que les cellules à
ADN ont peu à peu supplanté les cellules à ARN, empêchant ainsi l'apparition de nouvelles lignées par
passage ARN→ADN.
Autres modèles [modifier]
Milieu de collision Planète/Noyaux de comètes [modifier]
Selon des travaux de modélisation travaux présentés en 2010 [3], la compression brutale des
matériaux constituant le noyau de glace d'une comète, au moment du choc avec une planète pourrait donner
lieu à la production d'acides aminés. Cette modélisation a pris comme base 210 molécules diluées dans un
mélange d'eau, de méthanol, d'ammoniac, de dioxyde de carbone et de monoxyde de carbone estimés
couramment présentes dans le noyau des comètes. Avec une vitesse d'impact de 29 km/s, la pression atteint
10 gigapascals et une température de 700°Kelvin. Selon les auteurs, d'autres simulations, à pressions et
températures plus élevées conduisent à des réactions chimiques encore plus complexes. Selon le modèle,
une simulation de 47 gigapascals et une température de 3.141 Kelvin pour les 20 premières picosecondes
de l'impact fait naitre des molécules complexes, dont de grosses molécules à liaisons carbone-azote qui
pourraient être les premières briques de la vie. Le calcul a nécessité environ million d'heures d'ordinateur sur
le superordinateur du cluster Atlas du Lawrence Livermore.

Sources hydrothermales : le monde du soufre [modifier]

Les sources hydrothermales ont été découvertes en 1977 à 2 600 mètres de profondeur, là où deux
plaques tectoniques se séparent.
Les monts hydrothermaux sont situés sur la couche sédimentaire. Leur diamètre à la base varie de
25 à 100 mètres et leur hauteur varie de 70 à 100 mètres. Les cheminées de ces fumeurs sont parfois
recouvertes d'une croûte d'oxyde de manganèse. Les fumeurs situés sur ces monts sont composés d'un
solide friable dont la couleur varie du gris noir à l'ocre, ce sont des sulfures de fer, de cuivre et de zinc.
 Ces sources sont particulièrement intéressantes car on y a trouvé la vie où on la croyait impossible :
milieu privé d'oxygène, à haute température, chargé de métaux et de soufre, dans l'obscurité la plus totale.
Cependant les gradients de température importants autour de ces zones et le fait que les ultraviolets
destructeurs ne parviennent pas si profondément (alors qu'ils détruisent toute molécule formée à la surface)
sont de bonnes conditions pour l'apparition de la vie.
Ces organismes ont les mêmes formes que ceux que l'on connaît plus près de la surface (ADN,
protéines, sucres...) mais puisent leur énergie de l'oxydation du sulfure d'hydrogène (H2S) pour transformer
le carbone minéral en matière organique.
D'autre part, des expériences ont été menées, au laboratoire de géophysique de Washington, et ont
montré que dans les conditions qui existent autour des évents, il y a formation d'ammoniac (NH3), forme
réduite de l'azote qui est tant nécessaire à la formation des molécules organiques de la première partie et qui
n'existait pas dans l'atmosphère oxydante. Les sources hydrothermales sont donc de bonnes sources de
NH3.

Une origine extraterrestre primitive (exogenèse) [modifier]

Une hypothèse alternative est que la vie se soit d'abord formée hors de la Terre. Les composés
organiques sont relativement fréquents dans l'espace, notamment dans les zones lointaines du système
solaire où l'évaporation des composés volatils est très réduite. Certaines comètes sont enrobées dans des
couches de matière sombre, qu'on pense être une sorte de bitume formé par une combinaison de composés
carbonés simples et de rayons ultraviolets. La pluie de matériaux cométaires sur la Terre primitive pourrait
avoir apporté des quantités de molécules organiques complexes, ce qui aurait causé l'apparition de la vie sur
Terre.
 Une hypothèse plus large est la panspermie : la vie même serait apparue dans l'espace puis
disséminée sur Terre. Selon une variante, la vie serait apparue sur Mars d'abord et des impacts de comètes
et d'astéroïdes sur Mars auraient projeté du matériel de la surface martienne sur Terre. Il est encore plus
difficile de trouver des indices pour justifier ces hypothèses que les théories plus classiques.
Ces théories d'une origine extraterrestre n'expliquent pas directement comment la vie est apparue,
car elles ne font a priori que reporter le problème. Cependant, elles élargissent les conditions dans
lesquelles la vie a pu apparaître dans l'univers. Les futurs échantillons de sols ramenés de Mars et de
comètes permettront peut-être d'obtenir de nouveaux éléments de réponse.

Notes et références [modifier]

1. ↑ Matthew W. Powner, Béatrice Gerland and John D. Sutherland, Synthesis of activated
pyrimidine ribonucleotides in prebiotically plausible conditions, Nature, Vol 459, 14 May 2009, pp.
239-242
2. ↑ Robert M. Hazen, Timothy R. Filley, and Glenn A. Goodfriend, PNAS May 8, 2001 vol. 98
no. 10 5487–5490Y
3. ↑ travaux de Nir Goldman et de ses collègues du Lawrence Livermore National Laboratory à
Livermore, présentés le 24 Mars 2010 devant l'American Chemical Society à San Francisco,
Californie, et relayés par une brève du Journal Nature [archive] (26 Mars 2010 ; On line :
doi:10.1038/news.2010.152)
Sources [modifier]
• (en) John Desmond Bernal, The origin of life, éditions Weidenfeld & Nicolson, 1967.
• (fr) Guillemette Lauters, Olivier Lenaerts, Daniel Rousselet et Eric Depiereux, Bioscope :
L’origine de la vie.
• (fr) John Maynard Smith, Eörs Szathmáry, Les Origines de la vie, éditions Dunod, 2000.
(ISBN 2100048600)
• (en) J. William Schopf, Anatoliy B. Kudryavtsev, David G. Agresti, Thomas J. Wdowiak,
Andrew D. Czaja, « Laser−Raman imagery of Earth's earliest fossils », dans Nature, volume 416,
pages 73–76, mars 2002 [(en) lire en ligne].
• (fr) Marie-Christine Maurel, La Naissance de la vie : de l'évolution prébiotique à
l'évolution biologique, éditions Dunod, 2003 (3e édition). (ISBN 2100068822)
• (en) N. Fujii, T. Saito, « Homochirality and life », dans The Chemical Records, volume 4,
o
n 5, pages 267–78, 2004 [(en) lire en ligne]
• (fr) Uwe Meierhenrich, Un pas vers l’origine de l’homochiralité…, novembre 2005.
• (fr) Patrick Forterre, « L'origine du génome », dans Les dossiers de La Recherche :
l'histoire de la vie, les grandes étapes de l'évolution , no 19, pages 34–40, mai 2005.
• (fr) Marc-André Selosse, « Quelle parenté entre les trois grandes lignées du vivant ? »,
dans Les dossiers de La Recherche : l'histoire de la vie, les grandes étapes de l'évolution , no 19,
pages 42–45, mai 2005.
 • (fr) « Laurent Nahon : « L'asymétrie des biomolécules vient de l'espace » », propos
recueillis par Franck Daninos, dans La Recherche, no 390, octobre 2005.
• (en) U. Meierhenrich, L. Nahon, C. Alcaraz, J. Bredehöft, S. Hoffmann, B. Barbier, A.
Brack, « Asymmetric VUV photodecomposition of the amino acid D,L-Leucine in the solid state »,
dans Angewandte Chemie International Edition, volume 44, pages 5630–5634, juillet 2005 [(en) lire
en ligne].

Bibliographie [modifier]
• (fr) Le site de la Société Française d'Exobiologie (SFE).
• (fr) Conférence de Patrick Forterre : "Regards croisés sur les origines de la vie" (2009) à
l'ENS Cachan.
• (en) Uwe Meierhenrich: Amino acids and the asymmetry of life, Springer-Verlag, 2008.
(ISBN 978-3-540-76885-2)
• (en) Site dédié au mimivirus.
• (en) Anthony M. Poole, What is the Last Universal Common Ancestor ?, septembre 2002.
• (en) Antonio Lazcano The Origins of Life. Have too many cooks spoiled the prebiotic
soup?, Natural History magazine, février 2006.
• (en) Erwin Schrödinger, What is Life?, 1944. (fr) Traduction française : Qu'est-ce que
la vie?, ed. Seuil.
• (fr) Joël de Rosnay, Les Origines de la vie, éditions du Seuil, 1966. (ISBN 2020002167)
• (fr) Jean C. Baudet, La vie expliquée par la chimie, Vuibert, Paris, 2006.
 • (fr) André Brack, Bénédicte Leclercq, La vie est-elle universelle ? – Des premiers êtres
vivants à l'exploration spatiale, éditions EDP Sciences, 2003. (ISBN 2-86883-674-7).
• (fr) Sous la direction de Muriel Gargaud, Didier Despois, Jean-Paul Parisot,
L'Environnement de la Terre Primitive, Collection : L'origine de la Vie sur Terre et dans l'Univers,
Presses universitaires de Bordeaux, 2001. (ISBN 2-86781-267-4).

Voir aussi [modifier]

Articles connexes [modifier]
• Biologie, Biologie cellulaire, vie
• Biochimies hypothétiques
• Exobiologie
• Habitabilité d'une planète
• Génération spontanée
• Protéinoïde

Liens externes [modifier]

• (fr) Écouter La leçon consacrée aux origines chimiques de la vie et dispensée par
Jacques Reisse au Collège Belgique (2009).
• (fr) Comprendre facilement l'origine de la vie sur la Terre
 • (fr) Un site qui explique avec des mots simples les origines de la vie
• (fr) Les trois grandes théories de l'apparition de la vie sur Terre

[Enrouler]
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 [Dérouler]
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Catégories : Origine de la vie | Histoire évolutive | Métabolisme | [+]

 W000

Life
From Wikipedia, the free encyclopedia

Jump to: navigation, search

For other uses, see Life (disambiguation).

Life (Biota)
Life on a rocky peak in the Waitakere Ranges

Scientific classification

Domains and Kingdoms

Life on Earth:
• Non-cellular life (viruses) * *
• Cellular life
• Bacteria
• Archaea
• Eukarya
 • Protista
• Fungi
• Plantae
• Animalia
Life (cf. biota) is a characteristic that distinguishes objects that have signaling and self-sustaining
processes (biology) from those that do not,[1][2] either because such functions have ceased (death), or else
because they lack such functions and are classified as inanimate.[3]
In biology, the science of living organisms, life is the condition which distinguishes active organisms
from inorganic matter.[4] Living organisms undergo metabolism, maintain homeostasis, possess a capacity
to grow, respond to stimuli, reproduce and, through natural selection, adapt to their environment in
successive generations. More complex living organisms can communicate through various means.[1][5] A
diverse array of living organisms (life forms) can be found in the biosphere on Earth, and the properties
common to these organisms—plants, animals, fungi, protists, archaea, and bacteria—are a carbon- and water-
based cellular form with complex organization and heritable genetic information.
In philosophy and religion, the conception of life and its nature varies. Both offer interpretations as to
how life relates to existence and consciousness, and both touch on many related issues, including life
stance, purpose, conception of a god or gods, a soul or an afterlife.
Contents
[hide]
• 1 Early theories about life
• 1.1 Materialism
• 1.2 Hylomorphism
• 1.3 Vitalism
• 2 Definitions
• 2.1 Biology
• 2.2 Biophysics
• 2.3 Living systems theories
• 3 Origin of life
• 4 Conditions for life
• 4.1 Range of tolerance
• 4.2 Extremophiles
• 5 Classification of life
• 6 Extraterrestrial life
• 7 Death
• 7.1 Extinction
• 7.2 Fossils
• 8 See also
• 9 References
[edit] Early theories about life
[edit] Materialism

Plant life
Herds of zebra and impala gathering on the Masai Mara plain

An aerial photo of microbial mats around the Grand Prismatic Spring of Yellowstone National Park.
 Some of the earliest theories of life were materialist, holding that all that exists is matter, and that all
life is merely a complex form or arrangement of matter. Empedocles (430 B.C.) argued that every thing in the
universe is made up of a combination of four eternal 'elements' or 'roots of all': earth, water, air, and fire. All
change is explained by the arrangement and rearrangement of these four elements. The various forms of life
are caused by an appropriate mixture of elements. For example, growth in plants is explained by the natural
downward movement of earth and the natural upward movement of fire.[6]
Democritus (460 B.C.), the disciple of Leucippus, thought that the essential characteristic of life is
having a soul (psychê). In common with other ancient writers, he used the term to mean the principle of living
things that causes them to function as a living thing. He thought the soul was composed of fire atoms,
because of the apparent connection between life and heat, and because fire moves.[7] He also suggested
that humans originally lived like animals, gradually developing communities to help one another, originating
language, and developing crafts and agriculture.[8]
In the scientific revolution of the seventeenth century, mechanistic ideas were revived by
philosophers like Descartes.

[edit] Hylomorphism
Hylomorphism is the theory (originating with Aristotle (322 BC)) that all things are a combination of
matter and form. Aristotle was one of the first ancient writers to approach the subject of life in a scientific way.
Biology was one of his main interests, and there is extensive biological material in his extant writings.
According to him, all things in the material universe have both matter and form. The form of a living thing is its
soul (Greek 'psyche', Latin 'anima'). There are three kinds of souls: the 'vegetative soul' of plants, which
causes them to grow and decay and nourish themselves, but does not cause motion and sensation; the
'animal soul' which causes animals to move and feel; and the rational soul which is the source of
consciousness and reasoning which (Aristotle believed) is found only in man.[9] Each higher soul has all the
attributes of the lower one. Aristotle believed that while matter can exist without form, form cannot exist
without matter, and therefore the soul cannot exist without the body.[10]
Consistent with this account is a teleological explanation of life. A teleological explanation accounts
for phenomena in terms of their purpose or goal-directedness. Thus, the whiteness of the polar bear's coat is
explained by its purpose of camouflage. The direction of causality is the other way round from materialistic
science, which explains the consequence in terms of a prior cause. Modern biologists now reject this
functional view in terms of a material and causal one: biological features are to be explained not by looking
forward to future optimal results, but by looking backwards to the past evolutionary history of a species,
which led to the natural selection of the features in question.

[edit] Vitalism
Vitalism is the belief that the life-principle is essentially immaterial. This originated with Stahl (17th
century), and held sway until the middle of the nineteenth century. It appealed to philosophers such as Henri
Bergson, Nietzsche, Wilhelm Dilthey, anatomists like Bichat, and chemists like Liebig.
Vitalism underpinned the idea of a fundamental separation of organic and inorganic material, and the
belief that organic material can only be derived from living things. This was disproved in 1828 when Wöhler
prepared urea from inorganic materials. This so-called Wöhler synthesis is considered the starting point of
modern organic chemistry. It is of great historical significance because for the first time an organic compound
was produced from inorganic reactants.
Later, Helmholtz, anticipated by Mayer, demonstrated that no energy is lost in muscle movement,
suggesting that there were no vital forces necessary to move a muscle. These empirical results led to the
abandonment of scientific interest in vitalistic theories, although the belief lingered on in non-scientific
theories such as homeopathy, which interprets diseases and sickness as caused by disturbances in a
hypothetical vital force or life force.

[edit] Definitions
It is still a challenge for scientists and philosophers to define life in unequivocal terms.[11][12][13]
Defining life is difficult —in part— because life is a process, not a pure substance.[14] Any definition must be
sufficiently broad to encompass all life with which we are familiar, and it should be sufficiently general that,
with it, scientists would not miss life that may be fundamentally different from earthly life.[15]

[edit] Biology
Since there is no unequivocal definition of life, the current understanding is descriptive, where life is a
characteristic of organisms that exhibit all or most of the following phenomena:[14][16][17]
1. Homeostasis: Regulation of the internal environment to maintain a constant state; for
example, electrolyte concentration or sweating to reduce temperature.
2. Organization: Being structurally composed of one or more cells, which are the basic units of
life.
3. Metabolism: Transformation of energy by converting chemicals and energy into cellular
components (anabolism) and decomposing organic matter (catabolism). Living things require energy
to maintain internal organization (homeostasis) and to produce the other phenomena associated with
life.
 4. Growth: Maintenance of a higher rate of anabolism than catabolism. A growing organism
increases in size in all of its parts, rather than simply accumulating matter.
5. Adaptation: The ability to change over a period of time in response to the environment. This
ability is fundamental to the process of evolution and is determined by the organism's heredity as well
as the composition of metabolized substances, and external factors present.
6. Response to stimuli: A response can take many forms, from the contraction of a unicellular
organism to external chemicals, to complex reactions involving all the senses of multicellular
organisms. A response is often expressed by motion, for example, the leaves of a plant turning
toward the sun (phototropism) and by chemotaxis.
7. Reproduction: The ability to produce new individual organisms, either asexually from a single
parent organism, or sexually from two parent organisms.
Proposed
To reflect the minimum phenomena required, some have proposed other biological definitions of life:
• Living things are systems that tend to respond to changes in their environment, and inside
themselves, in such a way as to promote their own continuation.[17]
• A network of inferior negative feedbacks (regulatory mechanisms) subordinated to a superior
positive feedback (potential of expansion, reproduction).[18]
• A systemic definition of life is that living things are self-organizing and autopoietic (self-
producing). Variations of this definition include Stuart Kauffman's definition as an autonomous agent
or a multi-agent system capable of reproducing itself or themselves, and of completing at least one
thermodynamic work cycle.[19]
• Life is a self-sustained chemical system capable of undergoing Darwinian evolution.[20]
 • Things with the capacity for metabolism and motion.[14]
Viruses
Viruses are most often considered replicators rather than forms of life. They have been described as
"organisms at the edge of life",[21] since they possess genes, evolve by natural selection,[22] and replicate
by creating multiple copies of themselves through self-assembly. However, viruses do not metabolize and
require a host cell to make new products. Virus self-assembly within host cells has implications for the study
of the origin of life, as it may support the hypothesis that life could have started as self-assembling organic
molecules.[23][24]

[edit] Biophysics
Biophysicists have also commented on the nature and qualities of life forms—notably that they
function on negative entropy.[25][26] In more detail, according to physicists such as John Bernal, Erwin
Schrödinger, Eugene Wigner, and John Avery, life is a member of the class of phenomena which are open or
continuous systems able to decrease their internal entropy at the expense of substances or free energy taken
in from the environment and subsequently rejected in a degraded form (see: entropy and life).[27][28][29]

[edit] Living systems theories

Some scientists have proposed in the last few decades that a general living systems theory is
required to explain the nature of life.[30] Such general theory, arising out of the ecological and biological
sciences, attempts to map general principles for how all living systems work. Instead of examining
phenomena by attempting to break things down into component parts, a general living systems theory
explores phenomena in terms of dynamic patterns of the relationships of organisms with their environment.
[31]
Gaia hypothesis
The idea that the Earth is alive is probably as old as humankind, but the first public expression of it as
a fact of science was by a Scottish scientist, James Hutton. In 1785 he stated that the Earth was a
superorganism and that its proper study should be physiology. Hutton is rightly remembered as the father of
geology, but his idea of a living Earth was forgotten in the intense reductionism of the nineteenth century.[32]
The Gaia hypothesis, originally proposed in the 1960s by scientist James Lovelock,[33][34] explores the idea
that the life on Earth functions as a single organism which actually defines and maintains environmental
conditions necessary for its survival.[35]
Nonfractionability
Robert Rosen (1991) built on the assumption that the explanatory powers of the mechanistic
worldview cannot help understand the realm of living systems. One of several important clarifications he
made was to define a system component as "a unit of organization; a part with a function, i.e., a definite
relation between part and whole." From this and other starting concepts, he developed a "relational theory of
systems" that attempts to explain the special properties of life. Specifically, he identified the
"nonfractionability of components in an organism" as the fundamental difference between living systems and
'biological machines.'[36]
Life as a property of ecosystems
A systems view of life treats environmental fluxes and biological fluxes together as a "reciprocity of
influence",[37] and a reciprocal relation with environment is arguably as important for understanding life as it
is for understanding ecosystems. As Harold J. Morowitz (1992) explains it, life is a property of an ecological
system rather than a single organism or species.[38] He argues that an ecosystemic definition of life is
preferable to a strictly biochemical or physical one. Robert Ulanowicz (2009) also highlights mutualism as the
key to understand the systemic, order-generating behavior of life and ecosystems.[39]

[edit] Origin of life

Main article: Origin of life
For religious beliefs about the creation of life, see creation myth.
Evidence suggests that life on Earth has existed for about 3.7 billion years.[40] All known life forms
share fundamental molecular mechanisms, and based on these observations, theories on the origin of life
attempt to find a mechanism explaining the formation of a primordial single cell organism from which all life
originates. There are many different hypotheses regarding the path that might have been taken from simple
organic molecules via pre-cellular life to protocells and metabolism. Many models fall into the "genes-first"
category or the "metabolism-first" category, but a recent trend is the emergence of hybrid models that
combine both categories.[41]
There is no scientific consensus as to how life originated and all proposed theories are highly
speculative. However, most currently accepted scientific models build in one way or another on the following
hypotheses:
• The Miller-Urey experiment, and the work of Sidney Fox, suggest that that conditions on the
primitive Earth may have favored chemical reactions that synthesized some amino acids and other
organic compounds from inorganic precursors.
 • Phospholipids spontaneously form lipid bilayers, the basic structure of a cell membrane.
Life as we know it today synthesizes proteins, which are polymers of amino acids using instructions
encoded by cellular genes—which are polymers of deoxyribonucleic acid (DNA). Protein synthesis also
entails intermediary ribonucleic acid (RNA) polymers. One possibility is that genes came first[42] and then
proteins. Another possibility is that proteins came first[43] and then genes. However, because genes are
required to make proteins, and proteins are required to make genes, the problem of considering which came
first is like that of the chicken or the egg. Most scientists have adopted the hypothesis that because DNA and
proteins function together so intimately, it's unlikely that they arose independently.[44] Therefore, many
scientists consider the possibility, apparently first suggested by Francis Crick,[45] that the first life was based
on the DNA-protein intermediary: RNA.[44] In fact, RNA has the DNA-like properties of information storage
and replication and the catalytic properties of some proteins. Crick and others actually favored the RNA-first
hypothesis[46] even before the catalytic properties of RNA had been demonstrated by Thomas Cech.[47]
A significant issue with the RNA-first hypothesis is that experiments designed to synthesize RNA
from simple precursors have not been nearly as successful as the Miller-Urey experiments that synthesized
other organic molecules from inorganic precursors. One reason for the failure to create RNA in the laboratory
is that RNA precursors are very stable and don't react with each other under ambient conditions. However,
the successful synthesis of certain RNA molecules under conditions hypothesized to exist prior to life on
Earth has been achieved by adding alternative precursors in a specified order with the precursor phosphate
present throughout the reaction.[48] This study makes the RNA-first hypothesis more plausible to many
scientists.[49]
Recent experiments have demonstrated true Darwinian evolution of unique RNA enzymes
(ribozymes) made up of two separate catalytic components that replicate each other in vitro.[50] In describing
this work from his laboratory, Gerald Joyce stated: "This is the first example, outside of biology, of
evolutionary adaptation in a molecular genetic system."[51] Such experiments make the possibility of a
primordial RNA World even more attractive to many scientists.

[edit] Conditions for life

The diversity of life on Earth today is a result of the dynamic interplay between genetic opportunity,
metabolic capability, environmental challenges,[52] and symbiosis.[53][54][55] For most of its existence,
Earth's habitable environment has been dominated by microorganisms and subjected to their metabolism
and evolution. As a consequence of such microbial activities on a geologic time scale, the physical-chemical
environment on Earth has been changing, thereby determining the path of evolution of subsequent life.[52]
For example, the release of molecular oxygen by cyanobacteria as a by-product of photosynthesis induced
fundamental, global changes in the Earth's environment. The altered environment, in turn, posed novel
evolutionary challenges to the organisms present, which ultimately resulted in the formation of our planet's
major animal and plant species. Therefore this 'co-evolution' between organisms and their environment is
apparently an inherent feature of living systems.[52]

[edit] Range of tolerance

The inert components of an ecosystem are the physical and chemical factors necessary for life –
energy (sunlight or chemical energy), water, temperature, atmosphere, gravity, nutrients, and ultraviolet solar
radiation protection.[56] In most ecosystems the conditions vary during the day and often shift from one
season to the next. To live in most ecosystems, then, organisms must be able to survive a range of
conditions, called 'range of tolerance'.[57] Outside of that are the 'zones of physiological stress', where the
survival and reproduction are possible but not optimal. Outside of these zones are the 'zones of intolerance',
where life for that organism is implausible. It has been determined that organisms that have a wide range of
tolerance are more widely distributed than organisms with a narrow range of tolerance.[57]

[edit] Extremophiles

Deinococcus radiodurans can resist radiation exposure.

Main article: Extremophile
 To survive, some microorganisms can assume forms that enable them to withstand freezing,
complete desiccation, starvation, high-levels of radiation exposure, and other physical or chemical
challenges. Furthermore, some microorganisms can survive exposure to such conditions for weeks, months,
years, or even centuries.[52] Extremophiles are microbial life forms that thrive outside the ranges life is
commonly found in. They also excel at exploiting uncommon sources of energy. While all organisms are
composed of nearly identical molecules, evolution has enabled such microbes to cope with this wide range of
physical and chemical conditions. Characterization of the structure and metabolic diversity of microbial
communities in such extreme environments is ongoing. An understanding of the tenacity and versatility of life
on Earth, as well as an understanding of the molecular systems that some organisms utilize to survive such
extremes, will provide a critical foundation for the search for life beyond Earth.[52]

[edit] Classification of life

Main article: Biological classification
 The hierarchy of biological classification's eight major taxonomic ranks, which is an example of
definition by genus and differentia. Life is divided into domains, which are subdivided into further groups.
Intermediate minor rankings are not shown.
Traditionally, people have divided organisms into the classes of plants and animals, based mainly on
their ability of movement. The first known attempt to classify organisms was conducted by the Greek
philosopher Aristotle (384-322 BC). He classified all living organisms known at that time as either a plant or
an animal. Aristotle distinguished animals with blood from animals without blood (or at least without red
blood), which can be compared with the concepts of vertebrates and invertebrates respectively. He divided
the blooded animals into five groups: viviparous quadrupeds (mammals), birds, oviparous quadrupeds
(reptiles and amphibians), fishes and whales. The bloodless animals were also divided into five groups:
cephalopods, crustaceans, insects (which also included the spiders, scorpions, and centipedes, in addition to
what we now define as insects), shelled animals (such as most molluscs and echinoderms) and "zoophytes".
Though Aristotle's work in zoology was not without errors, it was the grandest biological synthesis of the time
and remained the ultimate authority for many centuries after his death.[58]
The exploration of the American continent revealed large numbers of new plants and animals that
needed descriptions and classification. In the latter part of the 16th century and the beginning of the 17th,
careful study of animals commenced and was gradually extended until it formed a sufficient body of
knowledge to serve as an anatomical basis for classification.
In the late 1740s, Carolus Linnaeus introduced his method, still used, to formulate the scientific name
of every species.[59] Linnaeus took every effort to improve the composition and reduce the length of the
many-worded names by abolishing unnecessary rhetoric, introducing new descriptive terms and defining
their meaning with an unprecedented precision. By consistently using his system, Linnaeus separated
nomenclature from taxonomy. This convention for naming species is referred to as binomial nomenclature.
The fungi were originally treated as plants. For a short period Linnaeus had placed them in the taxon
Vermes in Animalia. He later placed them back in Plantae. Copeland classified the Fungi in his Protoctista,
thus partially avoiding the problem but acknowledged their special status.[60] The problem was eventually
solved by Whittaker, when he gave them their own kingdom in his five-kingdom system. As it turned out, the
fungi are more closely related to animals than to plants.[61]
As new discoveries enabled us to study cells and microorganisms, new groups of life were revealed,
and the fields of cell biology and microbiology were created. These new organisms were originally described
separately in protozoa as animals and protophyta/thallophyta as plants, but were united by Haeckel in his
kingdom protista, later the group of prokaryotes were split off in the kingdom Monera, eventually this kingdom
would be divided in two separate groups, the Bacteria and the Archaea, leading to the six-kingdom system
and eventually to the current three-domain system.[62] The classification of eukaryotes is still controversial,
with protist taxonomy especially problematic.[63]
As microbiology, molecular biology and virology developed, non-cellular reproducing agents were
discovered, such as viruses and viroids. Sometimes these entities are considered to be alive but others
argue that viruses are not living organisms since they lack characteristics such as cell membrane,
metabolism and do not grow or respond to their environments. Viruses can however be classed into
"species" based on their biology and genetics but many aspects of such a classification remain controversial.
[64]
 Since the 1960s a trend called cladistics has emerged, arranging taxa in an evolutionary or
phylogenetic tree. It is unclear, should this be implemented, how the different codes will coexist.[65]

Linnaeus Haeckel Chatton Copeland Whittaker Woese et al.

1735[66] 1866[67] 1925[68][69] 1938[60][70] 1969[71] 1977[72][73]
2 kingdoms 3 kingdoms 2 empires 4 kingdoms 5 kingdoms 6 kingdoms

Eubacteria
Prokaryota Monera Monera
(not Archaebacteria
Protista
treated)
Protista Protista
Protoctista

Eukaryota Fungi Fungi

Vegetabilia Plantae
Plantae Plantae Plantae

Animalia Animalia Animalia Animalia Animalia

[edit] Extraterrestrial life
Main articles: Extraterrestrial life and astrobiology

Panspermia hypothesis
Earth is the only planet in the universe known to harbour life. The Drake equation, which relates the
number of extraterrestrial civilizations in our galaxy with which we might come in contact, has been used to
discuss the probability of life elsewhere, but scientists disagree on many of the values of variables in this
equation. Depending on those values, the equation may either suggest that life arises frequently or
infrequently.
The region around a main sequence star that could support Earth-like life on an Earth-like planet is
known as the habitable zone. The inner and outer radii of this zone vary with the luminosity of the star, as
does the time interval during which the zone will survive. Stars more massive than the Sun have a larger
habitable zone, but will remain on the main sequence for a shorter time interval during which life can evolve.
Small red dwarf stars have the opposite problem, compounded with higher levels of magnetic activity and the
effects of tidal locking from close orbits. Hence, stars in the intermediate mass range such as the Sun may
possess the optimal conditions for Earth-like life to develop. The location of the star within a galaxy may also
have an impact on the likelihood of life forming.
Panspermia, also called exogenesis, is a hypothesis proposing that life originated elsewhere in the
universe and was subsequently transferred to Earth in the form of spores perhaps via meteorites, comets or
cosmic dust. However, this hypothesis does not help explain the ultimate origin of life.

[edit] Death
Main article: Death
Death is the permanent termination of all vital functions or life processes in an organism or cell.[75]
[76] After death, the remains of an organism become part of the biogeochemical cycle. Organisms may be
consumed by a predator or a scavenger and leftover organic material may then be further decomposed by
detritivores, organisms which recycle detritus, returning it to the environment for reuse in the food chain.
One of the challenges in defining death is in distinguishing it from life. Death would seem to refer to either the
moment at which life ends, or when the state that follows life begins.[77] However, determining when death
has occurred requires drawing precise conceptual boundaries between life and death. This is problematic,
however, because there is little consensus over how to define life. The nature of death has for millennia been
a central concern of the world's religious traditions and of philosophical inquiry. Many religions maintain faith
in either some kind of afterlife, reincarnation, or resurrection.

[edit] Extinction
Main article: Extinction
 Extinction is the gradual process by which a group of taxa or species dies out, reducing biodiversity.
[78] The moment of extinction is generally considered to be the death of the last individual of that species.
Because a species' potential range may be very large, determining this moment is difficult, and is usually
done retrospectively after a period of apparent absence. Species become extinct when they are no longer
able to survive in changing habitat or against superior competition. Over the history of the Earth, over 99% of
all the species that have ever lived have gone extinct,[79] however, mass extinctions may have accelerated
evolution by providing opportunities for new groups of organisms to diversify.[80]

[edit] Fossils
Main article: Fossil
Fossils are the preserved remains or traces of animals, plants, and other organisms from the remote
past. The totality of fossils, both discovered and undiscovered, and their placement in fossil-containing rock
formations and sedimentary layers (strata) is known as the fossil record. Such a preserved specimen is
called a "fossil" if it is older than the arbitrary date of 10,000 years ago.[81] Hence, fossils range in age from
the youngest at the start of the Holocene Epoch to the oldest from the Archaean Eon, a few billion years old.

[edit] See also

• Alpha taxonomy • Kingdom (biology)
• Artificial life • Life on Mars
• Biological immortality • Meaning of life
• Biology - the study of life • Nature
• Carbon-based life • Non-cellular life
 • Cellular life • Organic life
• Conway's Game of Life • Organism
• Death • Origin of life
• Entropy and life • Personal life
• Evolutionary history of life • Phylogenetics
• Extraterrestrial life • Prehistoric life
• Extremophile • Prion
• Gaia hypothesis • Quality of life
• Genetics • Silicon-based life
• Genetic engineering • Synthetic life
• Hierarchy of life
• Hypothetical types of
biochemistry

[edit] References
1. ^ a b Koshland Jr, Daniel E. (March 22, 2002). "The Seven Pillars of Life". Science 295.
(5563): 2215–2216. doi:10.1126/science.1068489. PMID 11910092.
http://www.sciencemag.org/cgi/content/full/295/5563/2215. Retrieved 2009-05-25.
2. ^ The American Heritage Dictionary of the English Language, 4th edition, published by
Houghton Mifflin Company, via Answers.com:
 • "The property or quality that distinguishes living organisms from dead organisms and
inanimate matter, manifested in functions such as metabolism, growth, reproduction, and
response to stimuli or adaptation to the environment originating from within the organism."
• "The characteristic state or condition of a living organism."
3. ^ Definition of inanimate. WordNet Search by Princeton University.
4. ^ "Merriam-Webster Dictionary". Merriam-Webster Dictionary. http://www.merriam-
webster.com/dictionary/life. Retrieved 2009-06-21.
5. ^ "organism". Chambers 21st Century Dictionary (online ed.). 1999.
6. ^ SEP
7. ^ SEP
8. ^ Ibidem
9. ^ Aristotle, De Anima, Book II
10.^ Introduction to Ancient Philosophy, Don Marietta, p.104.
11.^ Defining Life : Astrobiology Magazine - earth science - evolution distribution Origin of life
universe - life beyond
12.^ Defining Life, Explaining Emergence
13.^ "Can We Define Life". Colorado Arts & Sciences. 2009.
http://artsandsciences.colorado.edu/magazine/2009/03/can-we-define-life/. Retrieved 2009-06-22.
14.^ a b c McKay, Chris P. (September 14, 2004). "What Is Life—and How Do We Search for It in
Other Worlds?". PLoS Biol. 2 (2(9)): 302. doi:10.1371/journal.pbio.0020302. PMID 15367939.
PMC 516796. http://www.ncbi.nlm.nih.gov/pmc/articles/PMC516796/?tool=pubmed. Retrieved 2010-
02-02.
 15.^ Nealson KH, Conrad PG (December 1999). "Life: past, present and future". Philos. Trans.
R. Soc. Lond., B, Biol. Sci. 354 (1392): 1923–39. doi:10.1098/rstb.1999.0532. PMID 10670014.
PMC 1692713. http://journals.royalsociety.org/content/7r10hqn3rp1g1vag/fulltext.pdf.
16.^ Davison, Paul G.. "How to Define Life". The University of North Alabama.
http://www2.una.edu/pdavis/BI%20101/Overview%20Fall%202004.htm. Retrieved 2008-10-17.
17.^ a b Witzany, G. (2007). The Logos of the Bios 2. Bio-Communication. Helsinki, Umweb.
18.^ Korzeniewski, Bernard (2001). "Cybernetic formulation of the definition of life". Journal of
Theoretical Biology. 2001 April 7. 209 (3) pp. 275–86.
19.^ 2004, "Autonomous Agents", in John D. Barrow, P.C.W. Davies, and C.L. Harper Jr., eds.,
Science and Ultimate Reality: Quantum Theory, Cosmology, and Complexity, Cambridge University
Press.
20.^ Gerald Francis Joyce, "The RNA World: Life Before DNA and Protein".
21.^ Rybicki EP (1990) "The classification of organisms at the edge of life, or problems with
virus systematics." S Aft J Sci 86:182–186
22.^ Holmes EC (October 2007). "Viral evolution in the genomic age". PLoS Biol. 5 (10): e278.
doi:10.1371/journal.pbio.0050278. PMID 17914905. PMC 1994994.
http://biology.plosjournals.org/perlserv/?request=get-document&doi=10.1371/journal.pbio.0050278.
Retrieved 2008-09-13.
23.^ Koonin EV, Senkevich TG, Dolja VV (2006). "The ancient Virus World and evolution of
cells". Biol. Direct 1: 29. doi:10.1186/1745-6150-1-29. PMID 16984643. PMC 1594570.
http://www.biology-direct.com/content/1//29. Retrieved 2008-09-14.
24.^ Rybicki, Ed (November 1997). "Origins of Viruses".
http://www.mcb.uct.ac.za/tutorial/virorig.html#Virus%20Origins. Retrieved 2009-04-12.
 25.^ Schrödinger, Erwin (1944). What is Life?. Cambridge University Press. ISBN 0-521-42708-
8.
26.^ Margulis, Lynn; Sagan, Dorion (1995). What is Life?. University of California Press. ISBN 0-
520-22021-8.
27.^ Lovelock, James (2000). Gaia – a New Look at Life on Earth. Oxford University Press.
ISBN 0-19-286218-9.
28.^ Avery, John (2003). Information Theory and Evolution. World Scientific. ISBN 9812383999.
29.^ "BIOPHYSICS: DEFINITION OF LIFE AND BRIEF EXPLANATION OF EACH TERM".
Biology Cabinet. September 29, 2006. http://biocab.org/Exobiology.html#anchor_41. Retrieved 2009-
07-22.
30.^ Woodruff, T. Sullivan; John Baross (October 8, 2007). Planets and Life: The Emerging
Science of Astrobiology. Cambridge University Press. Cleland and Chyba wrote a chapter in Planets
and Life: "In the absence of such a theory, we are in a position analogous to that of a 16th-century
investigator trying to define 'water' in the absence of molecular theory." [...] "Without access to living
things having a different historical origin, it is difficult and perhaps ultimately impossible to formulate
an adequately general theory of the nature of living systems".
31.^ "Patterns, Flows, and Interrelationship". 2002.
http://www.mollyyoungbrown.com/systems_article.htm. Retrieved 2009-06-27.
32.^ GAIA - A new look at life on Earth. James Lovelock 1979. pp. 10. Oxford University Press.
ISBN 0-19-286030-5.
33.^ Lovelock, J.E. (1965). "A physical basis for life detection experiments". Nature 207 (7): 568–
570. doi:10.1038/207568a0.
34.^ Geophysiology
 35.^ GAIA - A new look at life on Earth. James Lovelock. 1979. Oxford University Press. ISBN
0-19-286030-5.
36.^ Life Itself: A Comprehensive Inquiry into the Nature, Origin, and Fabrication of Life . Rosen,
Robert. November, 1991. ISBN 978-0-231-07565-7
37.^ "The Ecosystemic Life Hypothesis". Bulletin of the Ecological Society of America. April
2002. http://www.calresco.org/fiscus/esl.htm. Retrieved 2009-08-28.
38.^ Morowitz, Harold J. (1992) "Beginnings of Cellular Life: Metabolism Recapitulates
Biogenesis". Yale University Press. ISBN 0-300-05483-1
39.^ A Third Window: Natural Life Beyond Newton and Darwin , Templeton Foundation Press
(2009) ISBN 1-59947-154-X
40.^ "History of life through time". University of California Museum of Paleontology.
41.^ Coveney, Peter V.; Philip W. Fowler. "Modelling biological complexity: a physical scientist's
perspective". Journal of the Royal Society Interface. 2005. 2 (4) pp. 267–280.
doi:10.1098/rsif.2005.0045
42.^ Senapathy, Periannan, Independent Birth of Organisms, Madison, WI. Genome Press,
1994.
43.^ Eigen, Manfred, Steps Towards Life: A Perspective on Evolution (German edition, 1987),
Oxford University Press, 1992. p 31.
44.^ a b Barazesh, Solmaz, How RNA Got Started: Scientists Look for the Origins of Life,
Science News, May 13, 2009.
45.^ Watson, James D., Prologue: Early Speculations and Facts about RNA Templates, p xv-
xxiii, The RNA World, R.F. Gesteland and J.F. Atkins, Eds., Cold Spring Harbor Laboratory Press,
1993.
46.^ Gilbert, Walter, The RNA world, p 618 v 319, Nature, 1986.
 47.^ Cech, Thomas R., A model for the RNA-catalyzed replication of RNA, p 4360-4363 v 83,
Proc. Nat. Acad. Sci., USA, 1986.
48.^ Powner, Matthew W., Béatrice Gerland and John D. Sutherland, Synthesis of activated
pyrimidine ribonucleotides in prebiotically plausible conditions, Nature 459, 239-242 (14 May 2009).
49.^ Szostak, Jack W., Origins of life: Systems chemistry on early Earth, Nature 459, 171-172
(14 May 2009).
50.^ Lincoln, Tracey A. and Gerald F. Joyce, Self-Sustained Replication of an RNA Enzyme,
Science 27 February 2009:Vol. 323, No. 5918, pp. 1229-1232, DOI: 10.1126/science.1167856.
51.^ Joyce, Gerald F., Evolution in an RNA World, Cold Spring Harb Symp Quant Biol
sqb.2009.74.004; Published in Advance August 10, 2009, doi:10.1101/sqb.2009.74.004.
52.^ a b c d e Rothschild, Lynn (September, 2003). "Understand the evolutionary mechanisms
and environmental limits of life". NASA. http://astrobiology.arc.nasa.gov/roadmap/g5.html. Retrieved
2009-07-13.
53.^ "Symbiosis and the origin of life". Origins of Life and Evolution of Biospheres 8 (1): 39–53.
April, 1977. doi:10.1007/BF00930938. http://www.springerlink.com/content/n10p775113175l67/.
Retrieved 2010-02-22.
54.^ Margulis, Lynn (2001). The Symbiotic Planet: A New Look at Evolution. London, England:
Orion Books Ltd.. ISBN 0-75380-785-8.
55.^ Douglas J. Futuyma; Janis Antonovics (1992, Volume 8). Oxford surveys in evolutionary
biology: Symbiosis in evolution. London, England: Oxford University Press. pp. 347–374. ISBN 0-19-
507623-0.
56.^ "Essential requirements for life". CMEX-NASA.
http://cmapsnasacmex.ihmc.us/servlet/SBReadResourceServlet?
rid=1025200161109_2045745605_1714&partName=htmltext. Retrieved 2009-07-14.
 57.^ a b Chiras, Daniel C. (2009). Environmental Science – Creating a Sustainable Future .
58.^ "Aristotle -biography". University of California Museum of Paleontology.
http://www.ucmp.berkeley.edu/history/aristotle.html. Retrieved 2008-10-20.
59.^ Knapp S, Lamas G, Lughadha EN, Novarino G (April 2004). "Stability or stasis in the
names of organisms: the evolving codes of nomenclature". Philosophical transactions of the Royal
Society of London. Series B, Biological sciences 359 (1444): 611–22. doi:10.1098/rstb.2003.1445.
PMID 15253348. PMC 1693349. http://journals.royalsociety.org/openurl.asp?
genre=article&issn=0962-8436&volume=359&issue=1444&spage=611.
60.^ a b Copeland, H. F. (1938). "The Kingdoms of Organisms". Quarterly Review of Biology 13
(4): 383. doi:10.1086/394568. http://links.jstor.org/sici?sici=0033-
5770(193812)13%3A4%3C383%3ATKOO%3E2.0.CO%3B2-K.
61.^ Whittaker RH (January 1969). "New concepts of kingdoms or organisms. Evolutionary
relations are better represented by new classifications than by the traditional two kingdoms". Science
163 (863): 150–60. doi:10.1126/science.163.3863.150. PMID 5762760.
62.^ a b Woese C, Kandler O, Wheelis M (1990). "Towards a natural system of organisms:
proposal for the domains Archaea, Bacteria, and Eucarya.". Proc Natl Acad Sci U S A 87 (12): 4576–
9. doi:10.1073/pnas.87.12.4576. PMID 2112744. PMC 54159.
http://www.pnas.org/cgi/reprint/87/12/4576.
63.^ Adl SM, Simpson AG, Farmer MA, et al. (2005). "The new higher level classification of
eukaryotes with emphasis on the taxonomy of protists". J. Eukaryot. Microbiol. 52 (5): 399–451.
doi:10.1111/j.1550-7408.2005.00053.x. PMID 16248873.
64.^ Van Regenmortel MH (January 2007). "Virus species and virus identification: past and
current controversies". Infection, genetics and evolution : journal of molecular epidemiology and
evolutionary genetics in infectious diseases 7 (1): 133–44. doi:10.1016/j.meegid.2006.04.002.
PMID 16713373.
65.^ Pennisi E (March 2001). "Taxonomy. Linnaeus's last stand?". Science (New York, N.Y.)
291 (5512): 2304–7. doi:10.1126/science.291.5512.2304. PMID 11269295.
http://www.sciencemag.org/cgi/pmidlookup?view=long&pmid=11269295.
66.^ C. Linnaeus (1735). Systemae Naturae, sive regna tria naturae, systematics proposita per
classes, ordines, genera & species.
67.^ E. Haeckel (1866). Generelle Morphologie der Organismen. Reimer, Berlin.
68.^ É. Chatton (1925). "Pansporella perplexa. Réflexions sur la biologie et la phylogénie des
protozoaires". Ann. Sci. Nat. Zool 10-VII: 1–84.
69.^ É. Chatton (1937). Titres et Travaux Scientifiques (1906–1937). Sette, Sottano, Italy.
70.^ H. F. Copeland (1956). The Classification of Lower Organisms. Palo Alto: Pacific Books.
71.^ Whittaker RH (January 1969). "New concepts of kingdoms of organisms". Science 163
(863): 150–60. doi:10.1126/science.163.3863.150. PMID 5762760.
72.^ C. R. Woese, W. E. Balch, L. J. Magrum, G. E. Fox and R. S. Wolfe (August 1977). "An
ancient divergence among the bacteria". Journal of Molecular Evolution 9 (4): 305–311.
doi:10.1007/BF01796092. PMID 408502.
73.^ Woese CR, Fox GE (November 1977). "Phylogenetic structure of the prokaryotic domain:
the primary kingdoms". Proc. Natl. Acad. Sci. U.S.A. 74 (11): 5088–90. PMID 270744.
74.^ Cavalier-Smith, T. (2004), "Only six kingdoms of life", Proc. R. Soc. Lond. B 271: 1251–62,
doi:10.1098/rspb.2004.2705, http://www.cladocera.de/protozoa/cavalier-smith_2004_prs.pdf,
retrieved 2010-04-29
75.^ "Definition of death.". Archived from the original on 2009-10-31.
http://www.webcitation.org/5kwsdvU8f.
 76.^ Defining of death.
77.^ Encyclopedia of Death and Dying
78.^ "Extinction - definition.". Archived from the original on 2009-10-31.
http://www.webcitation.org/5kwseRB80.
79.^ What is an extinction?
80.^ Van Valkenburgh, B. (1999). "Major patterns in the history of carnivorous mammals".
Annual Review of Earth and Planetary Sciences 26: 463–493. doi:10.1146/annurev.earth.27.1.463.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.27.1.463.
81.^ FAQs - San Diego Natural History Museum

[edit] Further reading

• Kauffman, Stuart. The Adjacent Possible: A Talk with Stuart Kauffman
• Nealson KH, Conrad PG (December 1999). "Life: past, present and future". Philos. Trans. R.
Soc. Lond., B, Biol. Sci. 354 (1392): 1923–39. doi:10.1098/rstb.1999.0532. PMID 10670014.
PMC 1692713. http://journals.royalsociety.org/content/7r10hqn3rp1g1vag/.
• Walker, Martin G. LIFE! Why We Exist...And What We Must Do to Survive Dog Ear
Publishing, 2006, ISBN 1-59858-243-7

[edit] External links

Wikimedia Commons has media related to: Tree of life
 Wikiquote has a collection of quotations related to: Life

Look up life or living in Wiktionary, the free dictionary.

Wikispecies has information related to: The Taxonomy of Life

• Wikispecies - a free directory of life

• "The Adjacent Possible: A Talk with Stuart Kauffman"
• Stanford Encyclopedia of Philosophy entry
• Life under extreme conditions An in depth look at how life can form under the most extreme
conditions and circumstaces.

[show]
v•d•e
Elements of nature

Universe Space · Time · Matter · Energy

 Earth Earth science · Geology · History of the Earth · Geological history of Earth ·
Future of the Earth · Structure of the Earth · Plate tectonics

Weather Earth's atmosphere · Climate · Meteorology

Environment Ecology · Ecosystem · Wilderness

Hierarchy of life · Origin of life · Life on Earth · Eukaryota (Plants/Flora,

Life Animals/Fauna, Fungi, Protista) · Prokaryote (Archaea, Bacteria) · Virus · Evolutionary
history of life · Biology

Category · Portal

[show]
v•d•e
Hierarchy of life

Biosphere > Ecosystem > Community (Biocoenosis) > Population > Organism > Organ system
> Organ > Tissue > Cell > Organelle > Molecule (Macromolecule · Biomolecule) > Atom
 [show]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

[show]
v•d•e
Taxonomic ranks
Magnorder

Domain/Super Superphylum/
Superclass Superorder Superfam
kingdom Superdivision
 Phylum/Divisi
Kingdom Class Legion Order Family
on

Subkingdom Subphylum Subclass Cohort Suborder Subfamily

Infrakingdom/B
Infraphylum Infraclass Infraorder
ranch

Microphylum Parvclass Parvorder

Retrieved from "http://en.wikipedia.org/wiki/Life"

Categories: Life | Biology | Biological systems

 W000

Vie
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir Vie (homonymie).

Cet article ne cite pas suffisamment ses sources (mars 2009).
Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
bas de page. (Modifier l'article)
Les trois âges de la Vie et la Mort, peinture de Hans Baldung.
 La vie est le nom donné :
1. à l'état et aux formes auto-organisées et homéostatiques de la matière (organismes vivants)
ayant une capacité de duplication et d'évolution. Cette définition est parfois étendue à l'ensemble des
êtres vivants dans la biosphère.
2. à un phénomène empirique particulièrement important pour les humains (qui sont eux-
mêmes vivants et pour qui les autres êtres vivants ont une place particulière), mais qui ne se laisse
pas facilement définir (cf. infra). Ce phénomène s'oppose à la notion de matière inerte ou inanimé qui
s'articule avec la notion de mort ;
3. à la durée qui sépare la naissance de la mort ;
4. au contenu en événements ou en actions de cette étendue temporelle, pour un humain ;
5. à l'approche harmonieuse des relations humaines (voir « question sociale »).
Une des marques de l'hominisation est l'existence de rites funéraires, et donc d'une conscience
d'une transition entre la vie et la mort. La vie est un concept primordial qui a donné lieu depuis des temps
immémoriaux à de nombreuses réflexions empiriques, philosophiques, scientifiques, etc. C'est également un
sujet de débat politique, qu'il s'agisse du traitement accordé aux êtres vivants par rapport aux humains et aux
choses inertes (cf. écologisme) ou des considérations sur le début et la fin de la vie humaine (cf. avortement,
euthanasie, « vie éternelle »), biophilie.
Ces réflexions concernent par exemple :
• la catégorie non-statique (par opposition à la matière inerte ou à l'état de mort) ;
• le concept d’évolution (passage de la matière inerte à la vie, développement et disparition
des formes vivantes, mort, création, etc.).
• la qualité de vie
 Elles sont toujours liées aux notions d'esprit et d'intelligence. Elles débouchent également sur des
réflexions sur l'étendue temporelle et spatiale de la vie (y compris dans l'univers : « vie extraterrestre »).
Elles s'interrogent à la fois sur les conditions d'apparition de la vie (phénomène unique ou au contraire très
banal) et sur la possibilité d'une vie évoluée (par comparaison à l'humanité, implicitement considérée comme
l'achèvement de l'évolution de la vie terrestre) au sein de l'univers.
La biologie est l'étude scientifique de la vie. Elle s'appuie notamment sur la chimie organique mais
certains théoriciens n'excluent pas d'adopter des définitions pouvant inclure des formes mécaniques ou
électromécaniques, et même des formes créées par l'homme hors de tout processus reproductif naturel
(« vie artificielle » ou cellule artificielle).
Sommaire
[masquer]
• 1 Science
• 1.1 Domaine du vivant
• 1.2 Définitions
• 1.3 Origine de la vie
• 1.4 La vie comme propriété d’un organisme vivant
• 1.5 Caractéristiques biologiques d’un être vivant
• 1.5.1 Caractéristiques au niveau des
activités
• 1.5.2 Caractéristiques au niveau des
structures et composés chimiques
• 1.5.3 Classification
• 1.5.4 Des organismes presque vivants
• 1.6 Autres définitions
• 2 Philosophie
• 2.1 Idéalisme et matérialisme
• 2.2 Une définition phénoménologique
• 3 Religion
• 3.1 Christianisme
• 4 Annexes
Science [modifier]
Domaine du vivant [modifier]
Le phénomène de la vie est potentiellement présent ailleurs dans l'Univers, mais n'est connu de
l'Homme que dans la biosphère terrestre. Elle est principalement concentrée dans les quelques centimètres
(ou premiers mètres) du sol, se produit surtout dans le premier mètre au-dessus du sol, ou les premiers
dizaines de mètres (mais se rencontre jusqu'à vers 10 km d'altitude) ainsi que dans les eaux douces et
marines. Mais on connait quelques formes de vies endolithiques : dans la roche, dans le pétrole ; et
extrémophiles : certaines faisant preuves d'une grande résistance (y compris au vide poussé, à la
radioactivité, à des pressions, pH ou températures extrêmes (chaud ou froid)).

Définitions [modifier]
Les expériences de Louis Pasteur ont réfuté la génération spontanée d'organismes vivants. Mais, il
n'existe pas encore de définition scientifique de la vie unanimement partagée. Toute définition doit tenir
compte de la notion de niveaux d'organisation structurels, d'émergence, d'homéostasie, d'entropie et de
métabolisme pour éviter de se retrouver dans une « zone grise ». Les définitions suivantes semblent limiter
ces zones grises :
• Selon la NASA, est vivant tout système délimité sur le plan spatial par une membrane semi-
perméable de sa propre fabrication et capable de s'auto-entretenir, ainsi que de se reproduire en
fabriquant ses propres constituants à partir d'énergie et/ou à partir d'éléments extérieurs.
• La vie est un état organisé et homéostatique de la matière.
 • Mode d’organisation de la matière générant des formes diverses, de complexités variables,
en interaction et ayant comme propriété principale de se reproduire presque à l’identique en utilisant
les matériaux et l'énergie disponibles dans leur environnement, auquel elles peuvent s’adapter.
L'expression presque à l’identique réfère aux mutations qui apparaissent lors de la réplication de
l'organisme et qui peuvent conférer un avantage adaptatif à celui-ci.

Origine de la vie [modifier]

Article détaillé : origine de la vie.
L'origine de la vie et la relation entre ses lignées majeures font l'objet de recherches incessantes,
sans cesse bouleversées par de nouvelles découvertes scientifiques, en particulier la biologie moléculaire
pour ces dernières années.
On peut distinguer trois principaux groupes, les procaryotes, les eucaryotes et les archaeas. Deux
organites symbiotiques trouvés chez les eucaryotiques, à savoir la mitochondrie animale et le chloroplaste
végétal, sont considérés comme le résultat de l'endosymbiose d'une bactérie.

La vie comme propriété d’un organisme vivant [modifier]

L’organisme vivant est l’objet d’un processus de développement, la vie, qui le conduit en général par
étapes d’un état « embryonnaire » à l’adulte et à la mort, de manière individuelle ou coloniale, libre ou fixée,
tout ou partie de sa vie.
La graine, la spore, le spermatozoïde ou l’ovule sont aussi des formes du vivant, bien qu’ils n’aient
en eux-mêmes ni la forme ni les caractéristiques des êtres vivants qu’ils vont devenir. Il est ainsi difficile
d’isoler totalement la vie d’un individu de la lignée à laquelle il appartient et de la biosphère. Le vivant nait du
vivant : nous ne connaissons pas de vivant émergeant de l'inerte, ce qui rend difficile la reconstitution des
étapes prébiotiques.

Caractéristiques biologiques d’un être vivant [modifier]

Ou, comment peut-on affirmer qu’une entité est « vivante » ?

Caractéristiques au niveau des activités [modifier]

En biologie, une entité est traditionnellement considérée comme vivante si elle présente les activités
suivantes, au moins une fois durant son existence :
1. Développement ou croissance : l’entité grandit ou mûrit jusqu’au moment où elle devient
capable de se reproduire ;
2. Métabolisme : consommation, transformation et stockage d'énergie ou de masse; croissance
en absorbant de l’énergie ou des nutriments présents dans son environnement ou en réorganisant sa
masse, par production d’énergie, de travail et rejet de déchets ;
3. Motricité externe (locomotion) ou interne (circulation) ;
4. Reproduction : pouvoir créer de façon autonome d'autres entités similaires à soi-même.
5. Réponse à des stimuli : pouvoir détecter des propriétés de son environnement et d'agir de
façon adaptée.
Discussion sur ces critères :
 • Ils ne sont pas tous satisfaits en même temps pour un individu particulier : il faut parfois
considérer la lignée ou l’espèce pour qu’ils coexistent (les hybrides stériles sont des êtres vivants) ;
• En isoler un ou deux peut conduire à des conclusions erronées : le feu (combustion)
assimilable à une digestion, car ce sont deux processus d’oxydation, ne transforme pas le feu en être
vivant ;
• Parfois, un critère manque : les virus ne grandissent pas et n'ont pas d'activité métabolique,
mais certains les considèrent comme vivants puisqu’ils peuvent contenir de l’ADN et être munis de
mécanismes (transcription d’ADN en ARN) provoquant leur reproduction dans les cellules hôtes ;
• D’autres fois encore, c’est une seule propriété qui est présente et qui se transmet à d’autres
entités, comme un mime de la fonction de reproduction (le prion est une protéine, conformée en
miroir par rapport à la protéine normale, qui transmet sa propriété pathogène aux autres protéines),
etc.

Caractéristiques au niveau des structures et composés chimiques [modifier]

D’où le besoin, éprouvé par les biologistes, de compléter ces caractéristiques pour réduire ces
ambiguïtés :
1. Les organismes vivants sont composés au moins d'une cellule, c’est-à-dire d’une membrane
fermée, séparant le milieu extérieur du milieu intérieur, qui contient le métabolisme et le matériel
génétique;
2. Les organismes vivants contiennent des molécules telles que : des hydrates de carbone, des
lipides, des acides nucléiques et des protéines, toutes à base de carbone ; mais on peut y voir une
vision biaisée parce que carbocentrique de la vie. Des formes de vie pourraient en théorie être
 fondées sur le silicium, mais celui-ci ne présente pas l’étonnante variété de formes et de propriétés
du carbone.
Selon la source d'énergie utilisée, on distingue les chimiotrophes, tirant leur énergie de molécules et
les Phototrophes, tirant leur énergie du Soleil.
Un organisme vivant est un ensemble organisé de matière qui tend à se maintenir à l'état
homéostatique par une utilisation concertée d'énergie.

Classification [modifier]
Articles connexes : Classification scientifique des espèces, Classification phylogénétique et
Classification classique.
Les organismes vivants sont classés par la phylogénétique. La première subdivision comprend trois
domaines :
• les archéobactéries ;
• les eubactéries ;
• les eucaryotes.

Des organismes presque vivants [modifier]

Il existe des entités proches des organismes vivants, qui ne sont toutefois pas considérés comme
tels, du fait d'une absence de métabolisme : les virus et les prions. Cependant, ces entités partagent avec les
organismes vivants la capacité de se répliquer, c'est-à-dire de susciter de la part de leur environnement la
production de copies d'elles-mêmes (formulation de David Deutsch) : ce sont des réplicateurs.

Autres définitions [modifier]

Pour Francisco Varela et Humberto Maturana, une entité est vivante si :
• elle peut se reproduire elle-même ;
• elle est basée sur l'eau ;
• elle produit des lipides et des protéines (?) ;
• son métabolisme est basé sur le carbone ;
• elle se réplique grâce à des acides nucléiques ;
• elle possède un système permettant de « lire » des protéines.
Cette définition a été largement utilisée par Lynn Margulis.
« Un système de rétrocontrôles négatifs inférieurs subordonnés à un rétrocontrôle positif supérieur »
(J. theor Biol. 2001)
Tom Kinch définit la vie comme un système autophage, hautement organisé, émergeant
naturellement des conditions ordinaires sur les corps planétaires et qui consiste en une population de
réplicateurs capables de muter.
Dans L'aventure du vivant, le biologiste Joël de Rosnay énumère trois propriétés fondamentales :
• L'autoconservation, qui est la capacité des organismes à se maintenir en vie par
l'assimilation, la nutrition, les réactions énergétiques de fermentation et de respiration ;
 • L'autoreproduction, leur possibilité de propager la vie ;
• L'autorégulation : les fonctions de coordination, de synchronisation et de contrôle des
réactions d'ensemble.
Il faut ajouter à ces trois propriétés la capacité des êtres vivants à évoluer.

Philosophie [modifier]
La vie est un système ordonné capable de volonté. Ce système est dépendant de l'environnement
dans lequel il évolue.

Idéalisme et matérialisme [modifier]

Deux grands groupes de définitions sont discutés depuis les débuts de la philosophie : les
conceptions idéalistes qui s’appuient sur une séparation plus ou moins nette entre la matière et la vie (cf. la
définition phénoménologique, ci-après) et les conceptions matérialistes qui supposent la vie comme une des
manifestations émergentes de la matière.
Historiquement, on peut distinguer deux thèses, sans qu'il soit possible de déterminer si l'une est
antérieure à l'autre, d'autant qu'elles peuvent faire l'objet de synthèses variées (les deux thèses cohabitant à
des degrés divers au sein de théories plus sophistiquées). On les retrouve dans la pensée grecque antique.
Selon les thèses dites dualistes, la vie est conçue comme fondamentalement différente de la
matière : il y a du vivant (spirituel) et de l'inerte (matériel et énergie) comme il y a du fer et de l'eau. La seule
difficulté, c'est de « purifier » et « d'isoler » (au sens quasiment chimique) le vivant de l'inerte, séparation
d'autant plus difficile qu'elle est, par définition, inaccessible aux méthodes exclusivement matérielles. Ces
thèses font appel à des notions diverses : l’âme, le souffle vital, l’élan vital, etc. Cette séparation a donné lieu
à diverses théories, comme celle de la génération spontanée, encore vivaces au temps de Louis Pasteur.
Selon les thèses monistes, au contraire, la vie est une manifestation de la matière, une propriété
émergente qui apparaît spontanément dans certaines conditions. Il est alors possible de faire varier la
définition de la vie selon les conditions qu'on considère comme caractéristiques, ce qui introduit des marges
de faux débats (les contradicteurs croyant discuter sur le concept de vie alors que, en adoptant des critères
différents, ils s'interdisent a priori tout accord) même si en pratique seuls les objets en marge sont sujet à
discussion (les microbes, les virus, les prions, le feu, etc.). La pensée scientifique moderne relève de ce type
de thèse, en particulier suite aux expériences de Pasteur sur la stérilisation : tant qu'on n'a pas démontré la
nécessité de postuler une dualité, il convient de s'en tenir à l'hypothèse moniste. Même si les étapes de
l’apparition de la vie, ou de l'organisation des êtres vivants, restent à expliquer, les lois chimiques connues
sont pour l'instant suffisantes.
Les recherches sur les conditions matérielles originelles de notre planète, avec l’espoir de parvenir à
croiser ces informations avec celles existant sur d’autres planètes, nous donneront peut-être un jour un ou
des scénarios convaincants du passage de la matière inerte à la vie.

Une définition phénoménologique [modifier]

Article détaillé : phénoménologie de la vie.
Le philosophe Michel Henry définit la vie d'un point de vue phénoménologique comme ce qui
possède la faculté et le pouvoir « de se sentir et de s'éprouver soi-même en tout point de son être ». Pour lui,
la vie est essentiellement force subjective et affectivité, elle consiste en une pure expérience subjective de
soi qui oscille en permanence entre la souffrance et la joie. Une « force subjective » n’est pas une force
impersonnelle, aveugle et insensible comme le sont les forces objectives que l’on rencontre dans la nature,
mais une force vivante et sensible éprouvée de l’intérieur et résultant d’un désir subjectif et d’un effort
subjectif de la volonté pour le satisfaire. Il établit également une opposition radicale entre la chair vivante
douée de sensibilité et le corps matériel, qui est par principe insensible, dans son livre Incarnation, une
philosophie de la chair.

Religion [modifier]
Christianisme [modifier]
La religion insiste sur le caractère inaliénable de la vie en tant que fruit de la création divine. Le livre
de la Genèse contient le récit de la création.
Dans les dix commandements, il est écrit qu'il est interdit de tuer. Le décalogue est en quelque sorte
un code de vie pour les Israélites et, dans un certain sens, pour les Chrétiens également.
Dans le Nouveau Testament, Jésus dit Je suis la voie, la vérité et la vie. (Jn 14, 6). L'Esprit Saint est
appelé souffle de vie. La vie surnaturelle trouve sa source dans l'union hypostatique de Dieu.
Le magistère a adressé les encycliques : Evangelium vitae et Humanae Vitae, sur le droit à la vie et,
au respect fondamental qu'il lui est dû.

Annexes [modifier]
Sur les autres projets Wikimédia :
 • Vie sur le Wiktionnaire (dictionnaire universel)
• Vie sur Wikisource (bibliothèque universelle)
• Vie sur Wikiquote (recueil de citations)

Articles connexes [modifier]

• Origines de la vie
• Histoire évolutive du vivant
• Mort - Fin de vie
• Vie après la mort
• Demi-vie
• Valeur de la vie
• Vérité Compréhension de la vie

• Portail de la médecine

• Portail de la philosophie
Ce document provient de « http://fr.wikipedia.org/wiki/Vie ».

Catégories : Vivant | Concept philosophique | [+]

 W000

Cell (biology)
From Wikipedia, the free encyclopedia

Jump to: navigation, search

 Drawing of the structure of cork as it appeared under the microscope to Robert Hooke from
Micrographia which is the origin of the word "cell" being used to describe the smallest unit of a living
organism

Cells in culture, stained for keratin (red) and DNA (green)

The cell is the functional basic unit of life. It was discovered by Robert Hooke and is the functional
unit of all known living organisms. It is the smallest unit of life that is classified as a living thing, and is often
called the building block of life.[1] Some organisms, such as most bacteria, are unicellular (consist of a single
cell). Other organisms, such as humans, are multicellular. Humans have about 100 trillion or 1014 cells; a
typical cell size is 10 µm and a typical cell mass is 1 nanogram. The largest cells are about 135 µm in the
anterior horn in the spinal cord while granule cells in the cerebellum, the smallest, can be some 4 µm and the
longest cell can reach from the toe to the lower brain stem (Pseudounipolar cells).[2] The largest known cells
are unfertilised ostrich egg cells which weigh 3.3 pounds.[3][4]
In 1835, before the final cell theory was developed, Jan Evangelista Purkyně observed small
"granules" while looking at the plant tissue through a microscope. The cell theory, first developed in 1839 by
Matthias Jakob Schleiden and Theodor Schwann, states that all organisms are composed of one or more
cells, that all cells come from preexisting cells, that vital functions of an organism occur within cells, and that
all cells contain the hereditary information necessary for regulating cell functions and for transmitting
information to the next generation of cells.[5]
The word cell comes from the Latin cellula, meaning, a small room. The descriptive term for the
smallest living biological structure was coined by Robert Hooke in a book he published in 1665 when he
compared the cork cells he saw through his microscope to the small rooms monks lived in.[6]
Contents
[hide]
• 1 Anatomy
• 1.1 Prokaryotic cells
• 1.2 Eukaryotic cells
• 2 Subcellular components
• 2.1 Membrane
• 2.2 Cytoskeleton
• 2.3 Genetic material
• 2.4 Organelles
• 3 Structures outside the cell wall
• 3.1 Capsule
• 3.2 Flagella
• 3.3 Fimbriae (pili)
• 4 Functions
• 4.1 Growth and metabolism
• 4.2 Creation
• 4.3 Protein synthesis
• 5 Movement or motility
• 6 Evolution
• 6.1 Origin of the first cell
[edit] Anatomy
There are two types of cells: eukaryotic and prokaryotic. Prokaryotic cells are usually independent,
while eukaryotic cells are often found in multicellular organisms.

[edit] Prokaryotic cells

Main article: Prokaryote
Diagram of a typical prokaryotic cell
 The prokaryote cell is simpler, and therefore smaller, than a eukaryote cell, lacking a nucleus and
most of the other organelles of eukaryotes. There are two kinds of prokaryotes: bacteria and archaea; these
share a similar structure.
Nuclear material of prokaryotic cell consist of a single chromosome which is in direct contact with
cytoplasm. Here the undefined nuclear region in the cytoplasm is called nucleoid.
A prokaryotic cell has three architectural regions:
• On the outside, flagella and pili project from the cell's surface. These are structures (not
present in all prokaryotes) made of proteins that facilitate movement and communication between
cells;
• Enclosing the cell is the cell envelope – generally consisting of a cell wall covering a plasma
membrane though some bacteria also have a further covering layer called a capsule. The envelope
gives rigidity to the cell and separates the interior of the cell from its environment, serving as a
protective filter. Though most prokaryotes have a cell wall, there are exceptions such as Mycoplasma
(bacteria) and Thermoplasma (archaea). The cell wall consists of peptidoglycan in bacteria, and acts
as an additional barrier against exterior forces. It also prevents the cell from expanding and finally
bursting (cytolysis) from osmotic pressure against a hypotonic environment. Some eukaryote cells
(plant cells and fungi cells) also have a cell wall;
• Inside the cell is the cytoplasmic region that contains the cell genome (DNA) and ribosomes
and various sorts of inclusions. A prokaryotic chromosome is usually a circular molecule (an
exception is that of the bacterium Borrelia burgdorferi, which causes Lyme disease). Though not
forming a nucleus, the DNA is condensed in a nucleoid. Prokaryotes can carry extrachromosomal
DNA elements called plasmids, which are usually circular. Plasmids enable additional functions, such
as antibiotic resistance.

[edit] Eukaryotic cells

Main article: Eukaryote
 Diagram of a typical animal (eukaryotic) cell, showing subcellular components.
Organelles:
(1) nucleolus
(2) nucleus
(3) ribosome
(4) vesicle
(5) rough endoplasmic reticulum (ER)
(6) Golgi apparatus
(7) Cytoskeleton
(8) smooth endoplasmic reticulum
(9) mitochondria
(10) vacuole
(11) cytoplasm
(12) lysosome
(13) centrioles within centrosome
Eukaryotic cells are about 15 times wider than a typical prokaryote and can be as much as 1000
times greater in volume. The major difference between prokaryotes and eukaryotes is that eukaryotic cells
contain membrane-bound compartments in which specific metabolic activities take place. Most important
among these is a cell nucleus, a membrane-delineated compartment that houses the eukaryotic cell's DNA.
This nucleus gives the eukaryote its name, which means "true nucleus." Other differences include:
• The plasma membrane resembles that of prokaryotes in function, with minor differences in
the setup. Cell walls may or may not be present.
 • The eukaryotic DNA is organized in one or more linear molecules, called chromosomes,
which are associated with histone proteins. All chromosomal DNA is stored in the cell nucleus,
separated from the cytoplasm by a membrane. Some eukaryotic organelles such as mitochondria
also contain some DNA.
• Many eukaryotic cells are ciliated with primary cilia. Primary cilia play important roles in
chemosensation, mechanosensation, and thermosensation. Cilia may thus be "viewed as sensory
cellular antennae that coordinate a large number of cellular signaling pathways, sometimes coupling
the signaling to ciliary motility or alternatively to cell division and differentiation."[7]
• Eukaryotes can move using motile cilia or flagella. The flagella are more complex than those
of prokaryotes.
Table 1: Comparison of features of prokaryotic and eukaryotic cells
Prokaryotes Eukaryotes

Typical organisms bacteria, archaea protists, fungi, plants, animals

~ 10–100 µm (sperm cells, apart from the

Typical size ~ 1–10 µm
tail, are smaller)

nucleoid region; no
Type of nucleus real nucleus with double membrane
real nucleus

DNA circular (usually) linear molecules (chromosomes) with

 histone proteins

RNA-/protein- coupled in RNA-synthesis inside the nucleus

synthesis cytoplasm protein synthesis in cytoplasm

Ribosomes 50S+30S 60S+40S

Cytoplasmatic highly structured by endomembranes and

very few structures
structure a cytoskeleton

flagella made of flagella and cilia containing microtubules;

Cell movement
flagellin lamellipodia and filopodia containing actin

one to several thousand (though some

Mitochondria none
lack mitochondria)

Chloroplasts none in algae and plants

single cells, colonies, higher multicellular

Organization usually single cells
organisms with specialized cells

Binary fission Mitosis (fission or budding)

Cell division
(simple division) Meiosis
 Table 2: Comparison of structures between animal and plant cells
Typical animal cell Typical plant cell

• Nucleus
• Nucleus
• Nucleolus (within
• Nucleolus (within
nucleus)
nucleus)
• Rough endoplasmic
• Rough ER
reticulum (ER)
• Smooth ER
• Smooth ER
• Ribosomes
• Ribosomes
• Cytoskeleton
Organelles • Cytoskeleton
• Golgi apparatus
• Golgi apparatus
(dictiosomes)
• Cytoplasm
• Cytoplasm
• Mitochondria
• Mitochondria
• Vesicles
• Plastids and its derivatives
• Lysosomes
• Vacuole(s)
• Centrosome
• Cell wall
• Centrioles
 [edit] Subcellular components

The cells of eukaryotes (left) and prokaryotes (right)

All cells, whether prokaryotic or eukaryotic, have a membrane that envelops the cell, separates its
interior from its environment, regulates what moves in and out (selectively permeable), and maintains the
electric potential of the cell. Inside the membrane, a salty cytoplasm takes up most of the cell volume. All
cells possess DNA, the hereditary material of genes, and RNA, containing the information necessary to build
various proteins such as enzymes, the cell's primary machinery. There are also other kinds of biomolecules
in cells. This article will list these primary components of the cell, then briefly describe their function.

[edit] Membrane
Main article: Cell membrane
The cytoplasm of a cell is surrounded by a cell membrane or plasma membrane. The plasma
membrane in plants and prokaryotes is usually covered by a cell wall. This membrane serves to separate
and protect a cell from its surrounding environment and is made mostly from a double layer of lipids
(hydrophobic fat-like molecules) and hydrophilic phosphorus molecules. Hence, the layer is called a
phospholipid bilayer. It may also be called a fluid mosaic membrane. Embedded within this membrane is a
variety of protein molecules that act as channels and pumps that move different molecules into and out of the
cell. The membrane is said to be 'semi-permeable', in that it can either let a substance (molecule or ion) pass
through freely, pass through to a limited extent or not pass through at all. Cell surface membranes also
contain receptor proteins that allow cells to detect external signaling molecules such as hormones.

[edit] Cytoskeleton
Main article: Cytoskeleton
 Bovine Pulmonary Artery Endothelial cell: nuclei stained blue, mitochondria stained red, and F-actin,
an important component in microfilaments, stained green. Cell imaged on a fluorescent microscope.
The cytoskeleton acts to organize and maintain the cell's shape; anchors organelles in place; helps
during endocytosis, the uptake of external materials by a cell, and cytokinesis, the separation of daughter
cells after cell division; and moves parts of the cell in processes of growth and mobility. The eukaryotic
cytoskeleton is composed of microfilaments, intermediate filaments and microtubules. There is a great
number of proteins associated with them, each controlling a cell's structure by directing, bundling, and
aligning filaments. The prokaryotic cytoskeleton is less well-studied but is involved in the maintenance of cell
shape, polarity and cytokinesis.[8]

[edit] Genetic material

Two different kinds of genetic material exist: deoxyribonucleic acid (DNA) and ribonucleic acid
(RNA). Most organisms use DNA for their long-term information storage, but some viruses (e.g., retroviruses)
have RNA as their genetic material. The biological information contained in an organism is encoded in its
DNA or RNA sequence. RNA is also used for information transport (e.g., mRNA) and enzymatic functions
(e.g., ribosomal RNA) in organisms that use DNA for the genetic code itself. Transfer RNA (tRNA) molecules
are used to add amino acids during protein translation.
Prokaryotic genetic material is organized in a simple circular DNA molecule (the bacterial
chromosome) in the nucleoid region of the cytoplasm. Eukaryotic genetic material is divided into different,
linear molecules called chromosomes inside a discrete nucleus, usually with additional genetic material in
some organelles like mitochondria and chloroplasts (see endosymbiotic theory).
 A human cell has genetic material contained in the cell nucleus (the nuclear genome) and in the
mitochondria (the mitochondrial genome). In humans the nuclear genome is divided into 23 pairs of linear
DNA molecules called chromosomes. The mitochondrial genome is a circular DNA molecule distinct from the
nuclear DNA. Although the mitochondrial DNA is very small compared to nuclear chromosomes, it codes for
13 proteins involved in mitochondrial energy production and specific tRNAs.
Foreign genetic material (most commonly DNA) can also be artificially introduced into the cell by a
process called transfection. This can be transient, if the DNA is not inserted into the cell's genome, or stable,
if it is. Certain viruses also insert their genetic material into the genome.

[edit] Organelles
Main article: Organelle
The human body contains many different organs, such as the heart, lung, and kidney, with each
organ performing a different function. Cells also have a set of "little organs," called organelles, that are
adapted and/or specialized for carrying out one or more vital functions. Both eukaryotic and prokaryotic cells
have organelles but organelles in eukaryotes are generally more complex and may be membrane bound.
There are several types of organelles in a cell. Some (such as the nucleus and golgi apparatus) are
typically solitary, while others (such as mitochondria, peroxisomes and lysosomes) can be numerous
(hundreds to thousands). The cytosol is the gelatinous fluid that fills the cell and surrounds the organelles.
 Cell nucleus – eukaryotes only - a cell's information
center
The cell nucleus is the most conspicuous
organelle found in a eukaryotic cell. It houses the cell's
chromosomes, and is the place where almost all DNA
replication and RNA synthesis (transcription) occur.
The nucleus is spherical and separated from the
cytoplasm by a double membrane called the nuclear
envelope. The nuclear envelope isolates and protects
a cell's DNA from various molecules that could
accidentally damage its structure or interfere with its
processing. During processing, DNA is transcribed, or
copied into a special RNA, called messenger RNA
(mRNA). This mRNA is then transported out of the
nucleus, where it is translated into a specific protein
molecule. The nucleolus is a specialized region within
the nucleus where ribosome subunits are assembled.
Diagram of a cell nucleus
In prokaryotes, DNA processing takes place in the
cytoplasm.

Mitochondria are self-replicating organelles

that occur in various numbers, shapes, and sizes in
the cytoplasm of all eukaryotic cells. Mitochondria play
a critical role in generating energy in the eukaryotic
cell. Mitochondria generate the cell's energy by
oxidative phosphorylation, using oxygen to release
energy stored in cellular nutrients (typically pertaining
to glucose) to generate ATP. Mitochondria multiply by
splitting in two. Respiration occurs in the cell
mitochondria.
Organelles that are modified chloroplasts are
broadly called plastids, and are involved in energy
storage through photosynthesis, which uses solar
energy to generate carbohydrates and oxygen from
carbon dioxide and water.[citation needed]
Mitochondria and chloroplasts each contain
their own genome, which is separate and distinct from
the nuclear genome of a cell. Both organelles contain
this DNA in circular plasmids, much like prokaryotic
cells, strongly supporting the evolutionary theory of
endosymbiosis; since these organelles contain their
own genomes and have other similarities to
prokaryotes, they are thought to have developed
through a symbiotic relationship after being engulfed
by a primitive cell.[citation needed]
 Endoplasmic reticulum – eukaryotes only
The endoplasmic reticulum (ER) is the
transport network for molecules targeted for certain
modifications and specific destinations, as compared
to molecules that will float freely in the cytoplasm. The
ER has two forms: the rough ER, which has ribosomes
on its surface and secretes proteins into the
cytoplasm, and the smooth ER, which lacks them.
Smooth ER plays a role in calcium sequestration and
release.
 Golgi apparatus – eukaryotes only
The primary function of the Golgi apparatus is
to process and package the macromolecules such as
proteins and lipids that are synthesized by the cell. It is
particularly important in the processing of proteins for
secretion. The Golgi apparatus forms a part of the
endomembrane system of eukaryotic cells. Vesicles
that enter the Golgi apparatus are processed in a cis
to trans direction, meaning they coalesce on the cis
side of the apparatus and after processing pinch off on
the opposite (trans) side to form a new vesicle in the
animal cell.[citation needed]

Diagram of an endomembrane system

Ribosomes
The ribosome is a large complex of RNA and
protein molecules. They each consist of two subunits,
and act as an assembly line where RNA from the
nucleus is used to synthesise proteins from amino
acids. Ribosomes can be found either floating freely or
bound to a membrane (the rough endoplasmatic
reticulum in eukaryotes, or the cell membrane in
prokaryotes).[9]

Lysosomes and Peroxisomes – eukaryotes only

Lysosomes contain digestive enzymes (acid hydrolases). They digest excess or worn-out
organelles, food particles, and engulfed viruses or bacteria. Peroxisomes have enzymes that rid the
cell of toxic peroxides. The cell could not house these destructive enzymes if they were not contained
in a membrane-bound system. These organelles are often called a "suicide bag" because of their
ability to detonate and destroy the cell.[citation needed]

Centrosome – the cytoskeleton organiser

The centrosome produces the microtubules of a cell – a key component of the cytoskeleton. It
directs the transport through the ER and the Golgi apparatus. Centrosomes are composed of two
centrioles, which separate during cell division and help in the formation of the mitotic spindle. A single
centrosome is present in the animal cells. They are also found in some fungi and algae cells.[ citation
needed]
Vacuoles
Vacuoles store food and waste. Some vacuoles store extra water. They are often described
as liquid filled space and are surrounded by a membrane. Some cells, most notably Amoeba, have
contractile vacuoles, which can pump water out of the cell if there is too much water. The vacuoles of
 eukaryotic cells are usually larger in those of plants than animals.

[edit] Structures outside the cell wall

[edit] Capsule
A gelatinous capsule is present in some bacteria outside the cell wall. The capsule may be
polysaccharide as in pneumococci, meningococci or polypeptide as Bacillus anthracis or hyaluronic acid as
in streptococci.[citation needed] Capsules are not marked by ordinary stain and can be detected by special
stain. The capsule is antigenic. The capsule has antiphagocytic function so it determines the virulence of
many bacteria. It also plays a role in attachment of the organism to mucous membranes.[ citation needed]

[edit] Flagella
Flagella are the organelles of cellular mobility. They arise from cytoplasm and extrude through the
cell wall. They are long and thick thread-like appendages, protein in nature. Are most commonly found in
bacteria cells but are found in animal cells as well.

[edit] Fimbriae (pili)

They are short and thin hair like filaments, formed of protein called pilin (antigenic). Fimbriae are
responsible for attachment of bacteria to specific receptors of human cell (adherence). There are special
types of pili called (sex pili) involved in conjunction.[ citation needed]
[edit] Functions
[edit] Growth and metabolism
Main articles: Cell growth and Metabolism
Between successive cell divisions, cells grow through the functioning of cellular metabolism. Cell
metabolism is the process by which individual cells process nutrient molecules. Metabolism has two distinct
divisions: catabolism, in which the cell breaks down complex molecules to produce energy and reducing
power, and anabolism, in which the cell uses energy and reducing power to construct complex molecules
and perform other biological functions. Complex sugars consumed by the organism can be broken down into
a less chemically complex sugar molecule called glucose. Once inside the cell, glucose is broken down to
make adenosine triphosphate (ATP), a form of energy, through two different pathways.
The first pathway, glycolysis, requires no oxygen and is referred to as anaerobic metabolism. Each
reaction is designed to produce some hydrogen ions that can then be used to make energy packets (ATP). In
prokaryotes, glycolysis is the only method used for converting energy.
The second pathway, called the Krebs cycle, or citric acid cycle, occurs inside the mitochondria and
can generate enough ATP to run all the cell functions.
 An overview of protein synthesis.
Within the nucleus of the cell (light blue), genes (DNA, dark blue) are transcribed into RNA. This RNA is then
subject to post-transcriptional modification and control, resulting in a mature mRNA (red) that is then
transported out of the nucleus and into the cytoplasm (peach), where it undergoes translation into a protein.
mRNA is translated by ribosomes (purple) that match the three-base codons of the mRNA to the three-base
anti-codons of the appropriate tRNA. Newly synthesized proteins (black) are often further modified, such as
by binding to an effector molecule (orange), to become fully active.

[edit] Creation
Main article: Cell division
Cell division involves a single cell (called a mother cell) dividing into two daughter cells. This leads to
growth in multicellular organisms (the growth of tissue) and to procreation (vegetative reproduction) in
unicellular organisms.
Prokaryotic cells divide by binary fission. Eukaryotic cells usually undergo a process of nuclear
division, called mitosis, followed by division of the cell, called cytokinesis. A diploid cell may also undergo
meiosis to produce haploid cells, usually four. Haploid cells serve as gametes in multicellular organisms,
fusing to form new diploid cells.
DNA replication, or the process of duplicating a cell's genome, is required every time a cell divides.
Replication, like all cellular activities, requires specialized proteins for carrying out the job.
[edit] Protein synthesis
Main article: Protein biosynthesis
Cells are capable of synthesizing new proteins, which are essential for the modulation and
maintenance of cellular activities. This process involves the formation of new protein molecules from amino
acid building blocks based on information encoded in DNA/RNA. Protein synthesis generally consists of two
major steps: transcription and translation.
Transcription is the process where genetic information in DNA is used to produce a complementary
RNA strand. This RNA strand is then processed to give messenger RNA (mRNA), which is free to migrate
through the cell. mRNA molecules bind to protein-RNA complexes called ribosomes located in the cytosol,
where they are translated into polypeptide sequences. The ribosome mediates the formation of a polypeptide
sequence based on the mRNA sequence. The mRNA sequence directly relates to the polypeptide sequence
by binding to transfer RNA (tRNA) adapter molecules in binding pockets within the ribosome. The new
polypeptide then folds into a functional three-dimensional protein molecule.

[edit] Movement or motility

Cells can move during many processes: such as wound healing, the immune response and cancer
metastasis. For wound healing to occur, white blood cells and cells that ingest bacteria move to the wound
site to kill the microorganisms that cause infection.
At the same time fibroblasts (connective tissue cells) move there to remodel damaged structures. In the case
of tumor development, cells from a primary tumor move away and spread to other parts of the body. Cell
motility involves many receptors, crosslinking, bundling, binding, adhesion, motor and other proteins.[10] The
process is divided into three steps – protrusion of the leading edge of the cell, adhesion of the leading edge
and de-adhesion at the cell body and rear, and cytoskeletal contraction to pull the cell forward. Each step is
driven by physical forces generated by unique segments of the cytoskeleton.[11][12]

[edit] Evolution
Main article: Evolutionary history of life
The origin of cells has to do with the origin of life, which began the history of life on Earth.

[edit] Origin of the first cell

Further information: Abiogenesis
There are several theories about the origin of small molecules that could lead to life in an early Earth.
One is that they came from meteorites (see Murchison meteorite). Another is that they were created at deep-
sea vents. A third is that they were synthesized by lightning in a reducing atmosphere ( see Miller–Urey
experiment); although it is not clear if Earth had such an atmosphere. There are essentially no experimental
data defining what the first self-replicating forms were. RNA is generally assumed to be the earliest self-
replicating molecule, as it is capable of both storing genetic information and catalyzing chemical reactions
(see RNA world hypothesis). But some other entity with the potential to self-replicate could have preceded
RNA, like clay or peptide nucleic acid.[13]
Cells emerged at least 4.0–4.3 billion years ago. The current belief is that these cells were
heterotrophs. An important characteristic of cells is the cell membrane, composed of a bilayer of lipids. The
early cell membranes were probably more simple and permeable than modern ones, with only a single fatty
acid chain per lipid. Lipids are known to spontaneously form bilayered vesicles in water, and could have
preceded RNA. But the first cell membranes could also have been produced by catalytic RNA, or even have
required structural proteins before they could form.[14]

[edit] Origin of eukaryotic cells

The eukaryotic cell seems to have evolved from a symbiotic community of prokaryotic cells. DNA-
bearing organelles like the mitochondria and the chloroplasts are almost certainly what remains of ancient
symbiotic oxygen-breathing proteobacteria and cyanobacteria, respectively, where the rest of the cell seems
to be derived from an ancestral archaean prokaryote cell – a theory termed the endosymbiotic theory.
There is still considerable debate about whether organelles like the hydrogenosome predated the
origin of mitochondria, or viceversa: see the hydrogen hypothesis for the origin of eukaryotic cells.
Sex, as the stereotyped choreography of meiosis and syngamy that persists in nearly all extant
eukaryotes, may have played a role in the transition from prokaryotes to eukaryotes. An 'origin of sex as
vaccination' theory suggests that the eukaryote genome accreted from prokaryan parasite genomes in
numerous rounds of lateral gene transfer. Sex-as-syngamy (fusion sex) arose when infected hosts began
swapping nuclearized genomes containing co-evolved, vertically transmitted symbionts that conveyed
protection against horizontal infection by more virulent symbionts.[15]

[edit] History
• 1632–1723: Antonie van Leeuwenhoek teaches himself to grind lenses, builds a microscope
and draws protozoa, such as Vorticella from rain water, and bacteria from his own mouth.
• 1665: Robert Hooke discovers cells in cork, then in living plant tissue using an early
microscope.[6]
 • 1839: Theodor Schwann and Matthias Jakob Schleiden elucidate the principle that plants
and animals are made of cells, concluding that cells are a common unit of structure and
development, and thus founding the cell theory.
• The belief that life forms can occur spontaneously (generatio spontanea) is contradicted by
Louis Pasteur (1822–1895) (although Francesco Redi had performed an experiment in 1668 that
suggested the same conclusion).
• 1855: Rudolf Virchow states that cells always emerge from cell divisions (omnis cellula ex
cellula).
• 1931: Ernst Ruska builds first transmission electron microscope (TEM) at the University of
Berlin. By 1935, he has built an EM with twice the resolution of a light microscope, revealing
previously unresolvable organelles.
• 1953: Watson and Crick made their first announcement on the double-helix structure for DNA
on February 28.
• 1981: Lynn Margulis published Symbiosis in Cell Evolution detailing the endosymbiotic
theory.

[edit] See also

Wikimedia Commons has media related to: Cell biology

Main article: Topic outline of cell biology

• Cell biology
 • Cell culture
• Cell type
• Cellular component
• Cytorrhysis
• Cytotoxicity
• Plasmolysis
• Stem cell
• Syncytium

[edit] References
1. ^ Cell Movements and the Shaping of the Vertebrate Body in Chapter 21 of Molecular
Biology of the Cell fourth edition, edited by Bruce Alberts (2002) published by Garland Science.
The Alberts text discusses how the "cellular building blocks" move to shape developing embryos. It is
also common to describe small molecules such as amino acids as "molecular building blocks".
2. ^ Integrative Biology 131 - Lecture 03: Skeletal System at YouTube first 12 minutes of the
lecture covers cells (by Marian Diamond).
3. ^ Campbell, Neil A.; Brad Williamson; Robin J. Heyden (2006). Biology: Exploring Life.
Boston, Massachusetts: Pearson Prentice Hall. ISBN 0-13-250882-6.
http://www.phschool.com/el_marketing.html.
4. ^ Mitzi Perdue. "Facts about Birds and Eggs". http://www.eggscape.com/birds.htm. Retrieved
2010-04-15.
5. ^ Maton, Anthea; Hopkins, Jean Johnson, Susan LaHart, David Quon Warner, Maryanna
Wright, Jill D (1997). Cells Building Blocks of Life. New Jersey: Prentice Hall. ISBN 0-13-423476-6.
 6. ^ a b "... I could exceedingly plainly perceive it to be all perforated and porous, much like a
Honey-comb, but that the pores of it were not regular [..] these pores, or cells, [..] were indeed the
first microscopical pores I ever saw, and perhaps, that were ever seen, for I had not met with any
Writer or Person, that had made any mention of them before this. . . " – Hooke describing his
observations on a thin slice of cork. Robert Hooke
7. ^ Satir, P; Christensen, ST; Søren T. Christensen (2008-03-26). "Structure and function of
mammalian cilia". Histochemistry and Cell Biology (Springer Berlin / Heidelberg) 129 (6): 687–693.
doi:10.1007/s00418-008-0416-9. 1432-119X. PMID 18365235. PMC 2386530.
http://www.springerlink.com/content/x5051hq648t3152q/. Retrieved 2009-09-12.
8. ^ Michie K, Löwe J (2006). "Dynamic filaments of the bacterial cytoskeleton". Annu Rev
Biochem 75: 467–92. doi:10.1146/annurev.biochem.75.103004.142452. PMID 16756499.
9. ^ Ménétret JF, Schaletzky J, Clemons WM, et al., CW; Akey (December 2007). "Ribosome
binding of a single copy of the SecY complex: implications for protein translocation". Mol. Cell 28 (6):
1083–92. doi:10.1016/j.molcel.2007.10.034. PMID 18158904.
10.^ Revathi Ananthakrishnan1 *, Allen Ehrlicher2 ✉. "The Forces Behind Cell Movement".
Biolsci.org. http://www.biolsci.org/v03p0303.htm. Retrieved 2009-04-17.
11.^ Alberts B, Johnson A, Lewis J. et al. Molecular Biology of the Cell, 4e. Garland Science.
2002
12.^ Ananthakrishnan R, Ehrlicher A. The Forces Behind Cell Movement. Int J Biol Sci 2007;
3:303–317. http://www.biolsci.org/v03p0303.htm
13.^ Orgel LE (1998). "The origin of life--a review of facts and speculations". Trends Biochem
Sci 23 (12): 491–5. doi:10.1016/S0968-0004(98)01300-0. PMID 9868373.
14.^ Griffiths G (December 2007). "Cell evolution and the problem of membrane topology".
Nature reviews. Molecular cell biology 8 (12): 1018–24. doi:10.1038/nrm2287. PMID 17971839.
 15.^ Sterrer W (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology 216:
387–396. doi:10.1006/jtbi.2002.3008. PMID 12151256.
• This article incorporates public domain material from the NCBI document "Science
Primer".

[edit] External links

• Inside the Cell
• Virtual Cell's Educational Animations
• The Inner Life of A Cell, a flash video showing what happens inside of a cell
• The Virtual Cell
• Cells Alive!
• Journal of Cell Biology
• The Biology Project > Cell Biology
• Centre of the Cell online
• The Image & Video Library of The American Society for Cell Biology , a collection of peer-
reviewed still images, video clips and digital books that illustrate the structure, function and biology of
the cell.
 [edit] Textbooks
• Alberts B, Johnson A, Lewis J, Raff M, Roberts K, Walter P (2002). Molecular Biology of the
Cell (4th ed.). Garland. ISBN 0815332181. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?
rid=mboc4.TOC&depth=2.
• Lodish H, Berk A, Matsudaira P, Kaiser CA, Krieger M, Scott MP, Zipurksy SL, Darnell J
(2004). Molecular Cell Biology (5th ed.). WH Freeman: New York, NY. ISBN 978-0716743668.
http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=mcb.TOC.
• Cooper GM (2000). The cell: a molecular approach (2nd ed.). Washington, D.C: ASM Press.
ISBN 0-87893-102-3. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=cooper.TOC&depth=2.

[show]
v•d•e
Structures of the cell / organelles

Cell membrane · Nucleus (and Nucleolus) · Endoplasmic reticulum ·

Golgi apparatus · Parenthesome · Autophagosome
Endomembrane
system Vesicles (Exosome · Lysosome · Endosome · Phagosome · Vacuole)
Cytoplasmic granules: Melanosome · Microbody (Glyoxysome,
Peroxisome) · Weibel-Palade body

Endosymbionts Mitochondrion · Plastids (Chloroplast · Chromoplast · Leucoplast)

 Cytoskeleton Microfilaments · Intermediate filaments · Microtubules · Prokaryotic
cytoskeleton

MTOCs Centrosome/Centriole · Basal body · Spindle pole body

Undulipodium Cilium/Flagellum · Axoneme · Radial spoke

Other external Cell wall · Acrosome

Other Cytoplasm · Ribosome · Vault · Proteasome

B strc: perx, skel, epit, ctrs, cili, mito, nucl (chro)

[show]
v•d•e
Hierarchy of life

Biosphere > Ecosystem > Community (Biocoenosis) > Population > Organism > Organ system
> Organ > Tissue > Cell > Organelle > Molecule (Macromolecule · Biomolecule) > Atom
Retrieved from "http://en.wikipedia.org/wiki/Cell_(biology)"

Categories: Cell biology

• This page was last modified on 3 December 2010 at 07:06.

 W000

Cellule (biologie)
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir cellule.

Cellules épithéliales en culture. L'ADN est coloré en vert, les filaments de kératine en rouge.
 Une petite section d'une membrane cellulaire. Cette membrane de cellule moderne est bien plus
sophistiquée que la simple phospholipide à deux couches originelle (les petites sphères à deux queues).
Protéines et glucides ont plusieurs fonctions de régulation du passage de matériau à travers la membrane et
de réaction à l'environnement.
La cellule (du latin cellula petite chambre) est l'unité structurale, fonctionnelle et reproductrice
constituant tout ou partie d'un être vivant (à l'exception des virus). Chaque cellule est une entité vivante qui,
dans le cas d'organismes multicellulaires, fonctionne de manière autonome, mais coordonnée avec les
autres. Les cellules de même type sont réunies en tissus, eux-mêmes réunis en organes.
La théorie cellulaire implique l'unité de tout le vivant : tous les êtres vivants sont composés de
cellules dont la structure fondamentale est commune ainsi que l'homéostasie du milieu intérieur, milieu de
composition physico-chimique régulé et propice au développement des cellules de l'espèce considérée.
Sommaire
[masquer]
• 1 Histoire du concept
• 1.1 Théorie cellulaire
• 2 La notion de cellule
• 2.1 La cellule, machine à produire de l'ordre
• 2.1.1 Un espace clos effectuant des
échanges avec l'extérieur
• 2.1.2 La compartimentation : mise en place
de microenvironnements aux propriétés spécifiques
• 2.1.3 La cellule un flux organisé de matière
et d'énergie
• 2.2 La cellule vectrice de gènes
• 2.2.1 La transmission des gènes et cycle
cellulaire
• 2.2.2 La reproduction sexuée et cycle du
développement
• 2.2.3 La mort cellulaire : la cellule au
service de l'organisme et des gènes
• 2.3 Interdépendance cellulaire : de la cellule à
l'organisme
• 2.3.1 Les êtres unicellulaires : la cellule
Histoire du concept [modifier]

Dessin de « cellules » observées dans des coupes d'écorce d'arbre par Robert Hooke en 1665.
 • 1665 : Robert Hooke découvre des cellules mortes dans du liège, ces cellules lui font penser
aux cellules d'un monastère, d'où le nom. Puis il observe des cellules dans des plantes vivantes, en
utilisant les premiers microscopes.
• 1839 : Theodor Schwann découvre que les plantes et les animaux sont tous faits de cellules,
concluant que la cellule est l'unité commune de structure et de développement, ce qui fonda la
théorie cellulaire. Il donna son nom aux cellules de Schwann.
• La croyance selon laquelle des formes de vie peuvent apparaître spontanément ( génération
spontanée) est réfutée par Louis Pasteur (1822-1895).
• 1858 : Rudolf Virchow affirma que les cellules naissent du résultat de la division cellulaire
(« omnis cellula ex cellula »), ce qui repose en termes cellulaires la question de l'œuf et de la poule.
C'est précisément cette partie qui est attaquée par les tenants du créationnisme ou de son dernier
avatar, le dessein intelligent.

Théorie cellulaire [modifier]

Article détaillé : Théorie cellulaire.
1. La cellule est l'unité constitutive des organismes vivants. Elle en est aussi l'unité
fonctionnelle.
2. L'organisme dépend de l'activité des cellules isolées ou groupées en tissus pour assurer les
différentes fonctions.
3. Les activités biochimiques des cellules sont coordonnées et déterminées par certaines
structures présentes à l'intérieur des cellules.
4. La multiplication des cellules permet le maintien des organismes et leur multiplication.
 5. Cette théorie est formulée en 1838 par Schleiden et Schwann : la cellule est unité de vie
(tout ce qui est vivant est cellulaire). Cette théorie évoque également la présence d'organites à
l'intérieur de ces mêmes cellules.

La notion de cellule [modifier]

Ici on se demande avant tout quelles sont les caractéristiques communes aux cellules, malgré leur
diversité.

La cellule, machine à produire de l'ordre [modifier]

La cellule représente un état hautement organisé de la matière : maintenir cet ordre tout en étant
soumis aux principes de la thermodynamique nécessite la mise en place de structures permettant d'utiliser
l'énergie, la matière extérieure (on crée de l'ordre au niveau de la cellule mais, globalement, l'entropie
augmente) ; la cellule est donc un système thermodynamiquement ouvert.

Un espace clos effectuant des échanges avec l'extérieur [modifier]

La cellule constitue une unité spatiale, délimitée par une membrane. Celle-ci, loin d'être une limite
hermétique, constitue une surface d'échanges permettant la mise en place de flux.
Les membranes plasmiques, malgré leur diversité possèdent, sauf exceptions (certaines archées
thermophiles possédant une seule couche de lipides), une structure identique :
• une bicouche phospholipidique composée de lipides amphiphiles, qui constitue un filtre de
base permettant le passage des substances hydrophobes, freinant celui des hydrophiles.
 • des protéines transmembranaires et périphériques aux rôles divers (transferts, transport,
transduction de signaux...)
La membrane agit non seulement comme un filtre, c'est-à-dire en laissant passer certaines
molécules selon la différence de concentration (appelée à tort gradient de concentration) mais aussi en
utilisant de l'énergie (osmotique, chimique...) pour favoriser les flux endergoniques. Elle permet aussi le
passage de la lumière, de la chaleur... En tant que surface de contact avec l'extérieur, elle assure aussi la
transmission d'informations nécessaires à la réactivité de la cellule aux changements de l'environnement et à
la coordination avec d'autres cellules.
La membrane plasmique crée donc un espace clos en constant échange avec l'environnement
proche.

La compartimentation : mise en place de microenvironnements aux propriétés

spécifiques [modifier]
La présence d'une membrane biologique entourant un espace, que ce soit le cytoplasme ou la
lumière d'un organite, va permettre, en contrôlant les échanges des macromolécules, des ions (et de toute
autre molécule) l'établissement de conditions favorisant certaines réactions par rapport à d'autres : en variant
les différents facteurs physico-chimiques (pH, concentration en ions...), la nature des enzymes et des
produits, leur nombre...
Cet environnement permet ainsi la biosynthèse et la dégradation de molécules organiques, et ainsi
le maintien d'une structure hautement organisée par un recyclage constant des molécules qui le forment.
 Cette compartimentation se trouve particulièrement poussée dans le cas des eucaryotes : elle
permet la spécialisation fonctionnelle des différents organites (la composition de leur lumière étant différente,
ils sont le siège de réactions différentes : on va ainsi pouvoir favoriser la production de tel produit dans un
compartiment, sa destruction dans un autre).

La cellule un flux organisé de matière et d'énergie [modifier]

Cette structure de base (une membrane organisant les échanges entourant un compartiment, lieu de
réactions chimiques spécifiques) va permettre la mise en place et le maintien de flux de matière, d'énergie,
d'information... traversant la cellule. Il y a donc une réelle organisation des échanges cellule-extérieur, qui va
permettre au « système cellule » (au sens thermodynamique) de maintenir sa structure hautement
organisée.
Finalement, la cellule ne constitue pas une structure stable et immuable mais plutôt une entité
dynamique nécessitant un apport constant de matière et d'énergie pour permettre son fonctionnement et le
maintien de sa structure : seule son organisation persiste, ses constituants étant en perpétuel
renouvellement.
Le potentiel de repos de la cellule détermine la différence de potentiel (-70 mV
environ[réf. nécessaire]), avec l'intérieur de la cellule négatif.

La cellule vectrice de gènes [modifier]

Cette structure résulte de l'expression d'un programme génétique complexe (permettant notamment
la synthèse d'enzymes dont on vient de voir l'importance). Celui-ci doit être transmis, en même temps que la
structure de base, au cours des divisions cellulaires. La cellule peut donc être considérée non seulement
comme l'unité structurelle du vivant mais aussi comme un vecteur de gènes assurant leur transmission au fil
des générations.

La transmission des gènes et cycle cellulaire [modifier]

La division cellulaire aboutissant, à partir d'une cellule mère, à deux cellules filles contenant le même
génome (aux erreurs de copie près) nécessite la succession de phases de synthèse protéique permettant le
renouvellement et la croissance cellulaire, de synthèse d'ADN et enfin de partition plus ou moins équitable
de la cellule.
La synthèse protéique résulte de l'expression du matériel génétique, elle se déroule en plusieurs
étapes : transcription de l'ADN en ARN, traduction de l'ARN en une chaîne polypeptidique, repliement de
celle-ci (chez les eucaryotes s'insèrent des phases de maturation où l'on coupe et modifie la séquence
synthétisée). La copie du génome est réalisée par toute une machinerie protéique permettant à l'ADN
polymérase d'accéder à la séquence et de la copier, selon le principe d'appariement des bases.
La partition de la cellule se fait par des mécanismes différents chez les procaryotes et les eucaryotes
(nommée alors mitose) : celle-ci consiste en la partition et la transmission du génome intégral de la cellule
mère.

La reproduction sexuée et cycle du développement [modifier]

La cellule est là aussi le vecteur de gènes et permet un brassage génétique au sein de la population
grâce aux processus cellulaires que sont la méiose et la fécondation.
 La reproduction sexuée est caractéristique des eucaryotes, mais il existe des mécanismes de
brassage génétique chez les procaryotes.

La mort cellulaire : la cellule au service de l'organisme et des gènes [modifier]

La cellule ayant reçu un signal de son environnement va exprimer un programme entraînant sa mort
(l'apoptose étant un de ces mécanismes) : ce phénomène est nécessaire au développement des organismes
pluricellulaires ; autant chez les végétaux (avec par exemple la mort des cellules formant le tube criblé), que
chez les animaux (lors de la mise en place de la main chez l'homme : on a initialement une main palmée, la
mort des cellules permet l'individualisation des doigts). Ce phénomène a aussi été découvert chez certaines
bactéries (la mort cellulaire permet de limiter le nombre de bactéries lorsque les ressources sont
insuffisantes).
La cellule, tant pour les êtres pluricellulaires que pour les unicellulaires, constitue une structure
vouée avant tout à permettre la reproduction de l'organisme et donc la transmission d'une structure de base
contenant un programme génétique. Ainsi, certains auteurs ont été amenés à formuler la théorie du gène
égoïste, considérant les organismes (et donc les cellules) comme de simples structures destinées à assurer
la transmission et la prolifération des gènes (le gène proliférant alors pour lui-même est qualifié d'égoïste).

Interdépendance cellulaire : de la cellule à l'organisme [modifier]

La cellule, en constant échange avec l'extérieur dépend entièrement de celui-ci. Elle dépend aussi et
surtout des autres cellules, à plusieurs niveaux :
 Les êtres unicellulaires : la cellule « bonne à tout faire » [modifier]
Ici, l'être vivant ne comporte qu'une cellule : celle-ci doit donc assurer toutes les fonctions vitales (se
nourrir, intégrer et réagir aux variations du milieu, proliférer...)
La cellule est donc en quelque sorte autonome mais elle dépend tout de même des autres cellules
(rares sont les cellules ne prélevant que dans le milieu des composés exclusivement inorganiques).
Il peut donc exister une interdépendance cellulaire, même pour les êtres unicellulaires.

L'organisme pluricellulaire : une communauté de cellules interdépendantes [modifier]

Ses cellules sont totalement dépendantes du bon fonctionnement des autres cellules: chacune
d'entre elles, bien qu'ayant le même matériel génétique (à de rares exceptions près: les gamètes, les
lymphocytes par exemple), exprime un programme génétique particulier qui la maintient dans une voie de
différenciation (plus ou moins poussée). Cette spécialisation implique le fractionnement d'opérations
effectuées dans une seule cellule pour les unicellulaires : les cellules d'un même organisme s'organisent en
différentes structures (tissus organes systèmes....) réalisant des fonctions particulières. Ce fractionnement
des fonctions nécessite une coordination entre cellules d'où l'émergence de systèmes de communication
entre cellules.
On a donc une interdépendance forte au sein même de l'organisme qui se superpose à la
dépendance aux autres êtres vivants.
 La cellule eucaryote: formation de communautés de cellules intracellulaires [modifier]
La théorie endosymbiotique (théorie démontrée en ce qui concerne les mitochondries et les
chloroplastes) énonce que les cellules eucaryotes se sont formées à partir d'une cellule procaryote ayant
phagocyté puis domestiqué des bactéries : celles-ci seraient à l'origine des mitochondries. L'invagination de
cyanobactéries aurait donné naissance aux chloroplastes.
La cellule eucaryote dérive donc de l'association symbiotique de bactéries qui sont devenues
totalement interdépendantes au point de former une seule et même unité structurale et fonctionnelle.

Les principales structures cellulaires [modifier]

Il existe deux types fondamentaux de cellules selon qu'elles possèdent ou non un noyau :
• les procaryotes dont l'ADN est libre dans le cytoplasme (les bactéries, par exemple). Ils
comprennent les eubactéries et les archéobactéries ;
• les eucaryotes qui ont une organisation complexe, de nombreux organites et dont le noyau
est entouré d'une membrane nucléaire.
Principales différences entre les cellules procaryotes et eucaryotes
Procaryotes Eucaryotes

protistes, champignons, plantes,

représentants bactéries, archées
animaux

Taille typique ~ 1-10 µm ~ 10-100 µm

 nucléoïde; pas de véritable
Type de noyau vrai noyau avec une enveloppe
noyau

circulaire (chromosome), molécules linéaires

ADN avec des protéines HU pour (chromosomes) avec des protéines
eubactéries histone
synthèse d'ARN dans le noyau
ARN/synthèse des
couplé au cytoplasme synthèse de protéines dans le
protéines
cytoplasme

Ribosomes 23S+16S+5S 28S+18S+5,8S+5S

très structuré par des

Structure
très peu de structures membranes intracellulaires et un
cytoplasmique
cytosquelette

Mouvement de la
flagelle fait de flagelline flagelle et cils fait de tubuline
cellule

Métabolisme anaérobie ou aérobie habituellement aérobie

Mitochondries aucune de une à plusieurs douzaines

Chloroplastes aucun dans les algues et les plantes

 chlorophylliennes

cellules isolées, colonies,

habituellement des cellules
Organisation organismes complexes avec des
isolées
cellules spécialisées

Mitose (multiplication conforme

Division de la
division simple de la cellule)
cellule
Méiose (formation de gamètes)

Eubactéries [modifier]
• Le cytoplasme des procaryotes (le contenu de la cellule) est diffus et granulaire, du fait des
ribosomes (complexe macromoléculaire responsable de la synthèse des protéines).
• La membrane plasmique isole l'intérieur de la cellule de son environnement, et sert de filtre
et de porte de communication.
• Il y a souvent [1] une paroi cellulaire. Elle est formée de peptidoglycane chez les
eubactéries, et joue le rôle de barrière supplémentaire contre les forces extérieures. Elle empêche
également la cellule d'éclater sous la pression osmotique dans un environnement hypotonique.
• L'ADN des procaryotes se compose d'une molécule circulaire super enroulée. Bien que sans
véritable noyau, l'ADN est toutefois condensé en un nucléoïde.
Les procaryotes peuvent posséder un ADN extra-chromosomal, organisé en molécules circulaires
appelées plasmides. Ils peuvent avoir des fonctions supplémentaires, telles que la résistance aux
 antibiotiques. Certains procaryotes ont un flagelle leur permettant de se déplacer activement, plutôt que de
dériver passivement.

Spécificités des archées [modifier]

Les archéobactéries (archaea) sont considérées comme similaires à certains des premiers
organismes qui existèrent sur Terre. On les rencontre notamment dans des milieux extrêmes (elles sont
souvent appelées extrémophiles), tels que geysers, monts hydrothermaux, les fonds abyssaux. Certaines
peuvent résister à des pressions et des températures extrêmes, et avoir un métabolisme basé sur le
méthane ou le soufre.

Cellule eucaryote [modifier]

Organisation d'une cellule animale eucaryote typique.
1. Nucléole

2. Noyau

3. Ribosome

4. Vésicule

5. Réticulum
endoplasmique rugueux
(granuleux)
 6. Appareil de
Golgi

7. Microtubule

8. Réticulum
endoplasmique lisse

9. Mitochondrie

10. Lysosome

11. Cytoplasme
(rempli par le cytosol)

12.
Peroxysome

13. Centrosome
 • Le cytoplasme n'est pas aussi granulaire que celui des procaryotes, puisque la majeure
partie de ses ribosomes sont rattachés au réticulum endoplasmique.
• La membrane plasmique ressemble, dans sa fonction, à celle des procaryotes, avec
quelques différences mineures dans sa configuration.
• La paroi cellulosique, quand elle existe (végétaux), est composée de polysaccharides,
principalement la cellulose.
• L'ADN des eucaryotes est organisé en une ou plusieurs molécules linéaires. Ces molécules
se condensent en s'enroulant autour d'histones lors de la division cellulaire. Tous les chromosomes
de l'ADN sont stockés dans le noyau, séparés du cytoplasme par une membrane. Les eucaryotes ne
possèdent pas de plasmides : seuls quelques organites peuvent contenir de l'ADN.
• Certaines cellules eucaryotes peuvent devenir mobiles, en utilisant un cil ou un flagelle
(spermatozoïde par exemple). Leur flagelle est plus évolué que celui des procaryotes.
Les eucaryotes contiennent plusieurs organites. Ce sont des compartiments cellulaires baignant
dans le hyaloplasme. Ils sont délimités par une membrane plasmique (simple, double ou triple) et possèdent
des fonctions spécifiques.
• Le réticulum endoplasmique (RE) est une extension de la membrane du noyau. Il est divisé
en RE lisse (REL) et RE rugueux (RER) (parfois appelé RE granuleux REG), en fonction de son
apparence au microscope. La surface du RE rugueux est couverte de ribosomes qui insèrent les
 protéines néosynthétisées dans le RE. Du RE, les protéines sont transportées vers l'appareil de
Golgi grâce à des vésicules.
• L'appareil de Golgi est le lieu de transformation finale des protéines. La glycosylation (ajout
de chaînes glucidiques complexes) se réalise à ce niveau.
• Les mitochondries jouent un rôle important dans le métabolisme de la cellule. Elles
contiennent leur propre génome (l'ADN mitochondrial). C'est là que se déroulent la respiration
cellulaire et la fabrication de l'énergie, l'ATP (Adénosine TriPhosphate). Cette énergie est
indispensable aux réactions métaboliques.
• Le cytosquelette permet à la cellule de conserver sa forme (Tenségrité) et de se mouvoir. Il
est également important lors de la division cellulaire, et dans le système de transport intracellulaire.
• Les plastes sont présents dans les plantes et les algues. Les plus connus sont les
chloroplastes, dans les cellules d'organismes photosynthétiques, qui convertissent l'énergie
lumineuse du Soleil en énergie chimique utilisée pour fabriquer des sucres à partir de dioxyde de
carbone (phase sombre de la photosynthèse). Ils possèdent également leur propre génome. Ils sont
le fruit de l'endosymbiose d'une cyanobactéries.
• Chez les plantes, les algues et les champignons, la cellule est encerclée par une paroi
cellulaire pectocellulosique qui fournit un squelette à l'organisme[2]. Des dépositions de composés
tels que la subérine ou la lignine modulent les propriétés physico-chimiques de la paroi, la rendant
plus solide ou plus imperméable, par exemple.
Certains eucaryotes unicellulaires peuvent former des structures multicellulaires. Ces colonies
consistent soit en des groupes de cellules identiques, capables de rester en vie une fois séparées de la
colonie principale (par exemple, les champignons), soit en des groupes de cellules spécialisées
interdépendantes.
Les méthodes d'étude de la cellule in vitro [modifier]

Microscopie [modifier]
La microscopie optique (résolution de +/- 0,25µm en lumière visible) permet l'observation de la
structure des cellules eucaryotes.
La microscopie électronique (résolution de quelques Angströms) révèle l'ultrastructure de celles-ci et
permet une observation plus poussée de la structure des cellules procaryotes comme eucaryotes.

Marquage de molécules [modifier]

Pour étudier l'organisation subcellulaire des cellules au microscope, les tissus peuvent, en fonction
de la méthode choisie, être vivants, ce qui permet une observation dynamique, ou fixés et préparés en
coupes histologiques, ce qui permet en général une observation plus précise, mais figée et ponctuelle.
Localisation subcellulaire par l'utilisation de gènes rapporteurs tels que la GFP (green fluorescing
protein) et la luciférase, par immunocytochimie, ou grâce à des molécules radioactives.
Différentes colorations, vitales ou non, permettent l'observation des structures au microscope
optique : rouge neutre pour les vacuoles, violet dahlia ou cristal pour le noyau...
Étude des constituants cellulaires [modifier]
Isolement de structures: par choc osmotique, ou grâce à des détergents puis par centrifugation.
Purification de protéines: par dialyse, chromatographie, électrophorèse....

Numération des cellules [modifier]

Il est fréquent de devoir compter le nombre de cellules vivantes dans une boîte de culture et de le
comparer au nombre de cellules total, par exemple pour déterminer la toxicité d'un produit. L'une de ces
méthodes de numération est réalisée grâce au test MTT.

Notes et références [modifier]

1. ↑ pas chez les mycoplasmes, par exemple
2. ↑ Geoffrey M. Cooper, La cellule : une approche moléculaire, De Boeck Université, 1999, p.
502

Voir aussi [modifier]

• Voir la vidéo "the inner life of the cell" faite par BioVision (harvard)
• Voir la vidéo sur la cellule dans l'encyclopédie médicale Vulgaris
Sur les autres projets Wikimédia :
• « Cellule (biologie) », sur Wikimedia Commons (ressources multimédia)
• « Cellule (biologie) », sur le Wiktionnaire (dictionnaire universel)
Liens internes [modifier]
• Biologie cellulaire
• Cellule souche
• Culture cellulaire
• Cellule végétale
• Respiration cellulaire
• Unicellulaire

Références externes [modifier]

• (fr) eBiologie.fr : le site de la Science du vivant
• (fr) Dossier Sagascience du CNRS : la cellule animale
• (fr) Qu'est ce qu'une cellule : La cellule en microcinéma
• (fr) Schéma détaillé d'une cellule animale.
• (fr) Schéma détaillé d'une cellule végétale.
• (fr) Exploration 3D d'une cellule : sinauer.com, aimediaserver.com
• (de) Cell Biology - Graphics
• biologie cellulaire des molécules aux organismes, Jean Claude Callen

• Portail de la biologie cellulaire et moléculaire

W000

Introduction to evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is intended as an accessible, non-technical introduction to the subject. For the main
encyclopedia article, see Evolution.
Artist's depiction of a T. rex.
Natural selection does not lead to
perfection; dramatic changes in the
environment often lead to mass
extinctions, as in the case of the dinosaurs
nearly 65 million years ago.
Overview

Life forms reproduce to make

offspring.

The offspring differs from the

parent in minor random ways.

If the differences are helpful, the

offspring is more likely to survive and
reproduce.

This means that more offspring in

the next generation will have the helpful
difference.

These differences accumulate

resulting in changes within the population.

Over time, populations branch off

to become new species as they become
geographically separated and genetically
isolated.

This process is responsible for the

many diverse life forms in the world today.
Haeckel's Paleontological Tree of
Vertebrates (c. 1879).
 The evolutionary history of species has
been described as a "tree", with many
branches arising from a single trunk. While
Haeckel's tree is somewhat outdated, it
illustrates clearly the principles that more
complex modern reconstructions can
obscure.
Evolution is the process of change in all forms of life over generations, and evolutionary biology is the
study of how evolution occurs.
The biodiversity of life evolves by means of natural selection, mutations and genetic drift. The
principles of natural selection are based on three factual observations. First, every individual is supplied with
hereditary material in the form of genes that are received from their parents; then, passed on to their
offspring. Second, organisms tend to produce more offspring than the environment can support. Third, there
are variations among offspring as a consequence of either the random introduction of new genes via
mutations or reshuffling of existing genes during sexual reproduction.[1][2][3]
Natural selection will occur when these repeatedly observed facts of nature (heredity, overproduction
of offspring, and variation) hold true. Natural selection means individuals do not have equal chances of
reproductive success. As a consequence, some individuals produce more offspring and thus have a higher
degree of fitness. Traits that ensure organisms are better adapted to their living conditions become more
common in descendant populations.[2][3] For this reason, populations will never remain exactly the same
over successive generations. The forces of evolution are most evident when populations become isolated,
either through geographic distance or by mechanisms that prevent genetic exchange. Over time, isolated
populations can branch off into new species.[4][5] This is the basic premise behind evolutionary theory that
explains the origins of new species by means of natural selection.[2]
Random genetic drift describes another natural process that regulates the evolution of minor
mutations in the genes, leading to changes in allele frequencies over time. These smaller mutations occur
with regular frequency in and among populations. The vast majority of genetic mutations neither assist,
change the appearance of, nor bring harm to individuals. These mutated genes are neutrally sorted among
populations and survive across generations by chance alone. When species migrate they carry different
genetic varieties to different places. When organisms mate they exchange genetic material and new
individuals are born.
The outward expression of each unique genetic mixture in different environments, the phenotype,
creates a diversity of traits that can be measured and observed as individuals grow and develop. Physical
and behavioural traits are regulated by environmental conditions and, like clay, they are malleable as
individuals interact and respond to ever changing environmental and ecological situations. In contrast to
genetic drift, natural selection is not a random process because it acts on traits that become adapted for their
functional utilities that are necessary for survival.[6] Natural selection and random genetic drift are forever
constant and dynamic parts of life. More than 99.9% of all species have become extinct since life began over
3,500 million years ago. Evolution is more death than survival and over time this has shaped the branching
structure in the tree of life.[7]
The modern understanding of evolution began with the 1859 publication of Charles Darwin's On the
Origin of Species. In addition, Gregor Mendel's work with plants helped to explain the hereditary patterns of
genetics.[8] Fossil discoveries in paleontology, advances in population genetics and a global network of
scientific research have provided further details into the mechanisms of evolution. Scientists now have a
good understanding of the origin of new species (speciation) and have observed the speciation process in
the laboratory and in the wild. Evolution is the principal theory that biologists use to understand life and is
used in many disciplines, including medicine, psychology, conservation biology, anthropology, forensics,
agriculture and other social-cultural applications.
Contents
[hide]
• 1 Darwin's idea: evolution by natural selection
• 2 Source of variation
• 3 Modern synthesis
• 4 Evidence for evolution
• 4.1 Fossil record
• 4.2 Comparative anatomy
• 4.2.1 Taxonomy
• 4.2.2 Embryology
• 4.2.3 Vestigial structures
• 4.2.4 Convergent evolution
• 4.3 Molecular biology
• 4.4 Co-evolution
• 4.5 Artificial selection
• 5 Species
• 6 Different views on the mechanism of evolution
• 6.1 Rate of change
• 6.2 Unit of change
• 7 Summary
• 8 See also
[edit] Darwin's idea: evolution by natural selection
Further information: Common descent
In the 19th century, natural history collections and museums were a popular pastime. The European
expansion and naval expeditions employed naturalists and curators of grand museums showcasing
preserved and live specimens of the varieties of life. Charles Darwin was an English graduate who was
educated and trained in the disciplines of natural history science. Such natural historians would collect,
catalogue, describe and study the vast collections of specimens stored and managed by curators at these
museums. Charles Darwin served as a ship's naturalist on board the HMS Beagle, assigned to a five-year
research expedition around the world. During his voyage, Darwin observed and collected an abundance of
organisms, being very interested in the diverse forms of life along the coasts of South America and the
neighboring Galapagos Islands.[9][10]
Charles Darwin gained extensive experience as he collected and studied the natural history of life
forms from distant places. Through his studies, Darwin formulated the idea that each species had developed
from ancestors with similar features. In 1838, he described how a process he called natural selection would
make this happen.[11]
Darwin's idea of how evolution works relied on the following observations:[12]
• If all the individuals of a species reproduced successfully, the population of that species
would increase uncontrollably.
• Populations tend to remain about the same size from year to year.
• Environmental resources are limited.
• No two individuals in a given species are exactly alike.
 • Much of this variation in a population can be passed on to offspring.

Charles Darwin proposed the theory of evolution by natural selection.

 Darwin noted that orchids exhibited a variety of complex adaptations to ensure pollination; all derived
from basic floral parts.
Darwin deduced that since organisms produce more offspring than their environment could possibly
support, there must be a competitive struggle for survival—only a few individuals can survive out of each
generation. Darwin realized that it was not chance alone that determined survival. Instead, survival depends
on the traits of each individual and if these traits aid or hinder survival and reproduction. Well-adapted, or
"fit", individuals are likely to leave more offspring than their less well-adapted competitors. Darwin realized
that the unequal ability of individuals to survive and reproduce could cause gradual changes in the
population. Traits that help an organism survive and reproduce would accumulate over generations. On the
other hand, traits that hinder survival and reproduction would disappear. Darwin used the term natural
selection to describe this process.[13]
Natural selection is commonly equated with survival of the fittest, but this expression originated in
Herbert Spencer's Principles of Biology in 1864, after Charles Darwin published his original works. Survival
of the fittest describes the process of natural selection incorrectly, because natural selection is not only about
survival and it is not always the fittest that survives.[14]
Observations of variations in animals and plants formed the basis of the theory of natural selection.
For example, Darwin observed that orchids and insects have a close relationship that allows the pollination of
the plants. He noted that orchids have a variety of structures that attract insects, so that pollen from the
flowers gets stuck to the insects’ bodies. In this way, insects transport the pollen from a male to a female
orchid. In spite of the elaborate appearance of orchids, these specialized parts are made from the same
basic structures that make up other flowers. In Fertilisation of Orchids Darwin proposed that the orchid
flowers did not represent the work of an ideal engineer, but were adapted from pre-existing parts, through
natural selection.[15]
Darwin was still researching and experimenting with his ideas on natural selection when he received
a letter from Alfred Wallace describing a theory very similar to his own. This led to an immediate joint
publication of both theories. Both Wallace and Darwin saw the history of life like a family tree, with each fork
in the tree’s limbs being a common ancestor. The tips of the limbs represented modern species and the
branches represented the common ancestors that are shared amongst many different species. To explain
these relationships, Darwin said that all living things were related, and this meant that all life must be
descended from a few forms, or even from a single common ancestor. He called this process descent with
modification.[12]
 Darwin published his theory of evolution by natural selection in On the Origin of Species in 1859. His
theory means that all life, including humanity, is a product of continuing natural processes. The implication
that all life on earth has a common ancestor has met with objections from some religious groups who believe
even today that the different types of life are due to special creation.[16] Their objections are in contrast to
the level of support for the theory by more than 99 percent of those within the scientific community today.[17]

[edit] Source of variation

Darwin’s theory of natural selection laid the groundwork for modern evolutionary theory, and his
experiments and observations showed that the organisms in populations varied from each other, that some
of these variations were inherited, and that these differences could be acted on by natural selection.
However, he could not explain the source of these variations. Like many of his predecessors, Darwin
mistakenly thought that heritable traits were a product of use and disuse, and that features acquired during
an organism's lifetime could be passed on to its offspring. He looked for examples, such as large ground
feeding birds getting stronger legs through exercise, and weaker wings from not flying until, like the ostrich,
they could not fly at all.[18] This misunderstanding was called the inheritance of acquired characters and was
part of the theory of transmutation of species put forward in 1809 by Jean-Baptiste Lamarck. In the late 19th
century this theory became known as Lamarckism. Darwin produced an unsuccessful theory he called
pangenesis to try to explain how acquired characteristics could be inherited. In the 1880s August
Weismann's experiments indicated that changes from use and disuse could not be inherited, and
Lamarckism gradually fell from favor.[19]
The missing information needed to help explain how new features could pass from a parent to its
offspring was provided by the pioneering genetics work of Gregor Mendel. Mendel’s experiments with several
generations of pea plants demonstrated that inheritance works by separating and reshuffling hereditary
information during the formation of sex cells and recombining that information during fertilization. This is like
mixing different hands of cards, with an organism getting a random mix of half of the cards from one parent,
and half of the cards from the other. Mendel called the information factors; however, they later became
known as genes. Genes are the basic units of heredity in living organisms. They contain the information that
directs the physical development and behavior of organisms.
Genes are made of DNA, a long molecule that carries information. This information is encoded in the
sequence of nucleotides in the DNA, just as the sequence of the letters in words carries information on a
page. The genes are like short instructions built up of the "letters" of the DNA alphabet. Put together, the
entire set of these genes gives enough information to serve as an "instruction manual" of how to build and
run an organism. The instructions spelled out by this DNA alphabet can be changed, however, by mutations,
and this may alter the instructions carried within the genes. Within the cell, the genes are carried in
chromosomes, which are packages for carrying the DNA, with the genes arranged along them like beads on
a string. It is the reshuffling of the chromosomes that results in unique combinations of genes in offspring.
Although such mutations in DNA are random, natural selection is not a process of chance: the
environment determines the probability of reproductive success. The end products of natural selection are
organisms that are adapted to their present environments. Natural selection does not involve progress
towards an ultimate goal. Evolution does not necessarily strive for more advanced, more intelligent, or more
sophisticated life forms.[20] For example, fleas (wingless parasites) are descended from a winged, ancestral
scorpionfly, and snakes are lizards that no longer require limbs - although pythons still grow tiny structures
that are the remains of their ancestor's hind legs.[21][22] Organisms are merely the outcome of variations
that succeed or fail, dependent upon the environmental conditions at the time.
Rapid environmental changes typically cause extinctions.[23] Of all species that have existed on
Earth, 99.9 percent are now extinct.[24] Since life began on Earth, five major mass extinctions have led to
large and sudden drops in the variety of species. The most recent, the Cretaceous–Tertiary extinction event,
occurred 65 million years ago, and has attracted more attention than all others because it killed the
dinosaurs.[25]

[edit] Modern synthesis

Further information: Modern evolutionary synthesis
The modern evolutionary synthesis was the outcome of a merger of several different scientific fields
into a cohesive understanding of evolutionary theory. In the 1930s and 1940s, efforts were made to merge
Darwin's theory of natural selection, research in heredity, and understandings of the fossil records into a
unified explanatory model.[26] The application of the principles of genetics to naturally occurring populations,
by scientists such as Theodosius Dobzhansky and Ernst Mayr, advanced understanding of the processes of
evolution. Dobzhansky's 1937 work Genetics and the Origin of Species was an important step in bridging the
gap between genetics and field biology. Mayr, on the basis of an understanding of genes and direct
observations of evolutionary processes from field research, introduced the biological species concept, which
defined a species as a group of interbreeding or potentially interbreeding populations that are reproductively
isolated from all other populations. The paleontologist George Gaylord Simpson helped to incorporate fossil
research, which showed a pattern consistent with the branching and non-directional pathway of evolution of
organisms predicted by the modern synthesis.
The modern synthesis emphasizes the importance of populations as the unit of evolution, the central
role of natural selection as the most important mechanism of evolution, and the idea of gradualism to explain
how large changes evolve as an accumulation of small changes over long periods of time.
[edit] Evidence for evolution

During the voyage of the Beagle, naturalist Charles Darwin collected fossils in South America, and
found fragments of armor which he thought were like giant versions of the scales on the modern armadillos
living nearby. On his return, the anatomist Richard Owen showed him that the fragments were from gigantic
extinct glyptodons, related to the armadillos. This was one of the patterns of distribution that helped Darwin to
develop his theory.[11]
Further information: Evidence of common descent
Scientific evidence for evolution comes from many aspects of biology, and includes fossils,
homologous structures, and molecular similarities between species' DNA.

[edit] Fossil record

Research in the field of paleontology, the study of fossils, supports the idea that all living organisms
are related. Fossils provide evidence that accumulated changes in organisms over long periods of time have
led to the diverse forms of life we see today. A fossil itself reveals the organism's structure and the
relationships between present and extinct species, allowing paleontologists to construct a family tree for all of
the life forms on earth.[27]
Modern paleontology began with the work of Georges Cuvier (1769–1832). Cuvier noted that, in
sedimentary rock, each layer contained a specific group of fossils. The deeper layers, which he proposed to
be older, contained simpler life forms. He noted that many forms of life from the past are no longer present
today. One of Cuvier’s successful contributions to the understanding of the fossil record was establishing
extinction as a fact. In an attempt to explain extinction, Cuvier proposed the idea of “revolutions” or
catastrophism in which he speculated that geological catastrophes had occurred throughout the earth’s
history, wiping out large numbers of species.[28] Cuvier's theory of revolutions was later replaced by
uniformitarian theories, notably those of James Hutton and Charles Lyell who proposed that the earth’s
geological changes were gradual and consistent.[29] However, current evidence in the fossil record supports
the concept of mass extinctions. As a result, the general idea of catastrophism has re-emerged as a valid
hypothesis for at least some of the rapid changes in life forms that appear in the fossil records.
 A very large number of fossils have now been discovered and identified. These fossils serve as a
chronological record of evolution. The fossil record provides examples of transitional species that
demonstrate ancestral links between past and present life forms.[30] One such transitional fossil is
Archaeopteryx, an ancient organism that had the distinct characteristics of a reptile (such as a long, bony tail
and conical teeth) yet also had characteristics of birds (such as feathers and a wishbone). The implication
from such a find is that modern reptiles and birds arose from a common ancestor.[31]

[edit] Comparative anatomy

Further information: Convergent evolution and Divergent evolution
The comparison of similarities between organisms of their form or appearance of parts, called their
morphology, has long been a way to classify life into closely related groups. This can be done by comparing
the structure of adult organisms in different species or by comparing the patterns of how cells grow, divide
and even migrate during an organism's development.

[edit] Taxonomy
Taxonomy is the branch of biology that names and classifies all living things. Scientists use
morphological and genetic similarities to assist them in categorizing life forms based on ancestral
relationships. For example, orangutans, gorillas, chimpanzees, and humans all belong to the same
taxonomic grouping referred to as a family – in this case the family called Hominidae. These animals are
grouped together because of similarities in morphology that come from common ancestry (called homology).
[32]
 Strong evidence for evolution comes from the analysis of homologous structures: structures in
different species that no longer perform the same task but which share a similar structure.[33] Such is the
case of the forelimbs of mammals. The forelimbs of a human, cat, whale, and bat all have strikingly similar
bone structures. However, each of these four species' forelimbs performs a different task. The same bones
that construct a bat's wings, which are used for flight, also construct a whale's flippers, which are used for
swimming. Such a "design" makes little sense if they are unrelated and uniquely constructed for their
particular tasks. The theory of evolution explains these homologous structures: all four animals shared a
common ancestor, and each has undergone change over many generations. These changes in structure
have produced forelimbs adapted for different tasks.[34]

[edit] Embryology
In some cases, anatomical comparison of structures in the embryos of two or more species provides
evidence for a shared ancestor that may not be obvious in the adult forms. As the embryo develops, these
homologies can be lost to view, and the structures can take on different functions. Part of the basis of
classifying the vertebrate group (which includes humans), is the presence of a tail (extending beyond the
anus) and pharyngeal slits. Both structures appear during some stage of embryonic development but are not
always obvious in the adult form.[35]
Because of the morphological similarities present in embryos of different species during
development, it was once assumed that organisms re-enact their evolutionary history as an embryo. It was
thought that human embryos passed through an amphibian then a reptilian stage before completing their
development as mammals. Such a re-enactment, (often called Recapitulation theory), is not supported by
scientific evidence. What does occur, however, is that the first stages of development are similar in broad
groups of organisms.[36] At very early stages, for instance, all vertebrates appear extremely similar, but do
not exactly resemble any ancestral species. As development continues, specific features emerge from this
basic pattern.

[edit] Vestigial structures

Homology includes a unique group of shared structures referred to as vestigial structures. Vestigial
refers to anatomical parts that are of minimal, if any, value to the organism that possesses them. These
apparently illogical structures are remnants of organs that played an important role in ancestral forms. Such
is the case in whales, which have small vestigial bones that appear to be remnants of the leg bones of their
ancestors which walked on land.[37] Humans also have vestigial structures, including the ear muscles, the
wisdom teeth, the appendix, the tail bone, body hair (including goose bumps), and the semilunar fold in the
corner of the eye.[38]

[edit] Convergent evolution

The bird and the bat wing are examples of convergent evolution.
 A bat is a mammal and its forearm bones have been adapted for flight.
Anatomical comparisons can be misleading, as not all anatomical similarities indicate a close
relationship. Organisms that share similar environments will often develop similar physical features, a
process known as convergent evolution. Both sharks and dolphins have similar body forms, yet are only
distantly related – sharks are fish and dolphins are mammals. Such similarities are a result of both
populations being exposed to the same selective pressures. Within both groups, changes that aid swimming
have been favored. Thus, over time, they developed similar appearances (morphology), even though they
are not closely related.[39]
[edit] Molecular biology
 A section of DNA
Every living organism (with the possible exception of RNA viruses) contains molecules of DNA,
which carries genetic information. Genes are the pieces of DNA that carry this information, and they influence
the properties of an organism. Genes determine an individual's general appearance and to some extent their
behavior. If two organisms are closely related, their DNA will be very similar.[40] On the other hand, the more
distantly related two organisms are, the more differences they will have. For example, brothers are closely
related and have very similar DNA, while cousins share a more distant relationship and have far more
differences in their DNA. Similarities in DNA are used to determine the relationships between species in
much the same manner as they are used to show relationships between individuals. For example, comparing
chimpanzees with gorillas and humans shows that there is as much as a 96 percent similarity between the
DNA of humans and chimps. Comparisons of DNA indicate that humans and chimpanzees are more closely
related to each other than either species is to gorillas.[41][42]
The field of molecular systematics focuses on measuring the similarities in these molecules and
using this information to work out how different types of organisms are related through evolution. These
comparisons have allowed biologists to build a relationship tree of the evolution of life on earth.[43] They
have even allowed scientists to unravel the relationships between organisms whose common ancestors lived
such a long time ago that no real similarities remain in the appearance of the organisms.

[edit] Co-evolution
Co-evolution is a process in which two or more species influence the evolution of each other. All
organisms are influenced by life around them; however, in co-evolution there is evidence that genetically
determined traits in each species directly resulted from the interaction between the two organisms.[40]
 An extensively documented case of co-evolution is the relationship between Pseudomyrmex, a type
of ant, and the acacia, a plant that the ant uses for food and shelter. The relationship between the two is so
intimate that it has led to the evolution of special structures and behaviors in both organisms. The ant
defends the acacia against herbivores and clears the forest floor of the seeds from competing plants. In
response, the plant has evolved swollen thorns that the ants use as shelter and special flower parts that the
ants eat.[44] Such co-evolution does not imply that the ants and the tree choose to behave in an altruistic
manner. Rather, across a population small genetic changes in both ant and tree benefited each. The benefit
gave a slightly higher chance of the characteristic being passed on to the next generation. Over time,
successive mutations created the relationship we observe today.

[edit] Artificial selection

 The results of artificial selection: a Chihuahua mix and a Great Dane.
Artificial selection is the controlled breeding of domestic plants and animals. Humans determine
which animal or plant will reproduce and which of the offspring will survive; thus, they determine which genes
will be passed on to future generations. The process of artificial selection has had a significant impact on the
evolution of domestic animals. For example, people have produced different types of dogs by controlled
breeding. The differences in size between the Chihuahua and the Great Dane are the result of artificial
selection. Despite their dramatically different physical appearance, they and all other dogs evolved from a
few wolves domesticated by humans in what is now China less than 15,000 years ago.[45]
Artificial selection has produced a wide variety of plants. In the case of maize (corn), recent genetic
evidence suggests that domestication occurred 10,000 years ago in central Mexico.[46] Prior to
domestication, the edible portion of the wild form was small and difficult to collect. Today The Maize Genetics
Cooperation • Stock Center maintains a collection of more than 10,000 genetic variations of maize that have
arisen by random mutations and chromosomal variations from the original wild type.[47]
In artificial selection the new breed or variety that emerges is the one with random mutations
attractive to humans, while in natural selection the surviving species is the one with random mutations useful
to it in its non-human environment. In both natural and artificial selection the variations are a result of random
mutations, and the underlying genetic processes are essentially the same.[48] Darwin carefully observed the
outcomes of artificial selection in animals and plants to form many of his arguments in support of natural
selection.[49] Much of his book On the Origin of Species was based on these observations of the many
varieties of domestic pigeons arising from artificial selection. Darwin proposed that if humans could achieve
dramatic changes in domestic animals in short periods, then natural selection, given millions of years, could
produce the differences seen in living things today.
[edit] Species

There are numerous species of cichlids that demonstrate dramatic variations in morphology.
Further information: Species, Speciation, and Phylogenetics
Given the right circumstances, and enough time, evolution leads to the emergence of new species.
Scientists have struggled to find a precise and all-inclusive definition of species. Ernst Mayr (1904–2005)
defined a species as a population or group of populations whose members have the potential to interbreed
naturally with one another to produce viable, fertile offspring. (The members of a species cannot produce
viable, fertile offspring with members of other species).[50] Mayr's definition has gained wide acceptance
among biologists, but does not apply to organisms such as bacteria, which reproduce asexually.
be substantial, so that genetic exchange between the two populations is completely disrupted. In their
separate environments, the genetically isolated groups follow their own unique evolutionary pathways. Each
group will accumulate different mutations as well as be subjected to different selective pressures. The
accumulated genetic changes may result in separated populations that can no longer interbreed if they are
reunited.[13] Barriers that prevent interbreeding are either prezygotic (prevent mating or fertilization) or
postzygotic (barriers that occur after fertilization). If interbreeding is no longer possible, then they will be
considered different species.[51]
Usually the process of speciation is slow, occurring over very long time spans; thus direct
observations within human life-spans are rare. However speciation has been observed in present day
organisms, and past speciation events are recorded in fossils.[52][53][54] Scientists have documented the
formation of five new species of cichlid fishes from a single common ancestor that was isolated fewer than
5000 years ago from the parent stock in Lake Nagubago.[55] The evidence for speciation in this case was
morphology (physical appearance) and lack of natural interbreeding. These fish have complex mating rituals
and a variety of colorations; the slight modifications introduced in the new species have changed the mate
selection process and the five forms that arose could not be convinced to interbreed.[56]
[edit] Different views on the mechanism of evolution

James Hutton
Stephen Jay Gould
 Richard Dawkins
The theory of evolution is widely accepted among the scientific community, serving to link the diverse
specialty areas of biology.[17] Evolution provides the field of biology with a solid scientific base. The
significance of evolutionary theory is best described by the title of a paper by Theodosius Dobzhansky (1900–
1975), published in American Biology Teacher; "Nothing in Biology Makes Sense Except in the Light of
Evolution".[57] Nevertheless, the theory of evolution is not static. There is much discussion within the
scientific community concerning the mechanisms behind the evolutionary process. For example, the rate at
which evolution occurs is still under discussion. In addition, there are conflicting opinions as to which is the
primary unit of evolutionary change – the organism or the gene.
[edit] Rate of change
Two views exist concerning the rate of evolutionary change. Darwin and his contemporaries viewed
evolution as a slow and gradual process. Evolutionary trees are based on the idea that profound differences
in species are the result of many small changes that accumulate over long periods.
The view that evolution is gradual had its basis in the works of the geologist James Hutton (1726–
1797) and his theory called "gradualism". Hutton's theory suggests that profound geological change was the
cumulative product of a relatively slow continuing operation of processes which can still be seen in operation
today, as opposed to catastrophism which promoted the idea that sudden changes had causes which can no
longer be seen at work. A uniformitarian perspective was adopted for biological changes. Such a view can
seem to contradict the fossil record, which shows evidence of new species appearing suddenly, then
persisting in that form for long periods. The paleontologist Stephen Jay Gould (1941–2002) developed a
model that suggests that evolution, although a slow process in human terms, undergoes periods of relatively
rapid change over only a few thousand or million years, alternating with long periods of relative stability, a
model called "punctuated equilibrium" which explains the fossil record without contradicting Darwin's ideas.
[58]

[edit] Unit of change

A common unit of selection in evolution is the organism. Natural selection occurs when the
reproductive success of an individual is improved or reduced by an inherited characteristic, and reproductive
success is measured by the number of an individual's surviving offspring. The organism view has been
challenged by a variety of biologists as well as philosophers. Richard Dawkins (born 1941) proposes that
much insight can be gained if we look at evolution from the gene's point of view; that is, that natural selection
operates as an evolutionary mechanism on genes as well as organisms.[59] In his 1976 book The Selfish
Gene, he explains:
“ Individuals are not stable things, they are fleeting. Chromosomes too are
shuffled to oblivion, like hands of cards soon after they are dealt. But the cards
themselves survive the shuffling. The cards are the genes. The genes are not destroyed
by crossing-over; they merely change partners and march on. Of course they march on.
That is their business. They are the replicators and we are their survival machines.
When we have served our purpose we are cast aside. But genes are denizens of
geological time: genes are forever.[60] ”
Others view selection working on many levels, not just at a single level of organism or gene; for
example, Stephen Jay Gould called for a hierarchical perspective on selection.[61]

[edit] Summary
Several basic observations establish the theory of evolution, which explains the variety and
relationship of all living things. There are genetic variations within a population of individuals. Some
individuals, by chance, have features that allow them to survive and thrive better than their kind. The
individuals that survive will be more likely to have offspring of their own. The offspring might inherit the useful
feature.
neighbors in a particular environment. Fossils, the genetic code, and the peculiar distribution of life on earth
provide a record of evolution and demonstrate the common ancestry of all organisms, both living and long
dead. Evolution can be directly observed in artificial selection, the selective breeding for certain traits of
domestic animals and plants. The diverse breeds of cats, dogs, horses, and agricultural plants serve as
examples of evolution.
Although some groups raise objections to the theory of evolution, the evidence of observation and
experiments over a hundred years by thousands of scientists supports evolution.[16] The result of four billion
years of evolution is the diversity of life around us, with an estimated 1.75 million different species in
existence today.[5][62]

[edit] See also

Evolutionary biology portal

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Creation-evolution controversy
• Evidence of common descent
• Evolution as theory and fact
• Level of support for evolution
 • Misconceptions about evolution

[edit] Notes
1. ^ Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray. p. 1.
http://darwin-online.org.uk/content/frameset?itemID=F373&viewtype=text&pageseq=16. . Related
earlier ideas were acknowledged in Darwin, Charles (1861). On the Origin of Species (3rd ed.).
London: John Murray. xiii. http://darwin-online.org.uk/content/frameset?
itemID=F381&viewtype=text&pageseq=20.
2. ^ a b c Gould, Stephen J. (2002). The Structure of Evolutionary Theory. Harvard University
Press. pp. 1433. ISBN 0674006135, 9780674006133.
3. ^ a b Gregory, T. R. (2009). "Understanding Natural Selection: Essential Concepts and
Common Misconceptions". Evolution: Education and Outreach 2 (2): 156–175. doi:10.1007/s12052-
009-0128-1. http://www.springerlink.com/content/2331741806807x22/.
4. ^ "An introduction to evolution" (web resource). Understanding Evolution: your one-stop
source for information on evolution. The University of California Museum of Paleontology, Berkeley.
2008. http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_02. Retrieved 2008-01-23
5. ^ a b Cavalier-Smith T (2006). "Cell evolution and Earth history: stasis and revolution". Philos
Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842. PMID 16754610.
PMC 1578732. http://www.ncbi.nlm.nih.gov/pubmed/16754610. Retrieved 2008-01-24.
6. ^ Garvin-Doxas, K.; Klymkowsky, M. W. (2008). "Understanding Randomness and its Impact
on Student Learning: Lessons Learned from Building the Biology Concept Inventory (BCI)". CBE Life
Sci Educ. 7 (2): 227–233. doi:10.1187/cbe.07-08-0063. PMID 18519614. PMC 2424310.
http://www.ncbi.nlm.nih.gov/pmc/articles/PMC2424310/.
 7. ^ Raup, D. M. (1992). Extinction: bad genes or bad luck.. New York, W. W.: Norton and Co..
pp. 210. ISBN 978-0393309270. http://books.google.com/?
id=8klou91MwJoC&printsec=frontcover&dq=Extinction:
+bad+genes+or+bad+luck&cd=1#v=onepage&q=.
8. ^ Rhee, Sue Yon (1999). "Gregor Mendel". Access Excellence. National Health Museum.
http://www.accessexcellence.org/RC/AB/BC/Gregor_Mendel.html. Retrieved 2008-01-05.
9. ^ Farber, P. L. (2000). Finding Order in Nature: The Natualist Tradition from Linnaeus to E.
O. Wilson. Baltimore and London: The Johns Hopkins University Press. pp. 136. ISBN 0-8018-6389-
9. http://books.google.com/?id=tyG4pfKJ8WEC&printsec=frontcover&dq=Finding+Order+in+Nature:
+The+Natualist+Tradition+from+Linnaeus+to+E.+O.+Wilson&cd=1#v=onepage&q=.
10.^ Watson, J. D. (2005). Darwin, The Indelible Stamp: The Evolution of an Idea . Philadelphia
and London: Running Press. pp. 1257. ISBN 13-978-0-7624-2136-7. http://books.google.com/?
id=LEWfWf0mUJIC&printsec=frontcover&dq=Darwin:+The+Indelible+Stamp:
+The+Evolution+of+an+Idea&cd=1#v=onepage&q=.
11.^ a b Eldredge, Niles (Spring 2006). "Confessions of a Darwinist". The Virginia Quarterly
Review: 32–53. http://www.vqronline.org/articles/2006/spring/eldredge-confessions-darwinist/.
Retrieved 2008-01-23.
12.^ a b Wyhe, John van (2002). "Charles Darwin: gentleman naturalist". The Complete Work of
Charles Darwin Online. University of Cambridge. http://darwin-online.org.uk/darwin.html. Retrieved
2008-01-16.
13.^ a b c Quammen, David (2004). "Was Darwin Wrong?". National Geographic Magazine.
National Geographic. http://ngm.nationalgeographic.com/ngm/0411/feature1/fulltext.html#top.
Retrieved 2007-12-23.
 14.^ Futuyma, D. J. (2005). The Nature of Natural Selection. Ch. 8, pages 93-98 in Cracraft, J.
and Bybee R. W. (Eds.) Evolutionary Science and Society: Educating a New Generation. American
Institute of Biological Sciences.
15.^ Wyhe, John van (2002). "Fertilisation of Orchids". The Complete Works of Charles Darwin.
University of Cambridge. http://darwin-
online.org.uk/EditorialIntroductions/Freeman_FertilisationofOrchids.html. Retrieved 2008-01-07.
16.^ a b DeVries A (2004). "The enigma of Darwin". Clio Med 19 (2): 136–55. PMID 6085987.
17.^ a b Delgado, Cynthia (2006). "Finding the Evolution in Medicine". NIH Record (National
Institutes of Health). http://nihrecord.od.nih.gov/newsletters/2006/07_28_2006/story03.htm.
Retrieved 2007-12-21.
18.^ (Darwin 1872, p. 108.) Effects of the increased Use and Disuse of Parts, as controlled by
Natural Selection
19.^ Ghiselin, Michael T. (September/October 1994). "Nonsense in schoolbooks: 'The Imaginary
Lamarck'". The Textbook Letter. The Textbook League. http://www.textbookleague.org/54marck.htm.
Retrieved 2008-01-23
20.^ (Gould (a) 1981, p. 24)
21.^ Bejder L, Hall BK (2002). "Limbs in whales and limblessness in other vertebrates:
mechanisms of evolutionary and developmental transformation and loss". Evol. Dev. 4 (6): 445–58.
doi:10.1046/j.1525-142X.2002.02033.x. PMID 12492145.
22.^ Boughner JC, Buchtová M, Fu K, Diewert V, Hallgrímsson B, Richman JM (2007).
"Embryonic development of Python sebae - I: Staging criteria and macroscopic skeletal
morphogenesis of the head and limbs". Zoology (Jena) 110 (3): 212–30.
doi:10.1016/j.zool.2007.01.005. PMID 17499493.
 23.^ Drummond, A; Strimmer, K (Jul 2001). Frequently Asked Questions About Evolution.
"Evolution Library". Bioinformatics (Oxford, England) (WGBH Educational Foundation) 17 (7): 662–3.
ISSN 1367-4803. PMID 11448888. http://www.pbs.org/wgbh/evolution/library/faq/cat03.html.
Retrieved 2008-01-23
24.^ "Roundtable: Mass Extinction". Evolution: a jouney into where we're from and where we're
going. WGBH Educational Foundation. 2001.
http://www.pbs.org/wgbh/evolution/extinction/massext/index.html. Retrieved 2008-01-23 .
25.^ Bambach, R.K.; Knoll, A.H.; Wang, S.C. (December 2004). "Origination, extinction, and
mass depletions of marine diversity". Paleobiology 30 (4): 522–42. doi:10.1666/0094-
8373(2004)030<0522:OEAMDO>2.0.CO;2.
http://findarticles.com/p/articles/mi_qa4067/is_200410/ai_n9458414/pg_1. Retrieved 2008-01-24
26.^ Committee on Defining and Advancing the Conceptual Basis of Biological Sciences (1989).
"The tangled web of biological science". The role of theory in advancing 21st Century
Biology:Catalyzing Transformation Research. National Research Council.
http://books.nap.edu/openbook.php?record_id=12026&page=10. Retrieved 2008-01-06.
27.^ "The Fossil Record - Life's Epic". The Virtual Fossil Museum.
http://www.fossilmuseum.net/fossilrecord.htm. Retrieved 2007-08-31.
28.^ (Tattersall 1995, pp. 5–6)
29.^ (Lyell 1830, p. 76)
30.^ Committee on Revising Science and Creationism: A View from the National Academy of
Sciences, National Academy of Sciences and Institute of Medicine of the National Academies (2008).
"Science, Evolution, and Creationism". National Academy of Sciences.
http://books.nap.edu/openbook.php?record_id=11876&page=22. Retrieved 2008-01-06.
31.^ (Gould (b) 1995, p. 360)
 32.^ (Diamond 1992, p. 16)
33.^ a b Drummond, A; Strimmer, K (Jul 2001). Glossary. "Evolution Library" (web resource).
Bioinformatics (Oxford, England) (WGBH Educational Foundation) 17 (7): 662–3. ISSN 1367-4803.
PMID 11448888. http://www.pbs.org/wgbh/evolution/library/glossary/index.html. Retrieved 2008-01-
23 .
34.^ (Mayr 2001, pp. 25–27)
35.^ (Weichert & Presch 1975, p. 8)
36.^ Miller, Kenneth (1997). "Haeckel and his Embryos". Evolution Resources.
http://www.millerandlevine.com/km/evol/embryos/Haeckel.html. Retrieved 2007-09-02.
37.^ Theobald, Douglas (2004). "29+ Evidences for Macroevolution Part 2: Past History" (pdf).
TalkOrigins Archive. The TalkOrigins Foundation. http://www.talkorigins.org/pdf/comdesc.pdf.
Retrieved 2008-01-27.
38.^ Johnson, George (2002). "Vestigial Structures" (web resource). The Evidence for Evolution.
'On Science' column in St. Louis Post Dispatch.
http://www.txtwriter.com/backgrounders/Evolution/EVpage12.html. Retrieved 2008-01-23 .
39.^ Johnson, George (2002). "Convergent and Divergent Evolution" (web resource). The
Evidence for Evolution. 'On Science' column in St. Louis Post Dispatch.
http://www.txtwriter.com/backgrounders/Evolution/EVpage14.html. Retrieved 2008-01-23 .
40.^ a b Kennedy, Donald; (Working group on teaching evolution) (1998). "Teaching about
evolution and the nature of science". Evolution and the nature of science. The National Academy of
Science. http://www.nap.edu/readingroom/books/evolution98/evol3.html. Retrieved 2007-12-30.
41.^ Lovgren, Stefan (2005-08-31). "Chimps, Humans 96 Percent the Same, Gene Study
Finds". National Geographic News. National Geographic.
http://news.nationalgeographic.com/news/2005/08/0831_050831_chimp_genes.html. Retrieved
2007-12-23.
42.^ (Carroll, Grenier & Weatherbee 2000)
43.^ Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006). "Toward
automatic reconstruction of a highly resolved tree of life". Science 311 (5765): 1283–87.
doi:10.1126/science.1123061. PMID 16513982.
44.^ Janzen, Daniel (1974). "Swollen-Thorn Acacias of Central America" (pdf). Smithsonian
Contributions to Biology. Smithsonian Institution.
http://www.sil.si.edu/smithsoniancontributions/Botany/pdf_hi/sctb-0013.pdf. Retrieved 2007-08-31.
45.^ McGourty, Christine (2002-11-22). "Origin of dogs traced". BBC News.
http://news.bbc.co.uk/2/hi/science/nature/2498669.stm. Retrieved 2007-12-14.
46.^ Hall, Hardy. "Transgene Escape: Are Traditioanl Corn Varieties In Mexico Threatened by
Transgenic Corn Crops". Scientific Creative Quarterly. http://www.scq.ubc.ca/transgene-escape-are-
traditional-corn-varieties-in-mexico-threatened-by-transgenic-corn-crops/?. Retrieved 2007-12-14.
47.^ "The Maize Genetics Cooperation • Stock Center". National Plant Germplasm. U.S.
Department of Agriculture. 2006-06-21. http://maizecoop.cropsci.uiuc.edu/mgc-info.php. Retrieved
2007-12-19.
48.^ Silverman, E. David (2002). "Better Books by Trial and Error.".
http://home.comcast.net/~lifebook/evolve.html. Retrieved 2008-04-04.
49.^ Wilner A. (2006). "Darwin's artificial selection as an experiment". Stud Hist Philos Biol
Biomed Sci. 37 (1): 26–40. doi:10.1016/j.shpsc.2005.12.002. PMID 16473266.
50.^ (Mayr 2001, pp. 165–69)
51.^ Sulloway, Frank J (December 2005). "The Evolution of Charles Darwin". Smithsonian
Magazine. Smithsonian Institution.
http://www.smithsonianmagazine.com/issues/2005/december/darwin.php?page=2. Retrieved 2007-
08-31.
52.^ Jiggins CD, Bridle JR (2004). "Speciation in the apple maggot fly: a blend of vintages?".
Trends Ecol. Evol. (Amst.) 19 (3): 111–14. doi:10.1016/j.tree.2003.12.008. PMID 16701238.
53.^ Boxhorn, John (1995). "Observed Instances of Speciation". TalkOrigins Archive.
http://www.talkorigins.org/faqs/faq-speciation.html. Retrieved 2007-05-10.
54.^ Weinberg JR, Starczak VR, Jorg, D (1992). "Evidence for Rapid Speciation Following a
Founder Event in the Laboratory". Evolution (Evolution, Vol. 46, No. 4) 46 (4): 1214–20.
doi:10.2307/2409766. http://jstor.org/stable/2409766. Retrieved 2008-01-24.
55.^ (Mayr 1970, p. 348)
56.^ (Mayr 1970)
57.^ "NCSE Resource". Cans and Can`ts of Teaching Evolution. National Center for Science
Education. 2001-02-13. http://ncseweb.org/evolution/education/cans-cants-teaching-evolution.
Retrieved 2008-01-01.
58.^ Gould, Stephen Jay (1991). "Opus 200". Stephen Jay Gould Archive. Natural History.
http://www.stephenjaygould.org/library/gould_opus200.html. Retrieved 2007-08-31.
59.^ Wright, Sewall (September 1980). "Genic and Organismic Selection". Evolution (Evolution,
Vol. 34, No. 5) 34 (5): 825. doi:10.2307/2407990. http://links.jstor.org/sici?sici=0014-
3820%28198009%2934%3A5%3C825%3AGAOS%3E2.0.CO%3B2-
Z&size=LARGE&origin=JSTOR-enlargePage. Retrieved 2007-12-23.
60.^ (Dawkins 1976, p. 35)
61.^ Gould SJ, Lloyd EA (1999). "Individuality and adaptation across levels of selection: how
shall we name and generalize the unit of Darwinism?". Proc. Natl. Acad. Sci. U.S.A. 96 (21): 11904–
9. doi:10.1073/pnas.96.21.11904. PMID 10518549. PMC 18385.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=10518549. Retrieved 2008-01-18.
62.^ Sedjo, Roger (2007). "How many species are there?". Environmental Literacy Council.
http://www.enviroliteracy.org/article.php/58.html. Retrieved 2008-01-05.

[edit] References
• Carroll, SB; Grenier, J; Weatherbee, SD (2000). From DNA to Diversity: Molecular Genetics
and the Evolution of Animal Design (2nd ed.). Oxford: Blackwell Publishing. ISBN 1-4051-1950-0
• Darwin, Charles (1872). The Origin of Species (6th ed.). London: John Murray. http://darwin-
online.org.uk/content/frameset?itemID=F391&viewtype=text&pageseq=1
• Dawkins, Richard (1976). The Selfish Gene (1st ed.). Oxford University Press. pp. 33.
ISBN 0192860925. http://www.scribd.com/doc/104123/Richard-Dawkins-The-Selfish-Gene-Original-
Ed
• Diamond, Jared (1992). The Third Chimpanzee: the evolution and future of the human
animal. New York: HarperCollins. ISBN 0060183071
• Gould (a), Stephen Jay (1981). The Panda's Thumb: More Reflections in Natural History .
New York: W.W, Norton & Company. ISBN 0393308197
• Gould (b), Stephen Jay (1995). Dinosaur in a Haystack. New York: Harmony Books.
ISBN 0517703939
• Lyell, Charles (1830). Principles of geology. New York: Penguin Books. ISBN 014043528X.
http://www.esp.org/books/lyell/principles/facsimile/
• Mayr, Ernst (1970). Populations, Species, and Evolution. Cambridge, MA: Belknap Press of
Harvard University Press. ISBN 0674690109
 • Mayr, Ernst (2001). What evolution is. New York: Basic Books. ISBN 0-465-04425-5
• Tattersall, Ian (1995). The Fossil Trail: How We Know What We Think We Know About
Human Evolution. New York: Oxford University Press. ISBN 0195061012
• Weichert, Charles; Presch, William (1975). Elements of Chordate Anatomy. New York:
McGraw-Hill. ISBN 0070690081

[edit] Further reading

• Liam Neeson (narrator). (2001-11-20) (web resource). Evolution: a journey into where we're
from and where we're going. [DVD]. South Burlington, VT: WGBH Boston / PBS television series
Nova. ASIN B00005RG6J. http://www.pbs.org/wgbh/evolution/index.html. Retrieved 2008-01-24. -
Age level: Grade 7+
• Horvitz, Leslie Alan (2002). The complete idiot's guide to evolution. Indianapolis: Alpha
Books. ISBN 0028642260.
• Charlesworth, Deborah; Charlesworth, Brian (2003). Evolution: a very short introduction.
Oxford: Oxford University Press. ISBN 0192802518.
• Sis, Peter (2003). The tree of life: a book depicting the life of Charles Darwin, naturalist,
geologist & thinker. New York: Farrar Straus Giroux. ISBN 0-374-45628-3.
• Thomson, Keith Stewart (2005). Fossils: a very short introduction. Oxford: Oxford University
Press. ISBN 0192805045.
• Greg Krukonis (2008). Evolution For Dummies (For Dummies (Math & Science)) . For
Dummies. ISBN 0-470-11773-7.
• Darwin, Charles (2008). Quammen, David. ed. On the Origin of Species: The Illustrated
Edition. Sterling. ISBN 1402756399
 • Pallan, Mark (2009). The Rough Guide to Evolution. Rough Guides. ISBN 1858289467
• Zimmer, Carl (2009). The Tangled Bank: An Introduction to Evolution . Roberts and Company
Publishers. ISBN 0981519474
• Ellis, R. John (2010). How Science Works: Evolution. Springer. ISBN 9048131820

[edit] External links

• Brain, Marshall. "How Stuff Works: Evolution Library" (web resource). Howstuffworks.com.
http://science.howstuffworks.com/evolution-channel.htm. Retrieved 2008-01-24
• Carl Sagan. (2006-07-06) (Google video). Carl Sagan on evolution. [streaming video].
Google. http://video.google.com/videoplay?docid=-522726029201501667&q=carl+sagan. Retrieved
2008-01-24.
• Carl Sagan. (2006-10-21) (Youtube video). Theory of Evolution Explained. [streaming video].
Youtube. http://www.youtube.com/watch?v=E1Y5zMo74cY. Retrieved 2008-01-24.
• "Evolution Education Wiki: EvoWiki" (web resource). http://wiki.cotch.net/. Retrieved 2008-
01-24
• "The Big Picture on Evolution (PDF)". The Big Picture Series. Wellcome Trust. January 2007.
http://www.wellcome.ac.uk/stellent/groups/corporatesite/@msh_publishing_group/documents/web_d
ocument/wtd026042.pdf. Retrieved 2008-01-23
• "The Talk Origins Archive: Exploring the Creation/Evolution Controversy" (web resource).
http://www.talkorigins.org/. Retrieved 2008-01-24
• (web resource) Understanding Evolution: your one-stop source for information on evolution .
The University of California Museum of Paleontology, Berkeley.
http://evolution.berkeley.edu/evolibrary/article/0_0_0/evo_01. Retrieved 2008-01-24
 • "University of Utah Genetics Learning Center animated tour of the basics of genetics" (web
resource). Howstuffworks.com. http://learn.genetics.utah.edu/units/basics/tour. Retrieved 2008-01-
24
• "Introduction To Evolution" (web resource). vectorsite.net.
http://www.vectorsite.net/taevo.html. Retrieved 2010-06-01

[edit] Related information

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Evolution
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This article is about evolution in biology. For other uses, see Evolution (disambiguation).
For a generally accessible and less technical introduction to the topic, see Introduction to evolution.
Part of the Biology series on
 Evolution

Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

 Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
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Evolutionary biology fields

Cladistics
Ecological genetics
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Evolutionary psychology
Molecular evolution
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Biology portal · v • d • e
Evolution is the change in the inherited traits of a population of organisms through successive
generations.[1] After a population splits into smaller groups, these groups evolve independently and may
eventually diversify into new species. Ultimately, life is descended from a common ancestry through a long
series of these speciation events, stretching back in a tree of life that has grown over the 3.5 billion years of
life on Earth. This is visible in anatomical, genetic and other likenesses between groups of organisms,
geographical distribution of related species, the fossil record and the recorded genetic changes in living
organisms over many generations. To distinguish from other uses of the word evolution, it is sometimes
termed biological evolution, genetic evolution or organic evolution.[2][3][4]
Evolution is the product of two opposing forces: processes that constantly introduce variation in traits,
and processes that make particular variants become more common or rare. A trait is a particular
characteristic, such as eye color, height, or a behavior, that is expressed when an organism's genes interact
with its environment. Genes vary within populations, so organisms show heritable differences (variation) in
their traits. The main cause of variation is mutation, which changes the sequence of a gene. Altered genes,
or alleles, are then inherited by offspring. There can sometimes also be transfer of genes between species.
Two main processes cause variants to become more common or rare in a population. One is natural
selection, through which traits that aid survival and reproduction become more common, while traits that
hinder survival and reproduction become more rare.[1][5] Natural selection occurs because only a few
individuals in each generation will survive, since resources are limited and organisms produce many more
offspring than their environment can support. Over many generations, mutations produce successive, small,
random changes in traits, which are then filtered by natural selection and the beneficial changes retained.
This adjusts traits so they become suited to an organism's environment: these adjustments are called
adaptations.[6] Not every trait, however, is an adaptation. Another cause of evolution is genetic drift, which
produces entirely random changes in how common traits are in a population. Genetic drift comes from the
role that chance plays in whether a trait will be passed on to the next generation.
Evolutionary biologists document the fact that evolution occurs, and also develop and test theories
that explain its causes. The study of evolutionary biology began in the mid-nineteenth century, when
research into the fossil record and the diversity of living organisms convinced most scientists that species
changed over time.[7][8] The mechanism driving these changes remained unclear until the theories of natural
selection were independently proposed by Charles Darwin and Alfred Wallace. In 1859, Darwin's seminal
work On the Origin of Species brought the new theories of evolution by natural selection to a wide audience,
[9] leading to the overwhelming acceptance of evolution among scientists.[10][11][12][13] In the 1930s,
Darwinian natural selection became understood in combination with Mendelian inheritance, forming the
modern evolutionary synthesis,[14] which connected the units of evolution (genes) and the mechanism of
evolution (natural selection). This powerful explanatory and predictive theory has become the central
organizing principle of modern biology, directing research and providing a unifying explanation for the history
and diversity of life on Earth.[11][12][15] Evolution is therefore applied and studied in fields as diverse as
agriculture, anthropology, conservation biology, ecology, medicine, paleontology, philosophy, and
psychology along with other specific topics in the previous listed fields.
Contents
[hide]
• 1 History of evolutionary thought
• 2 Heredity
• 3 Variation
• 3.1 Mutation
• 3.2 Sex and recombination
• 3.3 Population genetics
• 3.4 Gene flow
• 4 Mechanisms
• 4.1 Natural selection
• 4.2 Genetic drift
• 5 Outcomes
• 5.1 Adaptation
• 5.2 Co-evolution
• 5.3 Co-operation
• 5.4 Speciation
• 5.5 Extinction
• 6 Evolutionary history of life
• 6.1 Origin of life
• 6.2 Common descent
[edit] History of evolutionary thought
For more details on this topic, see History of evolutionary thought.

Around 1854 Charles Darwin began writing out what became On the Origin of Species.
The scientific inquiry into the origin of species can be dated to at least the 6th century BCE, with the
Greek philosopher Anaximander.[16] Others who considered evolutionary ideas included the Greek
philosopher Empedocles, the Roman philosopher-poet Lucretius, the Afro-Arab biologist Al-Jahiz,[17] the
Persian philosopher Ibn Miskawayh, the Brethren of Purity,[18] and the Chinese philosopher Zhuangzi.[19]
As biological knowledge grew in the 18th century, evolutionary ideas were set out by a few natural
philosophers including Pierre Maupertuis in 1745 and Erasmus Darwin in 1796.[20] The ideas of the biologist
Jean-Baptiste Lamarck about transmutation of species influenced radicals, but were rejected by mainstream
scientists. Charles Darwin formulated his idea of natural selection in 1838 and was still developing his theory
in 1858 when Alfred Russel Wallace sent him a similar theory, and both were presented to the Linnean
Society of London in separate papers.[21] At the end of 1859, Darwin's publication of On the Origin of
Species explained natural selection in detail and presented evidence leading to increasingly wide acceptance
of the occurrence of evolution.
Debate about the mechanisms of evolution continued, and Darwin could not explain the source of the
heritable variations which would be acted on by natural selection. Like Lamarck, he thought that parents
passed on adaptations acquired during their lifetimes,[22] a theory which was subsequently dubbed
Lamarckism.[23] In the 1880s, August Weismann's experiments indicated that changes from use and disuse
were not heritable, and Lamarckism gradually fell from favour.[24][25] More significantly, Darwin could not
account for how traits were passed down from generation to generation. In 1865 Gregor Mendel found that
traits were inherited in a predictable manner.[26] When Mendel's work was rediscovered in 1900s,
disagreements over the rate of evolution predicted by early geneticists and biometricians led to a rift between
the Mendelian and Darwinian models of evolution.
Yet it was the rediscovery of Gregor Mendel's pioneering work on the fundamentals of genetics (of
which Darwin and Wallace were unaware) by Hugo de Vries and others in the early 1900s that provided the
impetus for a better understanding of how variation occurs in plant and animal traits. That variation is the
main fuel used by natural selection to shape the wide variety of adaptive traits observed in organic life. Even
though Hugo de Vries and other early geneticists rejected gradual natural selection, their rediscovery of and
subsequent work on genetics eventually provided a solid basis on which the theory of evolution stood even
more convincingly than when it was originally proposed.[27]
The apparent contradiction between Darwin's theory of evolution by natural selection and Mendel's
work was reconciled in the 1920s and 1930s by evolutionary biologists such as J.B.S. Haldane, Sewall
Wright, and particularly Ronald Fisher, who set the foundations for the establishment of the field of population
genetics. The end result was a combination of evolution by natural selection and Mendelian inheritance, the
modern evolutionary synthesis.[28] In the 1940s, the identification of DNA as the genetic material by Oswald
Avery and colleagues and the subsequent publication of the structure of DNA by James Watson and Francis
Crick in 1953, demonstrated the physical basis for inheritance. Since then, genetics and molecular biology
have become core parts of evolutionary biology and have revolutionized the field of phylogenetics.[14]
In its early history, evolutionary biology primarily drew in scientists from traditional taxonomically
oriented disciplines, whose specialist training in particular organisms addressed general questions in
evolution. As evolutionary biology expanded as an academic discipline, particularly after the development of
the modern evolutionary synthesis, it began to draw more widely from the biological sciences.[14] Currently
the study of evolutionary biology involves scientists from fields as diverse as biochemistry, ecology, genetics
and physiology, and evolutionary concepts are used in even more distant disciplines such as psychology,
medicine, philosophy and computer science. In the 21st century, current research in evolutionary biology
deals with several areas where the modern evolutionary synthesis may need modification or extension, such
as assessing the relative importance of various ideas on the unit of selection and evolvability and how to fully
incorporate the findings of evolutionary developmental biology.[29][30]

[edit] Heredity
Further information: Introduction to genetics, Genetics, and Heredity
 DNA structure. Bases are in the center, surrounded by phosphate–sugar chains in a double helix.
Evolution in organisms occurs through changes in heritable traits – particular characteristics of an
organism. In humans, for example, eye color is an inherited characteristic and an individual might inherit the
"brown-eye trait" from one of their parents.[31] Inherited traits are controlled by genes and the complete set
of genes within an organism's genome is called its genotype.[32]
The complete set of observable traits that make up the structure and behavior of an organism is
called its phenotype. These traits come from the interaction of its genotype with the environment.[33] As a
result, many aspects of an organism's phenotype are not inherited. For example, suntanned skin comes from
the interaction between a person's genotype and sunlight; thus, suntans are not passed on to people's
children. However, some people tan more easily than others, due to differences in their genotype; a striking
example are people with the inherited trait of albinism, who do not tan at all and are very sensitive to
sunburn.[34]
Heritable traits are passed from one generation to the next via DNA, a molecule that encodes genetic
information.[32] DNA is a long polymer composed of four types of bases. The sequence of bases along a
particular DNA molecule specify the genetic information, in a manner similar to a sequence of letters spelling
out a sentence. Before a cell divides, the DNA is copied, so that each of the resulting two cells will inherit the
DNA sequence.
Portions of a DNA molecule that specify a single functional unit are called genes; different genes
have different sequences of bases. Within cells, the long strands of DNA form condensed structures called
chromosomes. The specific location of a DNA sequence within a chromosome is known as a locus. If the
DNA sequence at a locus varies between individuals, the different forms of this sequence are called alleles.
DNA sequences can change through mutations, producing new alleles. If a mutation occurs within a gene,
the new allele may affect the trait that the gene controls, altering the phenotype of the organism.
However, while this simple correspondence between an allele and a trait works in some cases, most
traits are more complex and are controlled by multiple interacting genes.[35][36] The study of such complex
traits is a major area of current genetic research. Another unsolved question in genetics is whether or not
epigenetics is important in evolution. Epigenetics is when a trait is inherited without there being any change
in gene sequences.[37]

[edit] Variation
Further information: Genetic diversity and Population genetics
An individual organism's phenotype results from both its genotype and the influence from the
environment it has lived in. A substantial part of the variation in phenotypes in a population is caused by the
differences between their genotypes.[36] The modern evolutionary synthesis defines evolution as the change
over time in this genetic variation. The frequency of one particular allele will fluctuate, becoming more or less
prevalent relative to other forms of that gene. Evolutionary forces act by driving these changes in allele
frequency in one direction or another. Variation disappears when a new allele reaches the point of fixation —
when it either disappears from the population or replaces the ancestral allele entirely.[38]
Variation comes from mutations in genetic material, migration between populations (gene flow), and
the reshuffling of genes through sexual reproduction. Variation also comes from exchanges of genes
between different species; for example, through horizontal gene transfer in bacteria, and hybridization in
plants.[39] Despite the constant introduction of variation through these processes, most of the genome of a
species is identical in all individuals of that species.[40] However, even relatively small changes in genotype
can lead to dramatic changes in phenotype: for example, chimpanzees and humans differ in only about 5%
of their genomes.[41]
[edit] Mutation
Further information: Mutation and Molecular evolution

Duplication of part of a chromosome

 Random mutations constantly occur in the genomes of organisms; these mutations create genetic
variation. Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation,
viruses, transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA
replication.[42][43][44] These mutations involve several different types of change in DNA sequences; these
can either have no effect, alter the product of a gene, or prevent the gene from functioning. Studies in the fly
Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably
be harmful, with about 70 percent of these mutations having damaging effects, and the remainder being
either neutral or weakly beneficial.[45] Due to the damaging effects that mutations can have on cells,
organisms have evolved mechanisms such as DNA repair to remove mutations.[42] Therefore, the optimal
mutation rate for a species is a trade-off between costs of a high mutation rate, such as deleterious
mutations, and the metabolic costs of maintaining systems to reduce the mutation rate, such as DNA repair
enzymes.[46] Viruses that use RNA as their genetic material have rapid mutation rates,[47] which can be an
advantage since these viruses will evolve constantly and rapidly, and thus evade the defensive responses of
e.g. the human immune system.[48]
Mutations can involve large sections of a chromosome becoming duplicated (usually by genetic
recombination), which can introduce extra copies of a gene into a genome.[49] Extra copies of genes are a
major source of the raw material needed for new genes to evolve.[50] This is important because most new
genes evolve within gene families from pre-existing genes that share common ancestors.[51] For example,
the human eye uses four genes to make structures that sense light: three for color vision and one for night
vision; all four are descended from a single ancestral gene.[52] New genes can be created from an ancestral
gene when a duplicate copy mutates and acquires a new function. This process is easier once a gene has
been duplicated because it increases the redundancy of the system; one gene in the pair can acquire a new
function while the other copy continues to perform its original function.[53][54] Other types of mutation can
even create entirely new genes from previously noncoding DNA.[55][56] The creation of new genes can also
involve small parts of several genes being duplicated, with these fragments then recombining to form new
combinations with new functions.[57][58] When new genes are assembled from shuffling pre-existing parts,
domains act as modules with simple independent functions, which can be mixed together creating new
combinations with new and complex functions.[59] For example, polyketide synthases are large enzymes
that make antibiotics; they contain up to one hundred independent domains that each catalyze one step in
the overall process, like a step in an assembly line.[60]
Changes in chromosome number may involve even larger mutations, where segments of the DNA
within chromosomes break and then rearrange. For example, two chromosomes in the Homo genus fused to
produce human chromosome 2; this fusion did not occur in the lineage of the other apes, and they retain
these separate chromosomes.[61] In evolution, the most important role of such chromosomal
rearrangements may be to accelerate the divergence of a population into new species by making populations
less likely to interbreed, and thereby preserving genetic differences between these populations.[62]
Sequences of DNA that can move about the genome, such as transposons, make up a major fraction
of the genetic material of plants and animals, and may have been important in the evolution of genomes.[63]
For example, more than a million copies of the Alu sequence are present in the human genome, and these
sequences have now been recruited to perform functions such as regulating gene expression.[64] Another
effect of these mobile DNA sequences is that when they move within a genome, they can mutate or delete
existing genes and thereby produce genetic diversity.[43]

[edit] Sex and recombination

Further information: Genetic recombination and Sexual reproduction
 In asexual organisms, genes are inherited together, or linked, as they cannot mix with genes of other
organisms during reproduction. In contrast, the offspring of sexual organisms contain random mixtures of
their parents' chromosomes that are produced through independent assortment. In a related process called
homologous recombination, sexual organisms exchange DNA between two matching chromosomes.[65]
Recombination and reassortment do not alter allele frequencies, but instead change which alleles are
associated with each other, producing offspring with new combinations of alleles.[66] Sex usually increases
genetic variation and may increase the rate of evolution.[67][68] However, asexuality is advantageous in
some environments as it can evolve in previously sexual animals.[69] Here, asexuality might allow the two
sets of alleles in their genome to diverge and gain different functions.[70]
Recombination allows even alleles that are close together in a strand of DNA to be inherited
independently. However, the rate of recombination is low (approximately two events per chromosome per
generation). As a result, genes close together on a chromosome may not always be shuffled away from each
other, and genes that are close together tend to be inherited together, a phenomenon known as linkage.[71]
This tendency is measured by finding how often two alleles occur together on a single chromosome, which is
called their linkage disequilibrium. A set of alleles that is usually inherited in a group is called a haplotype.
This can be important when one allele in a particular haplotype is strongly beneficial: natural selection can
drive a selective sweep that will also cause the other alleles in the haplotype to become more common in the
population; this effect is called genetic hitchhiking.[72]
When alleles cannot be separated by recombination – such as in mammalian Y chromosomes, which
pass intact from fathers to sons – harmful mutations accumulate.[73][74] By breaking up allele combinations,
sexual reproduction allows the removal of harmful mutations and the retention of beneficial mutations.[75] In
addition, recombination and reassortment can produce individuals with new and advantageous gene
combinations. These positive effects are balanced by the fact that sex reduces an organism's reproductive
rate, can cause mutations and may separate beneficial combinations of genes.[75] The reasons for the
evolution of sexual reproduction are therefore unclear and this question is still an active area of research in
evolutionary biology,[76][77] that has prompted ideas such as the Red Queen hypothesis.[78]

[edit] Population genetics

White peppered moth

 Black morph in peppered moth evolution
Further information: Population genetics
From a genetic viewpoint, evolution is a generation-to-generation change in the frequencies of
alleles within a population that shares a common gene pool .[79] A population is a localized group of
individuals belonging to the same species. For example, all of the moths of the same species living in an
isolated forest represent a population. A single gene in this population may have several alternate forms,
which account for variations between the phenotypes of the organisms. An example might be a gene for
coloration in moths that has two alleles: black and white. A gene pool is the complete set of alleles for a gene
in a single population; the allele frequency measures the fraction of the gene pool composed of a single allele
(for example, what fraction of moth coloration genes are the black allele). Evolution occurs when there are
changes in the frequencies of alleles within a population of interbreeding organisms; for example, the allele
for black color in a population of moths becoming more common.
To understand the mechanisms that cause a population to evolve, it is useful to consider what
conditions are required for a population not to evolve. The Hardy-Weinberg principle states that the
frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only
forces acting on that population are the random reshuffling of alleles during the formation of the sperm or
egg, and the random combination of the alleles in these sex cells during fertilization.[80] Such a population is
said to be in Hardy-Weinberg equilibrium; it is not evolving.[81]

[edit] Gene flow

Further information: Gene flow, Hybrid (biology), and Horizontal gene transfer
 When they mature, male lions leave the pride where they were born and take over a new pride to
mate, causing gene flow between prides.[82]
Gene flow is the exchange of genes between populations, which are usually of the same species.[83]
Examples of gene flow within a species include the migration and then breeding of organisms, or the
exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and
horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing genetic
variation into a population. Immigration may add new genetic material to the established gene pool of a
population. Conversely, emigration may remove genetic material. As barriers to reproduction between two
diverging populations are required for the populations to become new species, gene flow may slow this
process by spreading genetic differences between the populations. Gene flow is hindered by mountain
ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has
hindered the flow of plant genes.[84]
Depending on how far two species have diverged since their most recent common ancestor, it may
still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.[85]
Such hybrids are generally infertile, due to the two different sets of chromosomes being unable to pair up
during meiosis. In this case, closely related species may regularly interbreed, but hybrids will be selected
against and the species will remain distinct. However, viable hybrids are occasionally formed and these new
species can either have properties intermediate between their parent species, or possess a totally new
phenotype.[86] The importance of hybridization in creating new species of animals is unclear, although cases
have been seen in many types of animals,[87] with the gray tree frog being a particularly well-studied
example.[88]
Hybridization is, however, an important means of speciation in plants, since polyploidy (having more
than two copies of each chromosome) is tolerated in plants more readily than in animals.[89][90] Polyploidy is
important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to
pair with an identical partner during meiosis.[91] Polyploids also have more genetic diversity, which allows
them to avoid inbreeding depression in small populations.[92]
Horizontal gene transfer is the transfer of genetic material from one organism to another organism
that is not its offspring; this is most common among bacteria.[93] In medicine, this contributes to the spread
of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other
species.[2] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces
cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred.[94][95] An
example of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have received a range
of genes from bacteria, fungi, and plants.[96] Viruses can also carry DNA between organisms, allowing
transfer of genes even across biological domains.[97] Large-scale gene transfer has also occurred between
the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.
[98]

[edit] Mechanisms
The two main mechanisms that produce evolution are natural selection and genetic drift. Natural
selection is the process which favors genes that aid survival and reproduction. Genetic drift is the random
change in the frequency of alleles, caused by the random sampling of a generation's genes during
reproduction. The relative importance of natural selection and genetic drift in a population varies depending
on the strength of the selection and the effective population size, which is the number of individuals capable
of breeding.[99] Natural selection usually predominates in large populations, whereas genetic drift dominates
in small populations. The dominance of genetic drift in small populations can even lead to the fixation of
slightly deleterious mutations.[100] As a result, changing population size can dramatically influence the
course of evolution. Population bottlenecks, where the population shrinks temporarily and therefore loses
genetic variation, result in a more uniform population.[38]

[edit] Natural selection

Further information: Natural selection and Fitness (biology)
 Natural selection of a population for dark coloration.
Natural selection is the process by which genetic mutations that enhance reproduction become, and
remain, more common in successive generations of a population. It has often been called a "self-evident"
mechanism because it necessarily follows from three simple facts:
 • Heritable variation exists within populations of organisms.
• Organisms produce more offspring than can survive.
• These offspring vary in their ability to survive and reproduce.
These conditions produce competition between organisms for survival and reproduction.
Consequently, organisms with traits that give them an advantage over their competitors pass these
advantageous traits on, while traits that do not confer an advantage are not passed on to the next generation.
[101]
The central concept of natural selection is the evolutionary fitness of an organism.[102] Fitness is
measured by an organism's ability to survive and reproduce, which determines the size of its genetic
contribution to the next generation.[102] However, fitness is not the same as the total number of offspring:
instead fitness is indicated by the proportion of subsequent generations that carry an organism's genes.[103]
For example, if an organism could survive well and reproduce rapidly, but its offspring were all too small and
weak to survive, this organism would make little genetic contribution to future generations and would thus
have low fitness.[102]
If an allele increases fitness more than the other alleles of that gene, then with each generation this
allele will become more common within the population. These traits are said to be "selected for". Examples of
traits that can increase fitness are enhanced survival, and increased fecundity. Conversely, the lower fitness
caused by having a less beneficial or deleterious allele results in this allele becoming rarer — they are
"selected against".[5] Importantly, the fitness of an allele is not a fixed characteristic; if the environment
changes, previously neutral or harmful traits may become beneficial and previously beneficial traits become
harmful.[1] However, even if the direction of selection does reverse in this way, traits that were lost in the past
may not re-evolve in an identical form (see Dollo's law).[104][105]
 A chart showing three types of selection. 1.Disruptive selection 2.Stabilizing selection 3.Directional
selection
Natural selection within a population for a trait that can vary across a range of values, such as height,
can be categorized into three different types. The first is directional selection, which is a shift in the average
value of a trait over time — for example organisms slowly getting taller.[106] Secondly, disruptive selection is
selection for extreme trait values and often results in two different values becoming most common, with
selection against the average value. This would be when either short or tall organisms had an advantage, but
not those of medium height. Finally, in stabilizing selection there is selection against extreme trait values on
both ends, which causes a decrease in variance around the average value and less diversity.[101][107] This
would, for example, cause organisms to slowly become all the same height.
A special case of natural selection is sexual selection, which is selection for any trait that increases
mating success by increasing the attractiveness of an organism to potential mates.[108] Traits that evolved
through sexual selection are particularly prominent in males of some animal species, despite traits such as
cumbersome antlers, mating calls or bright colors that attract predators, decreasing the survival of individual
males.[109] This survival disadvantage is balanced by higher reproductive success in males that show these
hard to fake, sexually selected traits.[110]
Natural selection most generally makes nature the measure against which individuals, and individual
traits, are more or less likely to survive. "Nature" in this sense refers to an ecosystem, that is, a system in
which organisms interact with every other element, physical as well as biological, in their local environment.
Eugene Odum, a founder of ecology, defined an ecosystem as: "Any unit that includes all of the
organisms...in a given area interacting with the physical environment so that a flow of energy leads to clearly
defined trophic structure, biotic diversity, and material cycles (ie: exchange of materials between living and
nonliving parts) within the system."[111] Each population within an ecosystem occupies a distinct niche, or
position, with distinct relationships to other parts of the system. These relationships involve the life history of
the organism, its position in the food chain, and its geographic range. This broad understanding of nature
enables scientists to delineate specific forces which, together, comprise natural selection.
An active area of research is the unit of selection, with natural selection being proposed to work at
the level of genes, cells, individual organisms, groups of organisms and species.[112][113] None of these are
mutually exclusive and selection can act on multiple levels simultaneously.[114] An example of selection
occurring below the level of the individual organism are genes called transposons, which can replicate and
spread throughout a genome.[115] Selection at a level above the individual, such as group selection, may
allow the evolution of co-operation, as discussed below.[116]

[edit] Genetic drift

Further information: Genetic drift and Effective population size
 Simulation of genetic drift of 20 unlinked alleles in populations of 10 (top) and 100 (bottom). Drift to
fixation is more rapid in the smaller population.
Genetic drift is the change in allele frequency from one generation to the next that occurs because
alleles in offspring are a random sample of those in the parents, as well as from the role that chance plays in
determining whether a given individual will survive and reproduce. In mathematical terms, alleles are subject
to sampling error. As a result, when selective forces are absent or relatively weak, allele frequencies tend to
"drift" upward or downward randomly (in a random walk). This drift halts when an allele eventually becomes
fixed, either by disappearing from the population, or replacing the other alleles entirely. Genetic drift may
therefore eliminate some alleles from a population due to chance alone. Even in the absence of selective
forces, genetic drift can cause two separate populations that began with the same genetic structure to drift
apart into two divergent populations with different sets of alleles.[117]
The time for an allele to become fixed by genetic drift depends on population size, with fixation
occurring more rapidly in smaller populations.[118] The precise measure of population that is important is
called the effective population size. The effective population is always smaller than the total population since
it takes into account factors such as the level of inbreeding, the number of animals that are too old or young
to breed, and the lower probability of animals that live far apart managing to mate with each other.[119]
An example when genetic drift is probably of central importance in determining a trait is the loss of
pigments from animals that live in caves, a change that produces no obvious advantage or disadvantage in
complete darkness.[120] However, it is usually difficult to measure the relative importance of selection and
drift,[121] so the comparative importance of these two forces in driving evolutionary change is an area of
current research.[122] These investigations were prompted by the neutral theory of molecular evolution,
which proposed that most evolutionary changes are the result of the fixation of neutral mutations that do not
have any immediate effects on the fitness of an organism.[123] Hence, in this model, most genetic changes
in a population are the result of constant mutation pressure and genetic drift.[124] This form of the neutral
theory is now largely abandoned, since it does not seem to fit the genetic variation seen in nature.[125][126]
However, a more recent and better-supported version of this model is the nearly neutral theory, where most
mutations only have small effects on fitness.[101]
[edit] Outcomes
Evolution influences every aspect of the form and behavior of organisms. Most prominent are the
specific behavioral and physical adaptations that are the outcome of natural selection. These adaptations
increase fitness by aiding activities such as finding food, avoiding predators or attracting mates. Organisms
can also respond to selection by co-operating with each other, usually by aiding their relatives or engaging in
mutually beneficial symbiosis. In the longer term, evolution produces new species through splitting ancestral
populations of organisms into new groups that cannot or will not interbreed.
These outcomes of evolution are sometimes divided into macroevolution, which is evolution that
occurs at or above the level of species, such as extinction and speciation, and microevolution, which is
smaller evolutionary changes, such as adaptations, within a species or population.[127] In general,
macroevolution is regarded as the outcome of long periods of microevolution.[128] Thus, the distinction
between micro- and macroevolution is not a fundamental one – the difference is simply the time involved.
[129] However, in macroevolution, the traits of the entire species may be important. For instance, a large
amount of variation among individuals allows a species to rapidly adapt to new habitats, lessening the
chance of it going extinct, while a wide geographic range increases the chance of speciation, by making it
more likely that part of the population will become isolated. In this sense, microevolution and macroevolution
might involve selection at different levels – with microevolution acting on genes and organisms, versus
macroevolutionary processes such as species selection acting on entire species and affecting their rates of
speciation and extinction.[130][131][132]
A common misconception is that evolution has goals or long-term plans; realistically however,
evolution has no long-term goal and does not necessarily produce greater complexity.[133][134] Although
complex species have evolved, they occur as a side effect of the overall number of organisms increasing,
and simple forms of life still remain more common in the biosphere.[135] For example, the overwhelming
majority of species are microscopic prokaryotes, which form about half the world's biomass despite their
small size,[136] and constitute the vast majority of Earth's biodiversity.[137] Simple organisms have therefore
been the dominant form of life on Earth throughout its history and continue to be the main form of life up to
the present day, with complex life only appearing more diverse because it is more noticeable.[138] Indeed,
the evolution of microorganisms is particularly important to modern evolutionary research, since their rapid
reproduction allows the study of experimental evolution and the observation of evolution and adaptation in
real time.[139][140]

[edit] Adaptation
For more details on this topic, see Adaptation.
Adaptation is one of the basic phenomena of biology,[141] and is the process whereby an organism
becomes better suited to its habitat.[142][143] Also, the term adaptation may refer to a trait that is important
for an organism's survival. For example, the adaptation of horses' teeth to the grinding of grass, or the ability
of horses to run fast and escape predators. By using the term adaptation for the evolutionary process, and
adaptive trait for the product (the bodily part or function), the two senses of the word may be distinguished.
Adaptations are produced by natural selection.[144] The following definitions are due to Theodosius
Dobzhansky.
1. Adaptation is the evolutionary process whereby an organism becomes better able to live in
its habitat or habitats.[145]
2. Adaptedness is the state of being adapted: the degree to which an organism is able to live
and reproduce in a given set of habitats.[146]
 3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or
enhances the probability of that organism surviving and reproducing.[147]
Adaptation may cause either the gain of a new feature, or the loss of an ancestral feature. An
example that shows both types of change is bacterial adaptation to antibiotic selection, with genetic changes
causing antibiotic resistance by both modifying the target of the drug, or increasing the activity of transporters
that pump the drug out of the cell.[148] Other striking examples are the bacteria Escherichia coli evolving the
ability to use citric acid as a nutrient in a long-term laboratory experiment,[149] Flavobacterium evolving a
novel enzyme that allows these bacteria to grow on the by-products of nylon manufacturing,[150][151] and
the soil bacterium Sphingobium evolving an entirely new metabolic pathway that degrades the synthetic
pesticide pentachlorophenol.[152][153] An interesting but still controversial idea is that some adaptations
might increase the ability of organisms to generate genetic diversity and adapt by natural selection
(increasing organisms' evolvability).[154][155]

A baleen whale skeleton, a and b label flipper bones, which were adapted from front leg bones: while
c indicates vestigial leg bones, suggesting an adaptation from land to sea.[156]
 Adaptation occurs through the gradual modification of existing structures. Consequently, structures
with similar internal organization may have different functions in related organisms. This is the result of a
single ancestral structure being adapted to function in different ways. The bones within bat wings, for
example, are very similar to those in mice feet and primate hands, due to the descent of all these structures
from a common mammalian ancestor.[157] However, since all living organisms are related to some extent,
[158] even organs that appear to have little or no structural similarity, such as arthropod, squid and vertebrate
eyes, or the limbs and wings of arthropods and vertebrates, can depend on a common set of homologous
genes that control their assembly and function; this is called deep homology.[159][160]
During adaptation, some structures may lose their original function and become vestigial structures.
[161] Such structures may have little or no function in a current species, yet have a clear function in ancestral
species, or other closely related species. Examples include pseudogenes,[162] the non-functional remains of
eyes in blind cave-dwelling fish,[163] wings in flightless birds,[164] and the presence of hip bones in whales
and snakes.[156] Examples of vestigial structures in humans include wisdom teeth,[165] the coccyx,[161] the
vermiform appendix,[161] and other behavioral vestiges such as goose bumps,[166] and primitive reflexes.
[167][168][169][170]
However, many traits that appear to be simple adaptations are in fact exaptations: structures
originally adapted for one function, but which coincidentally became somewhat useful for some other function
in the process.[171] One example is the African lizard Holaspis guentheri, which developed an extremely flat
head for hiding in crevices, as can be seen by looking at its near relatives. However, in this species, the head
has become so flattened that it assists in gliding from tree to tree—an exaptation.[171] Within cells, molecular
machines such as the bacterial flagella[172] and protein sorting machinery[173] evolved by the recruitment of
several pre-existing proteins that previously had different functions.[127] Another example is the recruitment
of enzymes from glycolysis and xenobiotic metabolism to serve as structural proteins called crystallins within
the lenses of organisms' eyes.[174][175]
A critical principle of ecology is that of competitive exclusion: no two species can occupy the same
niche in the same environment for a long time.[176] Consequently, natural selection will tend to force species
to adapt to different ecological niches. This may mean that, for example, two species of cichlid fish adapt to
live in different habitats, which will minimize the competition between them for food.[177]
An area of current investigation in evolutionary developmental biology is the developmental basis of
adaptations and exaptations.[178] This research addresses the origin and evolution of embryonic
development and how modifications of development and developmental processes produce novel features.
[179] These studies have shown that evolution can alter development to create new structures, such as
embryonic bone structures that develop into the jaw in other animals instead forming part of the middle ear in
mammals.[180] It is also possible for structures that have been lost in evolution to reappear due to changes
in developmental genes, such as a mutation in chickens causing embryos to grow teeth similar to those of
crocodiles.[181] It is now becoming clear that most alterations in the form of organisms are due to changes in
a small set of conserved genes.[182]
[edit] Co-evolution

Common garter snake (Thamnophis sirtalis sirtalis) which has evolved resistance to tetrodotoxin in
its amphibian prey.
Further information: Co-evolution
Interactions between organisms can produce both conflict and co-operation. When the interaction is
between pairs of species, such as a pathogen and a host, or a predator and its prey, these species can
develop matched sets of adaptations. Here, the evolution of one species causes adaptations in a second
species. These changes in the second species then, in turn, cause new adaptations in the first species. This
cycle of selection and response is called co-evolution.[183] An example is the production of tetrodotoxin in
the rough-skinned newt and the evolution of tetrodotoxin resistance in its predator, the common garter snake.
In this predator-prey pair, an evolutionary arms race has produced high levels of toxin in the newt and
correspondingly high levels of toxin resistance in the snake.[184]

[edit] Co-operation
Further information: Co-operation (evolution)
However, not all interactions between species involve conflict.[185] Many cases of mutually
beneficial interactions have evolved. For instance, an extreme cooperation exists between plants and the
mycorrhizal fungi that grow on their roots and aid the plant in absorbing nutrients from the soil.[186] This is a
reciprocal relationship as the plants provide the fungi with sugars from photosynthesis. Here, the fungi
actually grow inside plant cells, allowing them to exchange nutrients with their hosts, while sending signals
that suppress the plant immune system.[187]
Coalitions between organisms of the same species have also evolved. An extreme case is the
eusociality found in social insects, such as bees, termites and ants, where sterile insects feed and guard the
small number of organisms in a colony that are able to reproduce. On an even smaller scale, the somatic
cells that make up the body of an animal limit their reproduction so they can maintain a stable organism,
which then supports a small number of the animal's germ cells to produce offspring. Here, somatic cells
respond to specific signals that instruct them whether to grow, remain as they are, or die. If cells ignore these
signals and multiply inappropriately, their uncontrolled growth causes cancer.[42]
Such cooperation within species may have evolved through the process of kin selection, which is
where one organism acts to help raise a relative's offspring.[188] This activity is selected for because if the
helping individual contains alleles which promote the helping activity, it is likely that its kin will also contain
these alleles and thus those alleles will be passed on.[189] Other processes that may promote cooperation
include group selection, where cooperation provides benefits to a group of organisms.[190]

[edit] Speciation
Further information: Speciation
The four mechanisms of speciation.
 Speciation is the process where a species diverges into two or more descendant species.[191]
Evolutionary biologists view species as statistical phenomena and not categories or types. This view is
counterintuitive since the classical idea of species is still widely held, with a species seen as a class of
organisms exemplified by a "type specimen" that bears all the traits common to this species. Instead, a
species is now defined as a separately evolving lineage that forms a single gene pool. Although properties
such as genetics and morphology are used to help separate closely related lineages, this definition has fuzzy
boundaries.[192] Indeed, the exact definition of the term "species" is still controversial, particularly in
prokaryotes,[193] and this is called the species problem.[194] Biologists have proposed a range of more
precise definitions, but the definition used is a pragmatic choice that depends on the particularities of the
species concerned.[194] Typically the actual focus on biological study is the population, an observable
interacting group of organisms, rather than a species, an observable similar group of individuals.
Speciation has been observed multiple times under both controlled laboratory conditions and in
nature.[195] In sexually reproducing organisms, speciation results from reproductive isolation followed by
genealogical divergence. There are four mechanisms for speciation. The most common in animals is
allopatric speciation, which occurs in populations initially isolated geographically, such as by habitat
fragmentation or migration. Selection under these conditions can produce very rapid changes in the
appearance and behaviour of organisms.[196][197] As selection and drift act independently on populations
isolated from the rest of their species, separation may eventually produce organisms that cannot interbreed.
[198]
The second mechanism of speciation is peripatric speciation, which occurs when small populations
of organisms become isolated in a new environment. This differs from allopatric speciation in that the isolated
populations are numerically much smaller than the parental population. Here, the founder effect causes rapid
speciation through both rapid genetic drift and selection on a small gene pool.[199]
 The third mechanism of speciation is parapatric speciation. This is similar to peripatric speciation in
that a small population enters a new habitat, but differs in that there is no physical separation between these
two populations. Instead, speciation results from the evolution of mechanisms that reduce gene flow between
the two populations.[191] Generally this occurs when there has been a drastic change in the environment
within the parental species' habitat. One example is the grass Anthoxanthum odoratum, which can undergo
parapatric speciation in response to localized metal pollution from mines.[200] Here, plants evolve that have
resistance to high levels of metals in the soil. Selection against interbreeding with the metal-sensitive
parental population produced a gradual change in the flowering time of the metal-resistant plants, which
eventually produced complete reproductive isolation. Selection against hybrids between the two populations
may cause reinforcement, which is the evolution of traits that promote mating within a species, as well as
character displacement, which is when two species become more distinct in appearance.[201]
 Geographical isolation of finches on the Galápagos Islands produced over a dozen new species.
Finally, in sympatric speciation species diverge without geographic isolation or changes in habitat.
This form is rare since even a small amount of gene flow may remove genetic differences between parts of a
population.[202] Generally, sympatric speciation in animals requires the evolution of both genetic differences
and non-random mating, to allow reproductive isolation to evolve.[203]
One type of sympatric speciation involves cross-breeding of two related species to produce a new
hybrid species. This is not common in animals as animal hybrids are usually sterile. This is because during
meiosis the homologous chromosomes from each parent are from different species and cannot successfully
pair. However, it is more common in plants because plants often double their number of chromosomes, to
form polyploids.[204] This allows the chromosomes from each parental species to form matching pairs during
meiosis, since each parent's chromosomes are represented by a pair already.[205] An example of such a
speciation event is when the plant species Arabidopsis thaliana and Arabidopsis arenosa cross-bred to give
the new species Arabidopsis suecica.[206] This happened about 20,000 years ago,[207] and the speciation
process has been repeated in the laboratory, which allows the study of the genetic mechanisms involved in
this process.[208] Indeed, chromosome doubling within a species may be a common cause of reproductive
isolation, as half the doubled chromosomes will be unmatched when breeding with undoubled organisms.[90]
Speciation events are important in the theory of punctuated equilibrium, which accounts for the
pattern in the fossil record of short "bursts" of evolution interspersed with relatively long periods of stasis,
where species remain relatively unchanged.[209] In this theory, speciation and rapid evolution are linked,
with natural selection and genetic drift acting most strongly on organisms undergoing speciation in novel
habitats or small populations. As a result, the periods of stasis in the fossil record correspond to the parental
population, and the organisms undergoing speciation and rapid evolution are found in small populations or
geographically restricted habitats, and therefore rarely being preserved as fossils.[210]

[edit] Extinction
Further information: Extinction
 Tyrannosaurus rex. Non-avian dinosaurs died out in the Cretaceous–Tertiary extinction event at the
end of the Cretaceous period.
Extinction is the disappearance of an entire species. Extinction is not an unusual event, as species
regularly appear through speciation, and disappear through extinction.[211] Nearly all animal and plant
species that have lived on earth are now extinct,[212] and extinction appears to be the ultimate fate of all
species.[213] These extinctions have happened continuously throughout the history of life, although the rate
of extinction spikes in occasional mass extinction events.[214] The Cretaceous–Tertiary extinction event,
during which the non-avian dinosaurs went extinct, is the most well-known, but the earlier Permian–Triassic
extinction event was even more severe, with approximately 96 percent of species driven to extinction.[214]
The Holocene extinction event is an ongoing mass extinction associated with humanity's expansion across
the globe over the past few thousand years. Present-day extinction rates are 100–1000 times greater than the
background rate, and up to 30 percent of species may be extinct by the mid 21st century.[215] Human
activities are now the primary cause of the ongoing extinction event;[216] global warming may further
accelerate it in the future.[217]
The role of extinction in evolution is not very well understood and may depend on which type of
extinction is considered.[214] The causes of the continuous "low-level" extinction events, which form the
majority of extinctions, may be the result of competition between species for limited resources (competitive
exclusion).[14] If one species can out-compete another, this could produce species selection, with the fitter
species surviving and the other species being driven to extinction.[112] The intermittent mass extinctions are
also important, but instead of acting as a selective force, they drastically reduce diversity in a nonspecific
manner and promote bursts of rapid evolution and speciation in survivors.[218]

[edit] Evolutionary history of life

Main article: Evolutionary history of life
See also: Timeline of evolution and Timeline of human evolution

[edit] Origin of life

Further information: Abiogenesis and RNA world hypothesis
The origin of life is a necessary precursor for biological evolution, but understanding that evolution
occurred once organisms appeared and investigating how this happens does not depend on understanding
exactly how life began.[219] The current scientific consensus is that the complex biochemistry that makes up
life came from simpler chemical reactions, but it is unclear how this occurred.[220] Not much is certain about
the earliest developments in life, the structure of the first living things, or the identity and nature of any last
universal common ancestor or ancestral gene pool.[221][222] Consequently, there is no scientific consensus
on how life began, but proposals include self-replicating molecules such as RNA,[223] and the assembly of
simple cells.[224]

[edit] Common descent

Further information: Common descent, Evidence of common descent, and Homology (biology)

The hominoids are descendants of a common ancestor.

All organisms on Earth are descended from a common ancestor or ancestral gene pool.[158][225]
Current species are a stage in the process of evolution, with their diversity the product of a long series of
speciation and extinction events.[226] The common descent of organisms was first deduced from four simple
facts about organisms: First, they have geographic distributions that cannot be explained by local adaptation.
Second, the diversity of life is not a set of completely unique organisms, but organisms that share
morphological similarities. Third, vestigial traits with no clear purpose resemble functional ancestral traits,
and finally, that organisms can be classified using these similarities into a hierarchy of nested groups – similar
to a family tree.[9] However, modern research has suggested that, due to horizontal gene transfer, this "tree
of life" may be more complicated than a simple branching tree since some genes have spread independently
between distantly related species.[227][228]
Past species have also left records of their evolutionary history. Fossils, along with the comparative
anatomy of present-day organisms, constitute the morphological, or anatomical, record.[229] By comparing
the anatomies of both modern and extinct species, paleontologists can infer the lineages of those species.
However, this approach is most successful for organisms that had hard body parts, such as shells, bones or
teeth. Further, as prokaryotes such as bacteria and archaea share a limited set of common morphologies,
their fossils do not provide information on their ancestry.
More recently, evidence for common descent has come from the study of biochemical similarities
between organisms. For example, all living cells use the same basic set of nucleotides and amino acids.[230]
The development of molecular genetics has revealed the record of evolution left in organisms' genomes:
dating when species diverged through the molecular clock produced by mutations.[231] For example, these
DNA sequence comparisons have revealed that humans and chimpanzees share 96% of their genomes and
analyzing the few areas where they differ helps shed light on when the common ancestor of these species
existed.[232]

[edit] Evolution of life

For more details on this topic, see Timeline of evolution.
 Evolutionary tree showing the divergence of modern species from their common ancestor in the
center.[233] The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.
Despite the uncertainty on how life began, it is generally accepted that prokaryotes inhabited the
Earth from approximately 3–4 billion years ago.[234][235] No obvious changes in morphology or cellular
organization occurred in these organisms over the next few billion years.[236]
 The eukaryotes were the next major change in cell structure. These came from ancient bacteria
being engulfed by the ancestors of eukaryotic cells, in a cooperative association called endosymbiosis.[98]
[237] The engulfed bacteria and the host cell then underwent co-evolution, with the bacteria evolving into
either mitochondria or hydrogenosomes.[238] An independent second engulfment of cyanobacterial-like
organisms led to the formation of chloroplasts in algae and plants.[239] It is unknown when the first
eukaryotic cells appeared though they first emerged between 1.6 – 2.7 billion years ago.
The history of life was that of the unicellular eukaryotes, prokaryotes, and archaea until about 610
million years ago when multicellular organisms began to appear in the oceans in the Ediacaran period.[234]
[240] The evolution of multicellularity occurred in multiple independent events, in organisms as diverse as
sponges, brown algae, cyanobacteria, slime moulds and myxobacteria.[241]
Soon after the emergence of these first multicellular organisms, a remarkable amount of biological
diversity appeared over approximately 10 million years, in an event called the Cambrian explosion. Here, the
majority of types of modern animals appeared in the fossil record, as well as unique lineages that
subsequently became extinct.[242] Various triggers for the Cambrian explosion have been proposed,
including the accumulation of oxygen in the atmosphere from photosynthesis.[243] About 500 million years
ago, plants and fungi colonized the land, and were soon followed by arthropods and other animals.[244]
Insects were particularly successful and even today make up the majority of animal species.[245]
Amphibians first appeared around 300 million years ago, followed by early amniotes, then mammals around
200 million years ago and birds around 100 million years ago (both from "reptile"-like lineages). However,
despite the evolution of these large animals, smaller organisms similar to the types that evolved early in this
process continue to be highly successful and dominate the Earth, with the majority of both biomass and
species being prokaryotes.[137]
[edit] Applications
Further information: Artificial selection and Evolutionary computation
Evolutionary biology, and in particular the understanding of how organisms evolve through natural
selection, is an area of science with many practical applications.[246] A major technological application of
evolution is artificial selection, which is the intentional selection of certain traits in a population of organisms.
Humans have used artificial selection for thousands of years in the domestication of plants and animals.[247]
More recently, such selection has become a vital part of genetic engineering, with selectable markers such
as antibiotic resistance genes being used to manipulate DNA in molecular biology. It is also possible to use
repeated rounds of mutation and selection to evolve proteins with particular properties, such as modified
enzymes or new antibodies, in a process called directed evolution.[248]
Understanding the changes that have occurred during organism's evolution can reveal the genes
needed to construct parts of the body, genes which may be involved in human genetic disorders.[249] For
example, the Mexican tetra is an albino cavefish that lost its eyesight during evolution. Breeding together
different populations of this blind fish produced some offspring with functional eyes, since different mutations
had occurred in the isolated populations that had evolved in different caves.[250] This helped identify genes
required for vision and pigmentation, such as crystallins and the melanocortin 1 receptor.[251] Similarly,
comparing the genome of the Antarctic icefish, which lacks red blood cells, to close relatives such as the
zebrafish revealed genes needed to make these blood cells.[252]
As evolution can produce highly optimized processes and networks, it has many applications in
computer science. Here, simulations of evolution using evolutionary algorithms and artificial life started with
the work of Nils Aall Barricelli in the 1960s, and was extended by Alex Fraser, who published a series of
papers on simulation of artificial selection.[253] Artificial evolution became a widely recognized optimization
method as a result of the work of Ingo Rechenberg in the 1960s and early 1970s, who used evolution
strategies to solve complex engineering problems.[254] Genetic algorithms in particular became popular
through the writing of John Holland.[255] As academic interest grew, dramatic increases in the power of
computers allowed practical applications, including the automatic evolution of computer programs.[256]
Evolutionary algorithms are now used to solve multi-dimensional problems more efficiently than software
produced by human designers, and also to optimize the design of systems.[257]

[edit] Social and cultural responses

Further information: Social effect of evolutionary theory and Objections to evolution
 As Darwinism became widely accepted in the 1870s, caricatures of Charles Darwin with an ape or
monkey body symbolised evolution.[258]
In the 19th century, particularly after the publication of On the Origin of Species in 1859, the idea that
life had evolved was an active source of academic debate centered on the philosophical, social and religious
implications of evolution. Nowadays, the fact that organisms evolve is uncontested in the scientific literature
and the modern evolutionary synthesis is widely accepted by scientists.[14] However, evolution remains a
contentious concept for some theists.[259]
While various religions and denominations have reconciled their beliefs with evolution through
concepts such as theistic evolution, there are creationists who believe that evolution is contradicted by the
creation myths found in their respective religions and who raise various objections to evolution.[127][260]
[261] As had been demonstrated by responses to the publication of Vestiges of the Natural History of
Creation in 1844, the most controversial aspect of evolutionary biology is the implication of human evolution
that human mental and moral faculties, which had been thought purely spiritual, are not distinctly separated
from those of other animals.[8] In some countries—notably the United States—these tensions between science
and religion have fueled the current creation-evolution controversy, a religious conflict focusing on politics
and public education.[262] While other scientific fields such as cosmology[263] and earth science[264] also
conflict with literal interpretations of many religious texts, evolutionary biology experiences significantly more
opposition from religious literalists.
The teaching of evolution in American secondary school biology classes was uncommon in most of
the first half of the 20th century. The Scopes Trial decision of 1925 caused the subject to become very rare in
American secondary biology textbooks for a generation, but it was gradually re-introduced about a generation
later and legally protected with the 1968 Epperson v. Arkansas decision. Since then, the competing religious
belief of creationism was legally disallowed in secondary school curricula in various decisions in the 1970s
and 1980s, but it returned in the form of intelligent design, to be excluded once again in the 2005 Kitzmiller v.
Dover Area School District case.[265]
Another example somewhat associated with evolutionary theory that is now widely regarded as
unwarranted is "Social Darwinism", a derogatory term associated with the 19th century Malthusian theory
developed by Whig philosopher Herbert Spencer. It was later expanded by others into ideas about "survival
of the fittest" in commerce and human societies as a whole, and led to claims that social inequality, sexism,
racism, and imperialism were justified.[266] However, these ideas contradict Darwin's own views, and
contemporary scientists and philosophers consider these ideas to be neither mandated by evolutionary
theory nor supported by data.[267][268][269]

[edit] See also

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

[edit] References
1. ^ a b c Futuyma, Douglas J. (2005). Evolution. Sunderland, Massachusetts: Sinauer
Associates, Inc. ISBN 0-87893-187-2.
 2. ^ a b Walsh T (2006). "Combinatorial genetic evolution of multiresistance". Curr. Opin.
Microbiol. 9 (5): 476–82. doi:10.1016/j.mib.2006.08.009. PMID 16942901.
3. ^ Kottak, Conrad Phillip. Window on Humanity: A Concise Introduction to Anthropology. New
York: The McGraw–Hill Companies, Inc, 2005
4. ^ http://www.mhest.com/spotlight/darwin/articles/OrganicEvolution.pdf
5. ^ a b Lande R, Arnold SJ (1983). "The measurement of selection on correlated characters".
Evolution 37 (6): 1210–26. doi:10.2307/2408842. http://jstor.org/stable/2408842.
6. ^ Ayala FJ (2007). "Darwin's greatest discovery: design without designer". Proc. Natl. Acad.
Sci. U.S.A. 104 (Suppl 1): 8567–73. doi:10.1073/pnas.0701072104. PMID 17494753. PMC 1876431.
http://www.pnas.org/content/104/suppl.1/8567.full.
7. ^ Ian C. Johnston (1999). "History of Science: Early Modern Geology". Malaspina University-
College. http://records.viu.ca/~johnstoi/darwin/sect2.htm. Retrieved 2008-01-15.
8. ^ a b Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press.
ISBN 0-52023693-9.
9. ^ a b Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray. p. 1.
ISBN 0801413192. http://darwin-online.org.uk/content/frameset?
itemID=F373&viewtype=text&pageseq=16. . Related earlier ideas were acknowledged in Darwin,
Charles (1861). On the Origin of Species (3rd ed.). London: John Murray. xiii. ISBN 0801413192.
http://darwin-online.org.uk/content/frameset?itemID=F381&viewtype=text&pageseq=20.
10.^ AAAS Council (December 26, 1922). "AAAS Resolution: Present Scientific Status of the
Theory of Evolution". American Association for the Advancement of Science.
http://archives.aaas.org/docs/resolutions.php?doc_id=450.
 11.^ a b "IAP Statement on the Teaching of Evolution" (PDF). The Interacademy Panel on
International Issues. 2006. http://www.interacademies.net/Object.File/Master/6/150/Evolution
%20statement.pdf. Retrieved 2007-04-25. Joint statement issued by the national science academies
of 67 countries, including the United Kingdom's Royal Society
12.^ a b Board of Directors, American Association for the Advancement of Science (2006-02-16).
"Statement on the Teaching of Evolution" (PDF). American Association for the Advancement of
Science. http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf. from the world's
largest general scientific society
13.^ "Statements from Scientific and Scholarly Organizations". National Center for Science
Education. http://ncse.com/media/voices/science.
14.^ a b c d e Kutschera U, Niklas K (2004). "The modern theory of biological evolution: an
expanded synthesis". Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y.
PMID 15241603.
15.^ "Special report on evolution". New Scientist. 2008-01-19.
http://www.newscientist.com/topic/evolution.
16.^ Wright, S (1984). Evolution and the Genetics of Populations, Volume 1: Genetic and
Biometric Foundations. The University of Chicago Press. ISBN 0-226-91038-5.
17.^ Zirkle C (1941). "Natural Selection before the "Origin of Species"". Proceedings of the
American Philosophical Society 84 (1): 71–123.
18.^ Muhammad Hamidullah and Afzal Iqbal (1993), The Emergence of Islam: Lectures on the
Development of Islamic World-view, Intellectual Tradition and Polity , p. 143-144. Islamic Research
Institute, Islamabad.
19.^ "A Source Book In Chinese Philosophy", Chan, Wing-Tsit, p. 204, 1962.
 20.^ Terrall, M (2002). The Man Who Flattened the Earth: Maupertuis and the Sciences in the
Enlightenment. The University of Chicago Press. ISBN 978-0226793610.
21.^ Wallace, A (1858). "On the Tendency of Species to form Varieties, and on the Perpetuation
of Varieties and Species by Natural Means of Selection". Journal of the Proceedings of the Linnean
Society of London. Zoology 3: 53–62. doi:10.1098/rsnr.2006.0171. http://darwin-
online.org.uk/content/frameset?itemID=F350&viewtype=text&pageseq=1. Retrieved 2007-05-13.
22.^ Darwin, Charles (1872). "Effects of the increased Use and Disuse of Parts, as controlled by
Natural Selection". The Origin of Species. 6th edition, p. 108. John Murray. http://darwin-
online.org.uk/content/frameset?viewtype=text&itemID=F391&pageseq=136. Retrieved 2007-12-28.
23.^ Leakey, Richard E.; Darwin, Charles (1979). The illustrated origin of species. London:
Faber. ISBN 0-571-14586-8. p. 17-18
24.^ Ghiselin, Michael T. (September/October 1994). "Nonsense in schoolbooks: 'The Imaginary
Lamarck'". The Textbook Letter. The Textbook League. http://www.textbookleague.org/54marck.htm.
Retrieved 2008-01-23.
25.^ Magner, Lois N. (2002). A History of the Life Sciences (Third ed.). Marcel Dekker, CRC
Press. ISBN 9780203911006. http://books.google.com/?
id=YKJ6gVYbrGwC&printsec=frontcover#v=onepage&q.
26.^ Weiling F (1991). "Historical study: Johann Gregor Mendel 1822–1884". Am. J. Med. Genet.
40 (1): 1–25; discussion 26. doi:10.1002/ajmg.1320400103. PMID 1887835.
27.^ Quammen, D. (2006). The reluctant Mr. Darwin: An intimate portrait of Charles Darwin and
the making of his theory of evolution. New York, NY: W.W. Norton & Company.
28.^ Bowler, Peter J. (1989). The Mendelian Revolution: The Emergence of Hereditarian
Concepts in Modern Science and Society. Baltimore: Johns Hopkins University Press. ISBN 978-
0801838880.
 29.^ Pigliucci M (December 2007). "Do we need an extended evolutionary synthesis?".
Evolution 61 (12): 2743–9. doi:10.1111/j.1558-5646.2007.00246.x. PMID 17924956.
30.^ Winther RG (August 2008). "Systemic darwinism". Proc. Natl. Acad. Sci. U.S.A. 105 (33):
11833–8. doi:10.1073/pnas.0711445105. PMID 18697926.
31.^ Sturm RA, Frudakis TN (2004). "Eye colour: portals into pigmentation genes and ancestry".
Trends Genet. 20 (8): 327–32. doi:10.1016/j.tig.2004.06.010. PMID 15262401.
32.^ a b Pearson H (2006). "Genetics: what is a gene?". Nature 441 (7092): 398–401.
doi:10.1038/441398a. PMID 16724031.
33.^ Visscher PM, Hill WG, Wray NR (April 2008). "Heritability in the genomics era—concepts
and misconceptions". Nat. Rev. Genet. 9 (4): 255–66. doi:10.1038/nrg2322. PMID 18319743.
34.^ Oetting WS, Brilliant MH, King RA (1996). "The clinical spectrum of albinism in humans".
Molecular medicine today 2 (8): 330–5. doi:10.1016/1357-4310(96)81798-9. PMID 8796918.
35.^ Phillips PC (November 2008). "Epistasis—the essential role of gene interactions in the
structure and evolution of genetic systems". Nat. Rev. Genet. 9 (11): 855–67. doi:10.1038/nrg2452.
PMID 18852697.
36.^ a b Wu R, Lin M (2006). "Functional mapping – how to map and study the genetic
architecture of dynamic complex traits". Nat. Rev. Genet. 7 (3): 229–37. doi:10.1038/nrg1804.
PMID 16485021.
37.^ Richards EJ (May 2006). "Inherited epigenetic variation—revisiting soft inheritance". Nat.
Rev. Genet. 7 (5): 395–401. doi:10.1038/nrg1834. PMID 16534512.
38.^ a b Harwood AJ; Harwood, J (1998). "Factors affecting levels of genetic diversity in natural
populations". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 353 (1366): 177–86.
doi:10.1098/rstb.1998.0200. PMID 9533122.
 39.^ Draghi J, Turner P (2006). "DNA secretion and gene-level selection in bacteria".
Microbiology (Reading, Engl.) 152 (Pt 9): 2683–8. doi:10.1099/mic.0.29013-0. PMID 16946263.
*Mallet J (2007). "Hybrid speciation". Nature 446 (7133): 279–83. doi:10.1038/nature05706.
PMID 17361174.
40.^ Butlin RK, Tregenza T (1998). "Levels of genetic polymorphism: marker loci versus
quantitative traits". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 353 (1366): 187–98.
doi:10.1098/rstb.1998.0201. PMID 9533123.
41.^ Wetterbom A, Sevov M, Cavelier L, Bergström TF (2006). "Comparative genomic analysis
of human and chimpanzee indicates a key role for indels in primate evolution". J. Mol. Evol. 63 (5):
682–90. doi:10.1007/s00239-006-0045-7. PMID 17075697.
42.^ a b c Bertram J (2000). "The molecular biology of cancer". Mol. Aspects Med. 21 (6): 167–
223. doi:10.1016/S0098-2997(00)00007-8. PMID 11173079.
43.^ a b Aminetzach YT, Macpherson JM, Petrov DA (2005). "Pesticide resistance via
transposition-mediated adaptive gene truncation in Drosophila". Science 309 (5735): 764–7.
doi:10.1126/science.1112699. PMID 16051794.
44.^ Burrus V, Waldor M (2004). "Shaping bacterial genomes with integrative and conjugative
elements". Res. Microbiol. 155 (5): 376–86. doi:10.1016/j.resmic.2004.01.012. PMID 15207870.
45.^ Sawyer SA, Parsch J, Zhang Z, Hartl DL (2007). "Prevalence of positive selection among
nearly neutral amino acid replacements in Drosophila". Proc. Natl. Acad. Sci. U.S.A. 104 (16): 6504–
10. doi:10.1073/pnas.0701572104. PMID 17409186.
46.^ Sniegowski P, Gerrish P, Johnson T, Shaver A (2000). "The evolution of mutation rates:
separating causes from consequences". Bioessays 22 (12): 1057–66. doi:10.1002/1521-
1878(200012)22:12<1057::AID-BIES3>3.0.CO;2-W. PMID 11084621.
 47.^ Drake JW, Holland JJ (1999). "Mutation rates among RNA viruses". Proc. Natl. Acad. Sci.
U.S.A. 96 (24): 13910–3. doi:10.1073/pnas.96.24.13910. PMID 10570172. PMC 24164.
http://www.pnas.org/content/96/24/13910.long.
48.^ Holland J, Spindler K, Horodyski F, Grabau E, Nichol S, VandePol S (1982). "Rapid
evolution of RNA genomes". Science 215 (4540): 1577–85. doi:10.1126/science.7041255.
PMID 7041255.
49.^ Hastings, P J; Lupski, JR; Rosenberg, SM; Ira, G (2009). "Mechanisms of change in gene
copy number". Nature Reviews. Genetics 10 (8): 551–564. doi:10.1038/nrg2593. PMID 19597530.
50.^ Carroll SB, Grenier J, Weatherbee SD (2005). From DNA to Diversity: Molecular Genetics
and the Evolution of Animal Design. Second Edition . Oxford: Blackwell Publishing. ISBN 1-4051-
1950-0.
51.^ Harrison P, Gerstein M (2002). "Studying genomes through the aeons: protein families,
pseudogenes and proteome evolution". J Mol Biol 318 (5): 1155–74. doi:10.1016/S0022-
2836(02)00109-2. PMID 12083509.
52.^ Bowmaker JK (1998). "Evolution of colour vision in vertebrates". Eye (London, England) 12
(Pt 3b): 541–7. PMID 9775215.
53.^ Gregory TR, Hebert PD (1999). "The modulation of DNA content: proximate causes and
ultimate consequences". Genome Res. 9 (4): 317–24. doi:10.1101/gr.9.4.317 (inactive 2009-11-14).
PMID 10207154. http://genome.cshlp.org/content/9/4/317.full.
54.^ Hurles M (July 2004). "Gene duplication: the genomic trade in spare parts". PLoS Biol. 2
(7): E206. doi:10.1371/journal.pbio.0020206. PMID 15252449.
55.^ Liu N, Okamura K, Tyler DM (2008). "The evolution and functional diversification of animal
microRNA genes". Cell Res. 18 (10): 985–96. doi:10.1038/cr.2008.278. PMID 18711447.
PMC 2712117. http://www.nature.com/cr/journal/v18/n10/full/cr2008278a.html.
 56.^ Siepel A (October 2009). "Darwinian alchemy: Human genes from noncoding DNA".
Genome Res. 19 (10): 1693–5. doi:10.1101/gr.098376.109. PMID 19797681. PMC 2765273.
http://genome.cshlp.org/content/19/10/1693.full.
57.^ Orengo CA, Thornton JM (2005). "Protein families and their evolution-a structural
perspective". Annu. Rev. Biochem. 74: 867–900. doi:10.1146/annurev.biochem.74.082803.133029.
PMID 15954844.
58.^ Long M, Betrán E, Thornton K, Wang W (November 2003). "The origin of new genes:
glimpses from the young and old". Nat. Rev. Genet. 4 (11): 865–75. doi:10.1038/nrg1204.
PMID 14634634.
59.^ Wang M, Caetano-Anollés G (2009). "The evolutionary mechanics of domain organization
in proteomes and the rise of modularity in the protein world". Structure 17 (1): 66–78.
doi:10.1016/j.str.2008.11.008. PMID 19141283.
60.^ Weissman KJ, Müller R (April 2008). "Protein-protein interactions in multienzyme
megasynthetases". Chembiochem 9 (6): 826–48. doi:10.1002/cbic.200700751. PMID 18357594.
61.^ Zhang J, Wang X, Podlaha O (2004). "Testing the chromosomal speciation hypothesis for
humans and chimpanzees". Genome Res. 14 (5): 845–51. doi:10.1101/gr.1891104. PMID 15123584.
62.^ Ayala FJ, Coluzzi M (2005). "Chromosome speciation: humans, Drosophila, and
mosquitoes". Proc. Natl. Acad. Sci. U.S.A. 102 (Suppl 1): 6535–42. doi:10.1073/pnas.0501847102.
PMID 15851677. PMC 1131864. http://www.pnas.org/content/102/suppl.1/6535.full.
63.^ Hurst GD, Werren JH (2001). "The role of selfish genetic elements in eukaryotic evolution".
Nat. Rev. Genet. 2 (8): 597–606. doi:10.1038/35084545. PMID 11483984.
64.^ Häsler J, Strub K (2006). "Alu elements as regulators of gene expression". Nucleic Acids
Res. 34 (19): 5491–7. doi:10.1093/nar/gkl706. PMID 17020921.
 65.^ Radding C (1982). "Homologous pairing and strand exchange in genetic recombination".
Annu. Rev. Genet. 16: 405–37. doi:10.1146/annurev.ge.16.120182.002201. PMID 6297377.
66.^ Agrawal AF (2006). "Evolution of sex: why do organisms shuffle their genotypes?". Curr.
Biol. 16 (17): R696. doi:10.1016/j.cub.2006.07.063. PMID 16950096.
67.^ Peters AD, Otto SP (2003). "Liberating genetic variance through sex". Bioessays 25 (6):
533–7. doi:10.1002/bies.10291. PMID 12766942.
68.^ Goddard MR, Godfray HC, Burt A (2005). "Sex increases the efficacy of natural selection in
experimental yeast populations". Nature 434 (7033): 636–40. doi:10.1038/nature03405.
PMID 15800622.
69.^ Fontaneto D, Herniou EA, Boschetti C (April 2007). "Independently evolving species in
asexual bdelloid rotifers". PLoS Biol. 5 (4): e87. doi:10.1371/journal.pbio.0050087. PMID 17373857.
Lay summary.
70.^ Pouchkina-Stantcheva NN, McGee BM, Boschetti C (October 2007). "Functional
divergence of former alleles in an ancient asexual invertebrate". Science 318 (5848): 268–71.
doi:10.1126/science.1144363. PMID 17932297. Lay summary.
71.^ Lien S, Szyda J, Schechinger B, Rappold G, Arnheim N (February 2000). "Evidence for
heterogeneity in recombination in the human pseudoautosomal region: high resolution analysis by
sperm typing and radiation-hybrid mapping". Am. J. Hum. Genet. 66 (2): 557–66.
doi:10.1086/302754. PMID 10677316.
72.^ Barton, N H (2000). "Genetic hitchhiking". Philosophical Transactions of the Royal Society
of London. Series B, Biological Sciences 355 (1403): 1553–1562. doi:10.1098/rstb.2000.0716.
PMID 11127900. PMC 1692896. http://www.pubmedcentral.nih.gov/picrender.fcgi?
artid=1692896&blobtype=pdf. Retrieved 2009-09-23.
 73.^ Muller H (1964). "The relation of recombination to mutational advance". Mutat. Res. 106: 2–
9. PMID 14195748.
74.^ Charlesworth B, Charlesworth D (November 2000). "The degeneration of Y chromosomes".
Philos. Trans. R. Soc. Lond., B, Biol. Sci. 355 (1403): 1563–72. doi:10.1098/rstb.2000.0717.
PMID 11127901.
75.^ a b Otto S (1 July 2003). "The advantages of segregation and the evolution of sex".
Genetics 164 (3): 1099–118. PMID 12871918.
76.^ Doncaster CP, Pound GE, Cox SJ (March 2000). "The ecological cost of sex". Nature 404
(6775): 281–5. doi:10.1038/35005078. PMID 10749210.
77.^ Butlin R (April 2002). "Evolution of sex: The costs and benefits of sex: new insights from old
asexual lineages". Nat. Rev. Genet. 3 (4): 311–7. doi:10.1038/nrg749. PMID 11967555.
78.^ Salathé M, Kouyos RD, Bonhoeffer S (August 2008). "The state of affairs in the kingdom of
the Red Queen". Trends Ecol. Evol. (Amst.) 23 (8): 439–45. doi:10.1016/j.tree.2008.04.010.
PMID 18597889.
79.^ Stoltzfus A (2006). "Mutationism and the dual causation of evolutionary change". Evol.
Dev. 8 (3): 304–17. doi:10.1111/j.1525-142X.2006.00101.x. PMID 16686641.
80.^ O'Neil, Dennis (2008). "Hardy-Weinberg Equilibrium Model". The synthetic theory of
evolution: An introduction to modern evolutionary concepts and theories . Behavioral Sciences
Department, Palomar College. http://anthro.palomar.edu/synthetic/synth_2.htm. Retrieved 2008-01-
06.
81.^ Bright, Kerry (2006). "Causes of evolution". Teach Evolution and Make It Relevant. National
Science Foundation. http://evoled.dbs.umt.edu/lessons/causes.htm#hardy. Retrieved 2007-12-30.
82.^ Packer C, Gilbert DA, Pusey AE, O'Brieni SJ. (June 1991). "A molecular genetic analysis of
kinship and cooperation in African lions". Nature 351: 562–65. doi:10.1038/351562a0.
 83.^ Morjan C, Rieseberg L (2004). "How species evolve collectively: implications of gene flow
and selection for the spread of advantageous alleles". Mol. Ecol. 13 (6): 1341–56. doi:10.1111/j.1365-
294X.2004.02164.x. PMID 15140081.
84.^ Su H, Qu L, He K, Zhang Z, Wang J, Chen Z, Gu H (2003). "The Great Wall of China: a
physical barrier to gene flow?". Heredity 90 (3): 212–9. doi:10.1038/sj.hdy.6800237.
PMID 12634804.
85.^ Short RV (1975). "The contribution of the mule to scientific thought". J. Reprod. Fertil.
Suppl. (23): 359–64. PMID 1107543.
86.^ Gross B, Rieseberg L (2005). "The ecological genetics of homoploid hybrid speciation". J.
Hered. 96 (3): 241–52. doi:10.1093/jhered/esi026. PMID 15618301.
87.^ Burke JM, Arnold ML (2001). "Genetics and the fitness of hybrids". Annu. Rev. Genet. 35:
31–52. doi:10.1146/annurev.genet.35.102401.085719. PMID 11700276.
88.^ Vrijenhoek RC (2006). "Polyploid hybrids: multiple origins of a treefrog species". Curr. Biol.
16 (7): R245. doi:10.1016/j.cub.2006.03.005. PMID 16581499.
89.^ Wendel J (2000). "Genome evolution in polyploids". Plant Mol. Biol. 42 (1): 225–49.
doi:10.1023/A:1006392424384. PMID 10688139.
90.^ a b Sémon M, Wolfe KH (2007). "Consequences of genome duplication". Curr Opin Genet
Dev 17 (6): 505–12. doi:10.1016/j.gde.2007.09.007. PMID 18006297.
91.^ Comai L (2005). "The advantages and disadvantages of being polyploid". Nat. Rev. Genet.
6 (11): 836–46. doi:10.1038/nrg1711. PMID 16304599.
92.^ Soltis P, Soltis D (June 2000). "The role of genetic and genomic attributes in the success of
polyploids". Proc. Natl. Acad. Sci. U.S.A. 97 (13): 7051–7. doi:10.1073/pnas.97.13.7051.
PMID 10860970.
 93.^ Boucher Y, Douady CJ, Papke RT, Walsh DA, Boudreau ME, Nesbo CL, Case RJ, Doolittle
WF (2003). "Lateral gene transfer and the origins of prokaryotic groups". Annu Rev Genet 37: 283–
328. doi:10.1146/annurev.genet.37.050503.084247. PMID 14616063.
94.^ Kondo N, Nikoh N, Ijichi N, Shimada M, Fukatsu T (2002). "Genome fragment of Wolbachia
endosymbiont transferred to X chromosome of host insect". Proc. Natl. Acad. Sci. U.S.A. 99 (22):
14280–5. doi:10.1073/pnas.222228199. PMID 12386340.
95.^ Sprague G (1991). "Genetic exchange between kingdoms". Curr. Opin. Genet. Dev. 1 (4):
530–3. doi:10.1016/S0959-437X(05)80203-5. PMID 1822285.
96.^ Gladyshev EA, Meselson M, Arkhipova IR (May 2008). "Massive horizontal gene transfer in
bdelloid rotifers". Science 320 (5880): 1210–3. doi:10.1126/science.1156407. PMID 18511688.
97.^ Baldo A, McClure M (1 September 1999). "Evolution and horizontal transfer of dUTPase-
encoding genes in viruses and their hosts". J. Virol. 73 (9): 7710–21. PMID 10438861.
98.^ a b Poole A, Penny D (2007). "Evaluating hypotheses for the origin of eukaryotes".
Bioessays 29 (1): 74–84. doi:10.1002/bies.20516. PMID 17187354.
99.^ Whitlock M (1 June 2003). "Fixation probability and time in subdivided populations".
Genetics 164 (2): 767–79. PMID 12807795.
100.^ Ohta T (2002). "Near-neutrality in evolution of genes and gene regulation". Proc. Natl.
Acad. Sci. U.S.A. 99 (25): 16134–7. doi:10.1073/pnas.252626899. PMID 12461171. PMC 138577.
http://www.pnas.org/cgi/content/abstract/252626899v1.
101.^ a b c Hurst LD (February 2009). "Fundamental concepts in genetics: genetics and the
understanding of selection". Nat. Rev. Genet. 10 (2): 83–93. doi:10.1038/nrg2506. PMID 19119264.
102.^ a b c Orr HA (August 2009). "Fitness and its role in evolutionary genetics". Nat. Rev.
Genet. 10 (8): 531–9. doi:10.1038/nrg2603. PMID 19546856.
 103.^ Haldane J (1959). "The theory of natural selection today". Nature 183 (4663): 710–3.
doi:10.1038/183710a0. PMID 13644170.
104.^ Goldberg, Emma E; Igić, B (2008). "On phylogenetic tests of irreversible evolution".
Evolution 62 (11): 2727–2741. doi:10.1111/j.1558-5646.2008.00505.x. PMID 18764918.
105.^ Collin, Rachel; Miglietta, MP (2008). "Reversing opinions on Dollo's Law". Trends in
Ecology & Evolution 23 (11): 602–609. doi:10.1016/j.tree.2008.06.013. PMID 18814933.
106.^ Hoekstra H, Hoekstra J, Berrigan D, Vignieri S, Hoang A, Hill C, Beerli P, Kingsolver J
(July 2001). "Strength and tempo of directional selection in the wild". Proc. Natl. Acad. Sci. U.S.A. 98
(16): 9157–60. doi:10.1073/pnas.161281098. PMID 11470913.
107.^ Felsenstein (1 November 1979). "Excursions along the Interface between Disruptive and
Stabilizing Selection". Genetics 93 (3): 773–95. PMID 17248980.
108.^ Andersson M, Simmons L (2006). "Sexual selection and mate choice". Trends Ecol. Evol.
(Amst.) 21 (6): 296–302. doi:10.1016/j.tree.2006.03.015. PMID 16769428.
109.^ Kokko H, Brooks R, McNamara J, Houston A (2002). "The sexual selection continuum".
Proc. Biol. Sci. 269 (1498): 1331–40. doi:10.1098/rspb.2002.2020. PMID 12079655.
110.^ Hunt J, Brooks R, Jennions M, Smith M, Bentsen C, Bussière L (2004). "High-quality male
field crickets invest heavily in sexual display but die young". Nature 432 (7020): 1024–7.
doi:10.1038/nature03084. PMID 15616562.
111.^ Odum, EP (1971) Fundamentals of ecology, third edition, Saunders New York
112.^ a b Gould SJ (February 1998). "Gulliver's further travels: the necessity and difficulty of a
hierarchical theory of selection". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 353 (1366): 307–14.
doi:10.1098/rstb.1998.0211. PMID 9533127.
113.^ Mayr E (1997). "The objects of selection". Proc. Natl. Acad. Sci. U.S.A. 94 (6): 2091–4.
doi:10.1073/pnas.94.6.2091. PMID 9122151.
 114.^ Maynard Smith J (1998). "The units of selection". Novartis Found. Symp. 213: 203–11;
discussion 211–7. PMID 9653725.
115.^ Hickey DA (1992). "Evolutionary dynamics of transposable elements in prokaryotes and
eukaryotes". Genetica 86 (1–3): 269–74. doi:10.1007/BF00133725. PMID 1334911.
116.^ Gould SJ, Lloyd EA (1999). "Individuality and adaptation across levels of selection: how
shall we name and generalize the unit of Darwinism?". Proc. Natl. Acad. Sci. U.S.A. 96 (21): 11904–
9. doi:10.1073/pnas.96.21.11904. PMID 10518549.
117.^ Lande R (1989). "Fisherian and Wrightian theories of speciation". Genome 31 (1): 221–7.
PMID 2687093.
118.^ Otto S, Whitlock M (1 June 1997). "The probability of fixation in populations of changing
size". Genetics 146 (2): 723–33. PMID 9178020.
119.^ Charlesworth B (March 2009). "Fundamental concepts in genetics: Effective population
size and patterns of molecular evolution and variation". Nat. Rev. Genet. 10 (3): 195–205.
doi:10.1038/nrg2526. PMID 19204717.
120.^ Protas, Meredith; Conrad, M; Gross, JB; Tabin, C; Borowsky, R (2007). "Regressive
evolution in the Mexican cave tetra, Astyanax mexicanus". Current Biology 17 (5): 452–454.
doi:10.1016/j.cub.2007.01.051. PMID 17306543. PMC 2570642.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=17306543#R2.
121.^ Mitchell-Olds, Thomas; Willis, JH; Goldstein, DB (2007). "Which evolutionary processes
influence natural genetic variation for phenotypic traits?". Nature Reviews Genetics 8 (11): 845–856.
doi:10.1038/nrg2207. PMID 17943192.
122.^ Nei M (2005). "Selectionism and neutralism in molecular evolution". Mol. Biol. Evol. 22
(12): 2318–42. doi:10.1093/molbev/msi242. PMID 16120807.
 123.^ Kimura M (1991). "The neutral theory of molecular evolution: a review of recent evidence".
Jpn. J. Genet. 66 (4): 367–86. doi:10.1266/jjg.66.367. PMID 1954033.
http://www.jstage.jst.go.jp/article/jjg/66/4/66_367/_article.
124.^ Kimura M (1989). "The neutral theory of molecular evolution and the world view of the
neutralists". Genome 31 (1): 24–31. PMID 2687096.
125.^ Kreitman M (August 1996). "The neutral theory is dead. Long live the neutral theory".
Bioessays 18 (8): 678–83; discussion 683. doi:10.1002/bies.950180812. PMID 8760341.
http://www.cs.ucsb.edu/~ambuj/Courses/bioinformatics/neutral-theory.
126.^ Leigh E.G. (Jr) (2007). "Neutral theory: a historical perspective.". Journal of Evolutionary
Biology 20 (6): 2075–91. doi:10.1111/j.1420-9101.2007.01410.x. PMID 17956380.
127.^ a b c Scott EC, Matzke NJ (May 2007). "Biological design in science classrooms". Proc.
Natl. Acad. Sci. U.S.A. 104 Suppl 1: 8669–76. doi:10.1073/pnas.0701505104. PMID 17494747.
128.^ Hendry AP, Kinnison MT (2001). "An introduction to microevolution: rate, pattern,
process". Genetica 112–113: 1–8. doi:10.1023/A:1013368628607. PMID 11838760.
129.^ Leroi AM (2000). "The scale independence of evolution". Evol. Dev. 2 (2): 67–77.
doi:10.1046/j.1525-142x.2000.00044.x. PMID 11258392.
130.^ Gould 2002, pp. 657–8
131.^ Gould SJ (July 1994). "Tempo and mode in the macroevolutionary reconstruction of
Darwinism". Proc. Natl. Acad. Sci. U.S.A. 91 (15): 6764–71. doi:10.1073/pnas.91.15.6764.
PMID 8041695. PMC 44281. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=8041695.
132.^ Jablonski, D. (2000). "Micro- and macroevolution: scale and hierarchy in evolutionary
biology and paleobiology". Paleobiology 26 (sp4): 15–52. doi:10.1666/0094-
8373(2000)26[15:MAMSAH]2.0.CO;2. http://www.bioone.org/perlserv/?request=get-abstract.
 133.^ Michael J. Dougherty. Is the human race evolving or devolving? Scientific American July
20, 1998.
134.^ TalkOrigins Archive response to Creationist claims – Claim CB932: Evolution of
degenerate forms
135.^ Carroll SB (2001). "Chance and necessity: the evolution of morphological complexity and
diversity". Nature 409 (6823): 1102–9. doi:10.1038/35059227. PMID 11234024.
136.^ Whitman W, Coleman D, Wiebe W (1998). "Prokaryotes: the unseen majority". Proc Natl
Acad Sci U S a 95 (12): 6578–83. doi:10.1073/pnas.95.12.6578. PMID 9618454.
137.^ a b Schloss P, Handelsman J (December 2004). "Status of the microbial census".
Microbiol Mol Biol Rev 68 (4): 686–91. doi:10.1128/MMBR.68.4.686-691.2004. PMID 15590780.
138.^ Nealson K (1999). "Post-Viking microbiology: new approaches, new data, new insights".
Orig Life Evol Biosph 29 (1): 73–93. doi:10.1023/A:1006515817767. PMID 11536899.
139.^ Buckling A, Craig Maclean R, Brockhurst MA, Colegrave N (February 2009). "The Beagle
in a bottle". Nature 457 (7231): 824–9. doi:10.1038/nature07892. PMID 19212400.
140.^ Elena SF, Lenski RE (June 2003). "Evolution experiments with microorganisms: the
dynamics and genetic bases of adaptation". Nat. Rev. Genet. 4 (6): 457–69. doi:10.1038/nrg1088.
PMID 12776215.
141.^ Williams, George C. 1966. Adaptation and natural selection: a critique of some current
evolutionary thought. Princeton. "Evolutionary adaptation is a phenomenon of pervasive importance
in biology." p5
142.^ Mayr, Ernst 1982. The growth of biological thought. Harvard. p483: "Adaptation... could no
longer be considered a static condition, a product of a creative past, and became instead a
continuing dynamic process."
 143.^ The Oxford Dictionary of Science defines adaptation as "Any change in the structure or
functioning of an organism that makes it better suited to its environment".
144.^ Orr H (2005). "The genetic theory of adaptation: a brief history". Nat. Rev. Genet. 6 (2):
119–27. doi:10.1038/nrg1523. PMID 15716908.
145.^ Dobzhansky T. 1968. On some fundamental concepts of evolutionary biology.
Evolutionary biology 2, 1–34.
146.^ Dobzhansky T. 1970. Genetics of the evolutionary process. Columbia, N.Y. p4–6, 79–82,
84–87
147.^ Dobzhansky T. 1956. Genetics of natural populations XXV. Genetic changes in
populations of Drosophila pseudoobscura and Drosphila persimilis in some locations in California.
Evolution 10, 82–92.
148.^ Nakajima A, Sugimoto Y, Yoneyama H, Nakae T (2002). "High-level fluoroquinolone
resistance in Pseudomonas aeruginosa due to interplay of the MexAB-OprM efflux pump and the
DNA gyrase mutation". Microbiol. Immunol. 46 (6): 391–5. PMID 12153116.
http://www.jstage.jst.go.jp/article/mandi/46/6/46_391/_article/-char/en.
149.^ Blount ZD, Borland CZ, Lenski RE (June 2008). "Inaugural Article: Historical contingency
and the evolution of a key innovation in an experimental population of Escherichia coli". Proc. Natl.
Acad. Sci. U.S.A. 105 (23): 7899–906. doi:10.1073/pnas.0803151105. PMID 18524956.
PMC 2430337. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=18524956.
150.^ Okada H, Negoro S, Kimura H, Nakamura S (1983). "Evolutionary adaptation of plasmid-
encoded enzymes for degrading nylon oligomers". Nature 306 (5939): 203–6. doi:10.1038/306203a0.
PMID 6646204.
doi:10.1073/pnas.81.8.2421. PMID 6585807. PMC 345072. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=6585807.
152.^ Copley SD (June 2000). "Evolution of a metabolic pathway for degradation of a toxic
xenobiotic: the patchwork approach". Trends Biochem. Sci. 25 (6): 261–5. doi:10.1016/S0968-
0004(00)01562-0. PMID 10838562.
153.^ Crawford RL, Jung CM, Strap JL (October 2007). "The recent evolution of
pentachlorophenol (PCP)-4-monooxygenase (PcpB) and associated pathways for bacterial
degradation of PCP". Biodegradation 18 (5): 525–39. doi:10.1007/s10532-006-9090-6.
PMID 17123025.
154.^ Colegrave N, Collins S (May 2008). "Experimental evolution: experimental evolution and
evolvability". Heredity 100 (5): 464–70. doi:10.1038/sj.hdy.6801095. PMID 18212804.
http://www.nature.com/hdy/journal/v100/n5/full/6801095a.html.
155.^ Kirschner M, Gerhart J (July 1998). "Evolvability". Proc. Natl. Acad. Sci. U.S.A. 95 (15):
8420–7. doi:10.1073/pnas.95.15.8420. PMID 9671692.
156.^ a b Bejder L, Hall BK (2002). "Limbs in whales and limblessness in other vertebrates:
mechanisms of evolutionary and developmental transformation and loss". Evol. Dev. 4 (6): 445–58.
doi:10.1046/j.1525-142X.2002.02033.x. PMID 12492145.
157.^ Young, Nathan M.; Hallgrímsson, B (2005). "Serial homology and the evolution of
mammalian limb covariation structure". Evolution 59 (12): 2691. doi:10.1554/05-233.1.
PMID 16526515. http://www.bioone.org/doi/abs/10.1554/05-233.1. Retrieved 2009-09-24.
158.^ a b Penny D, Poole A (1999). "The nature of the last universal common ancestor". Curr.
Opin. Genet. Dev. 9 (6): 672–77. doi:10.1016/S0959-437X(99)00020-9. PMID 10607605.
159.^ Hall, Brian K (2003). "Descent with modification: the unity underlying homology and
homoplasy as seen through an analysis of development and evolution". Biological Reviews of the
Cambridge Philosophical Society 78 (3): 409–433. doi:10.1017/S1464793102006097.
PMID 14558591.
160.^ Shubin, Neil; Tabin, C; Carroll, S (2009). "Deep homology and the origins of evolutionary
novelty". Nature 457 (7231): 818–823. doi:10.1038/nature07891. PMID 19212399.
161.^ a b c Fong D, Kane T, Culver D (1995). "Vestigialization and Loss of Nonfunctional
Characters". Ann. Rev. Ecol. Syst. 26: 249–68. doi:10.1146/annurev.es.26.110195.001341.
162.^ Zhang Z, Gerstein M (August 2004). "Large-scale analysis of pseudogenes in the human
genome". Curr. Opin. Genet. Dev. 14 (4): 328–35. doi:10.1016/j.gde.2004.06.003. PMID 15261647.
163.^ Jeffery WR (2005). "Adaptive evolution of eye degeneration in the Mexican blind
cavefish". J. Hered. 96 (3): 185–96. doi:10.1093/jhered/esi028. PMID 15653557.
164.^ Maxwell EE, Larsson HC (2007). "Osteology and myology of the wing of the Emu
(Dromaius novaehollandiae), and its bearing on the evolution of vestigial structures". J. Morphol. 268
(5): 423–41. doi:10.1002/jmor.10527. PMID 17390336.
165.^ Silvestri AR, Singh I (2003). "The unresolved problem of the third molar: would people be
better off without it?". Journal of the American Dental Association (1939) 134 (4): 450–5.
doi:10.1146/annurev.es.26.110195.001341. PMID 12733778.
http://jada.ada.org/cgi/content/full/134/4/450.
166.^ Darwin, Charles. (1872) The Expression of the Emotions in Man and Animals John
Murray, London.
167.^ Peter Gray (2007). Psychology (fifth ed.). Worth Publishers. pp. 66. ISBN 0-7167-0617-2.
168.^ Behavior Development in Infants (via Google Books) by Evelyn Dewey, citing a study
"Reflexes and other motor activities in newborn infants: a report of 125 cases as a preliminary study
of infant behavior" published in the Bull. Neurol. Inst. New York, 1932, Vol. 2, pp. 1-56.
 169.^ Jerry Coyne (2009). Why Evolution is True. Penguin Group. pp. 85–86. ISBN 978-0-670-
02053.
170.^ Anthony Stevens (1982). Archetype: A Natural History of the Self. Routledge & Kegan
Paul. pp. 87. ISBN 0-7100-0980-1.
171.^ a b Gould 2002, pp. 1235–6
172.^ Pallen, Mark J.; Matzke, NJ (2006-10). "From The Origin of Species to the origin of
bacterial flagella". Nat Rev Micro 4 (10): 784–790. doi:10.1038/nrmicro1493. PMID 16953248.
http://home.planet.nl/~gkorthof/pdf/Pallen_Matzke.pdf. Retrieved 2009-09-18.
173.^ Clements, Abigail; Bursac, D; Gatsos, X; Perry, AJ; Civciristov, S; Celik, N; Likic, VA;
Poggio, S et al. (2009). "The reducible complexity of a mitochondrial molecular machine".
Proceedings of the National Academy of Sciences 106 (37): 15791–15795.
doi:10.1073/pnas.0908264106. PMID 19717453. PMC 2747197.
http://www.pnas.org/content/106/37/15791.long. Retrieved 2009-09-18.
174.^ Piatigorsky J, Kantorow M, Gopal-Srivastava R, Tomarev SI (1994). "Recruitment of
enzymes and stress proteins as lens crystallins". EXS 71: 241–50. PMID 8032155.
175.^ Wistow G (August 1993). "Lens crystallins: gene recruitment and evolutionary dynamism".
Trends Biochem. Sci. 18 (8): 301–6. doi:10.1016/0968-0004(93)90041-K. PMID 8236445.
176.^ Hardin G (April 1960). "The competitive exclusion principle". Science 131: 1292–7.
doi:10.1126/science.131.3409.1292. PMID 14399717. http://www.sciencemag.org/cgi/pmidlookup?
view=long&pmid=14399717.
177.^ Kocher TD (April 2004). "Adaptive evolution and explosive speciation: the cichlid fish
model". Nat. Rev. Genet. 5 (4): 288–98. doi:10.1038/nrg1316. PMID 15131652.
http://hcgs.unh.edu/staff/kocher/pdfs/Kocher2004.pdf.
 178.^ Johnson NA, Porter AH (2001). "Toward a new synthesis: population genetics and
evolutionary developmental biology". Genetica 112–113: 45–58. doi:10.1023/A:1013371201773.
PMID 11838782.
179.^ Baguñà J, Garcia-Fernàndez J (2003). "Evo-Devo: the long and winding road". Int. J. Dev.
Biol. 47 (7–8): 705–13. PMID 14756346. http://www.ijdb.ehu.es/web/paper.php?doi=14756346.
*Love AC. (2003). "Evolutionary Morphology, Innovation, and the Synthesis of Evolutionary and
Developmental Biology". Biology and Philosophy 18 (2): 309–345. doi:10.1023/A:1023940220348.
http://philsci-archive.pitt.edu/archive/00000375/00/LondonPaper.doc.
180.^ Allin EF (1975). "Evolution of the mammalian middle ear". J. Morphol. 147 (4): 403–37.
doi:10.1002/jmor.1051470404. PMID 1202224.
181.^ Harris MP, Hasso SM, Ferguson MW, Fallon JF (2006). "The development of
archosaurian first-generation teeth in a chicken mutant". Curr. Biol. 16 (4): 371–7.
doi:10.1016/j.cub.2005.12.047. PMID 16488870.
182.^ Carroll SB (July 2008). "Evo-devo and an expanding evolutionary synthesis: a genetic
theory of morphological evolution". Cell 134 (1): 25–36. doi:10.1016/j.cell.2008.06.030.
PMID 18614008.
183.^ Wade MJ (2007). "The co-evolutionary genetics of ecological communities". Nat. Rev.
Genet. 8 (3): 185–95. doi:10.1038/nrg2031. PMID 17279094.
184.^ Geffeney S, Brodie ED, Ruben PC, Brodie ED (2002). "Mechanisms of adaptation in a
predator-prey arms race: TTX-resistant sodium channels". Science 297 (5585): 1336–9.
doi:10.1126/science.1074310. PMID 12193784.
*Brodie ED, Ridenhour BJ, Brodie ED (2002). "The evolutionary response of predators to dangerous
prey: hotspots and coldspots in the geographic mosaic of coevolution between garter snakes and
newts". Evolution 56 (10): 2067–82. PMID 12449493.
*Sean B. Carroll (December 21, 2009). "Remarkable Creatures – Clues to Toxins in Deadly
Delicacies of the Animal Kingdom". New York Times.
http://www.nytimes.com/2009/12/22/science/22creature.html?hpw.
185.^ Sachs J (2006). "Cooperation within and among species". J. Evol. Biol. 19 (5): 1415–8;
discussion 1426–36. doi:10.1111/j.1420-9101.2006.01152.x. PMID 16910971.
*Nowak M (2006). "Five rules for the evolution of cooperation". Science 314 (5805): 1560–3.
doi:10.1126/science.1133755. PMID 17158317.
186.^ Paszkowski U (2006). "Mutualism and parasitism: the yin and yang of plant symbioses".
Curr. Opin. Plant Biol. 9 (4): 364–70. doi:10.1016/j.pbi.2006.05.008. PMID 16713732.
187.^ Hause B, Fester T (2005). "Molecular and cell biology of arbuscular mycorrhizal
symbiosis". Planta 221 (2): 184–96. doi:10.1007/s00425-004-1436-x. PMID 15871030.
188.^ Reeve HK, Hölldobler B (2007). "The emergence of a superorganism through intergroup
competition". Proc Natl Acad Sci U S A. 104 (23): 9736–40. doi:10.1073/pnas.0703466104.
PMID 17517608.
189.^ Axelrod R, Hamilton W (2005). "The evolution of cooperation". Science 211 (4489): 1390–
6. doi:10.1126/science.7466396. PMID 7466396.
190.^ Wilson EO, Hölldobler B (2005). "Eusociality: origin and consequences". Proc. Natl. Acad.
Sci. U.S.A. 102 (38): 13367–71. doi:10.1073/pnas.0505858102. PMID 16157878.
191.^ a b Gavrilets S (2003). "Perspective: models of speciation: what have we learned in 40
years?". Evolution 57 (10): 2197–215. doi:10.1554/02-727. PMID 14628909.
192.^ De Queiroz K (December 2007). "Species concepts and species delimitation". Syst. Biol.
56 (6): 879–86. doi:10.1080/10635150701701083. PMID 18027281.
 193.^ Fraser C, Alm EJ, Polz MF, Spratt BG, Hanage WP (February 2009). "The bacterial
species challenge: making sense of genetic and ecological diversity". Science 323 (5915): 741–6.
doi:10.1126/science.1159388. PMID 19197054.
194.^ a b de Queiroz K (May 2005). "Ernst Mayr and the modern concept of species". Proc. Natl.
Acad. Sci. U.S.A. 102 (Suppl 1): 6600–7. doi:10.1073/pnas.0502030102. PMID 15851674.
PMC 1131873. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=15851674.
195.^ Rice, W.R.; Hostert (1993). "Laboratory experiments on speciation: what have we learned
in 40 years". Evolution 47 (6): 1637–1653. doi:10.2307/2410209.
*Jiggins CD, Bridle JR (2004). "Speciation in the apple maggot fly: a blend of vintages?". Trends
Ecol. Evol. (Amst.) 19 (3): 111–4. doi:10.1016/j.tree.2003.12.008. PMID 16701238.
*Boxhorn, J (1995). "Observed Instances of Speciation". TalkOrigins Archive.
http://www.talkorigins.org/faqs/faq-speciation.html. Retrieved 2008-12-26.
*Weinberg JR, Starczak VR, Jorg, D (1992). "Evidence for Rapid Speciation Following a Founder
Event in the Laboratory". Evolution 46 (4): 1214–20. doi:10.2307/2409766.
http://jstor.org/stable/2409766.
196.^ Herrel, A.; Huyghe, K.; Vanhooydonck, B.; Backeljau, T.; Breugelmans, K.; Grbac, I.; Van
Damme, R.; Irschick, D.J. (2008). "Rapid large-scale evolutionary divergence in morphology and
performance associated with exploitation of a different dietary resource". Proceedings of the National
Academy of Sciences 105 (12): 4792–5. doi:10.1073/pnas.0711998105. PMID 18344323.
197.^ Losos, J.B. Warhelt, K.I. Schoener, T.W. (1997). "Adaptive differentiation following
experimental island colonization in Anolis lizards". Nature 387 (6628): 70–3. doi:10.1038/387070a0.
198.^ Hoskin CJ, Higgle M, McDonald KR, Moritz C (2005). "Reinforcement drives rapid
allopatric speciation". Nature 437 (7063): 1353–356. doi:10.1038/nature04004. PMID 16251964.
 199.^ Templeton AR (1 April 1980). "The theory of speciation via the founder principle".
Genetics 94 (4): 1011–38. PMID 6777243. PMC 1214177.
http://www.genetics.org/cgi/reprint/94/4/1011.
200.^ Antonovics J (2006). "Evolution in closely adjacent plant populations X: long-term
persistence of prereproductive isolation at a mine boundary". Heredity 97 (1): 33–7.
doi:10.1038/sj.hdy.6800835. PMID 16639420.
http://www.nature.com/hdy/journal/v97/n1/full/6800835a.html.
201.^ Nosil P, Crespi B, Gries R, Gries G (2007). "Natural selection and divergence in mate
preference during speciation". Genetica 129 (3): 309–27. doi:10.1007/s10709-006-0013-6.
PMID 16900317.
202.^ Savolainen V, Anstett M-C, Lexer C, Hutton I, Clarkson JJ, Norup MV, Powell MP,
Springate D, Salamin N, Baker WJr (2006). "Sympatric speciation in palms on an oceanic island".
Nature 441 (7090): 210–3. doi:10.1038/nature04566. PMID 16467788.
*Barluenga M, Stölting KN, Salzburger W, Muschick M, Meyer A (2006). "Sympatric speciation in
Nicaraguan crater lake cichlid fish". Nature 439 (7077): 719–23. doi:10.1038/nature04325.
PMID 16467837.
203.^ Gavrilets S (2006). "The Maynard Smith model of sympatric speciation". J. Theor. Biol.
239 (2): 172–82. doi:10.1016/j.jtbi.2005.08.041. PMID 16242727.
204.^ Wood TE, Takebayashi N, Barker MS, Mayrose I, Greenspoon PB, Rieseberg LH (August
2009). "The frequency of polyploid speciation in vascular plants". Proc. Natl. Acad. Sci. U.S.A. 106
(33): 13875–9. doi:10.1073/pnas.0811575106. PMID 19667210.
205.^ Hegarty Mf, Hiscock SJ (2008). "Genomic clues to the evolutionary success of polyploid
plants". Current Biology 18 (10): 435–44. doi:10.1016/j.cub.2008.03.043. PMID 18492478.
 206.^ Jakobsson M, Hagenblad J, Tavaré S (2006). "A unique recent origin of the allotetraploid
species Arabidopsis suecica: Evidence from nuclear DNA markers". Mol. Biol. Evol. 23 (6): 1217–31.
doi:10.1093/molbev/msk006. PMID 16549398.
207.^ Säll T, Jakobsson M, Lind-Halldén C, Halldén C (2003). "Chloroplast DNA indicates a
single origin of the allotetraploid Arabidopsis suecica". J. Evol. Biol. 16 (5): 1019–29.
doi:10.1046/j.1420-9101.2003.00554.x. PMID 14635917.
208.^ Bomblies K, Weigel D (2007). "Arabidopsis-a model genus for speciation". Curr Opin
Genet Dev 17 (6): 500–4. doi:10.1016/j.gde.2007.09.006. PMID 18006296.
209.^ Niles Eldredge and Stephen Jay Gould, 1972. "Punctuated equilibria: an alternative to
phyletic gradualism" In T.J.M. Schopf, ed., Models in Paleobiology. San Francisco: Freeman Cooper.
pp. 82–115. Reprinted in N. Eldredge Time frames. Princeton: Princeton Univ. Press. 1985
210.^ Gould SJ (1994). "Tempo and mode in the macroevolutionary reconstruction of
Darwinism". Proc. Natl. Acad. Sci. U.S.A. 91 (15): 6764–71. doi:10.1073/pnas.91.15.6764.
PMID 8041695.
211.^ Benton MJ (1995). "Diversification and extinction in the history of life". Science 268
(5207): 52–8. doi:10.1126/science.7701342. PMID 7701342.
212.^ Raup DM (1986). "Biological extinction in earth history". Science 231: 1528–33.
doi:10.1126/science.11542058. PMID 11542058.
213.^ Avise JC, Hubbell SP, Ayala FJ. (August 2008). "In the light of evolution II: Biodiversity
and extinction". Proc. Natl. Acad. Sci. U.S.A. 105 (Suppl 1): 11453–7. doi:10.1073/pnas.0802504105.
PMID 18695213. PMC 2556414. http://www.pnas.org/content/105/suppl.1/11453.full.
214.^ a b c Raup DM (1994). "The role of extinction in evolution". Proc. Natl. Acad. Sci. U.S.A. 91
(15): 6758–63. doi:10.1073/pnas.91.15.6758. PMID 8041694.
 215.^ Novacek MJ, Cleland EE (2001). "The current biodiversity extinction event: scenarios for
mitigation and recovery". Proc. Natl. Acad. Sci. U.S.A. 98 (10): 5466–70.
doi:10.1073/pnas.091093698. PMID 11344295.
216.^ Pimm S, Raven P, Peterson A, Sekercioglu CH, Ehrlich PR (2006). "Human impacts on
the rates of recent, present, and future bird extinctions". Proc. Natl. Acad. Sci. U.S.A. 103 (29):
10941–6. doi:10.1073/pnas.0604181103. PMID 16829570.
*Barnosky AD, Koch PL, Feranec RS, Wing SL, Shabel AB (2004). "Assessing the causes of late
Pleistocene extinctions on the continents". Science 306 (5693): 70–5. doi:10.1126/science.1101476.
PMID 15459379.
217.^ Lewis OT (2006). "Climate change, species-area curves and the extinction crisis". Philos.
Trans. R. Soc. Lond., B, Biol. Sci. 361 (1465): 163–71. doi:10.1098/rstb.2005.1712. PMID 16553315.
218.^ Jablonski D (May 2001). "Lessons from the past: evolutionary impacts of mass
extinctions". Proc. Natl. Acad. Sci. U.S.A. 98 (10): 5393–8. doi:10.1073/pnas.101092598.
PMID 11344284.
219.^ Isaak, Mark (2005). "Claim CB090: Evolution without abiogenesis". TalkOrigins Archive.
http://www.talkorigins.org/indexcc/CB/CB090.html. Retrieved 2008-12-26.
220.^ Peretó J (2005). "Controversies on the origin of life" (PDF). Int. Microbiol. 8 (1): 23–31.
PMID 15906258. http://www.im.microbios.org/0801/0801023.pdf.
221.^ Luisi PL, Ferri F, Stano P (2006). "Approaches to semi-synthetic minimal cells: a review".
Naturwissenschaften 93 (1): 1–13. doi:10.1007/s00114-005-0056-z. PMID 16292523.
222.^ Trevors JT, Abel DL (2004). "Chance and necessity do not explain the origin of life". Cell
Biol. Int. 28 (11): 729–39. doi:10.1016/j.cellbi.2004.06.006. PMID 15563395. Forterre P,
Benachenhou-Lahfa N, Confalonieri F, Duguet M, Elie C, Labedan B (1992). "The nature of the last
universal ancestor and the root of the tree of life, still open questions". BioSystems 28 (1–3): 15–32.
doi:10.1016/0303-2647(92)90004-I. PMID 1337989.
223.^ Joyce GF (2002). "The antiquity of RNA-based evolution". Nature 418 (6894): 214–21.
doi:10.1038/418214a. PMID 12110897.
224.^ Trevors JT, Psenner R (2001). "From self-assembly of life to present-day bacteria: a
possible role for nanocells". FEMS Microbiol. Rev. 25 (5): 573–82. doi:10.1111/j.1574-
6976.2001.tb00592.x. PMID 11742692.
225.^ Theobald, DL. (2010). "A formal test of the theory of universal common ancestry". Nature
465 (7295): 219–22. doi:10.1038/nature09014. PMID 20463738.
226.^ Bapteste E, Walsh DA (2005). "Does the 'Ring of Life' ring true?". Trends Microbiol. 13 (6):
256–61. doi:10.1016/j.tim.2005.03.012. PMID 15936656.
227.^ Doolittle WF, Bapteste E (February 2007). "Pattern pluralism and the Tree of Life
hypothesis". Proc. Natl. Acad. Sci. U.S.A. 104 (7): 2043–9. doi:10.1073/pnas.0610699104.
PMID 17261804. PMC 1892968. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=17261804.
228.^ Kunin V, Goldovsky L, Darzentas N, Ouzounis CA (2005). "The net of life: reconstructing
the microbial phylogenetic network". Genome Res. 15 (7): 954–9. doi:10.1101/gr.3666505.
PMID 15965028. PMC 1172039. http://www.genome.org/cgi/pmidlookup?
view=long&pmid=15965028.
229.^ Jablonski D (1999). "The future of the fossil record". Science 284 (5423): 2114–16.
doi:10.1126/science.284.5423.2114. PMID 10381868.
230.^ Mason SF (1984). "Origins of biomolecular handedness". Nature 311 (5981): 19–23.
doi:10.1038/311019a0. PMID 6472461.
231.^ Wolf YI, Rogozin IB, Grishin NV, Koonin EV (2002). "Genome trees and the tree of life".
Trends Genet. 18 (9): 472–79. doi:10.1016/S0168-9525(02)02744-0. PMID 12175808.
 232.^ Varki A, Altheide TK (2005). "Comparing the human and chimpanzee genomes: searching
for needles in a haystack". Genome Res. 15 (12): 1746–58. doi:10.1101/gr.3737405.
PMID 16339373.
233.^ Ciccarelli FD, Doerks T, von Mering C, Creevey CJ, Snel B, Bork P (2006). "Toward
automatic reconstruction of a highly resolved tree of life". Science 311 (5765): 1283–87.
doi:10.1126/science.1123061. PMID 16513982.
234.^ a b Cavalier-Smith T (2006). "Cell evolution and Earth history: stasis and revolution".
Philos Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842.
PMID 16754610.
235.^ Schopf J (2006). "Fossil evidence of Archaean life". Philos Trans R Soc Lond B Biol Sci
361 (1470): 869–85. doi:10.1098/rstb.2006.1834. PMID 16754604.
*Altermann W, Kazmierczak J (2003). "Archean microfossils: a reappraisal of early life on Earth".
Res Microbiol 154 (9): 611–17. doi:10.1016/j.resmic.2003.08.006. PMID 14596897.
236.^ Schopf J (1994). "Disparate rates, differing fates: tempo and mode of evolution changed
from the Precambrian to the Phanerozoic". Proc Natl Acad Sci U S a 91 (15): 6735–42.
doi:10.1073/pnas.91.15.6735. PMID 8041691.
237.^ Dyall S, Brown M, Johnson P (2004). "Ancient invasions: from endosymbionts to
organelles". Science 304 (5668): 253–57. doi:10.1126/science.1094884. PMID 15073369.
238.^ Martin W (2005). "The missing link between hydrogenosomes and mitochondria". Trends
Microbiol. 13 (10): 457–59. doi:10.1016/j.tim.2005.08.005. PMID 16109488.
239.^ Lang B, Gray M, Burger G (1999). "Mitochondrial genome evolution and the origin of
eukaryotes". Annu Rev Genet 33: 351–97. doi:10.1146/annurev.genet.33.1.351. PMID 10690412.
*McFadden G (1999). "Endosymbiosis and evolution of the plant cell". Curr Opin Plant Biol 2 (6):
513–19. doi:10.1016/S1369-5266(99)00025-4. PMID 10607659.
 240.^ DeLong E, Pace N (2001). "Environmental diversity of bacteria and archaea". Syst Biol 50
(4): 470–8. doi:10.1080/106351501750435040. PMID 12116647.
241.^ Kaiser D (2001). "Building a multicellular organism". Annu. Rev. Genet. 35: 103–23.
doi:10.1146/annurev.genet.35.102401.090145. PMID 11700279.
242.^ Valentine JW, Jablonski D, Erwin DH (1 March 1999). "Fossils, molecules and embryos:
new perspectives on the Cambrian explosion". Development 126 (5): 851–9. PMID 9927587.
http://dev.biologists.org/cgi/reprint/126/5/851.
243.^ Ohno S (1997). "The reason for as well as the consequence of the Cambrian explosion in
animal evolution". J. Mol. Evol. 44 Suppl 1: S23–7. doi:10.1007/PL00000055. PMID 9071008.
*Valentine J, Jablonski D (2003). "Morphological and developmental macroevolution: a
paleontological perspective". Int. J. Dev. Biol. 47 (7–8): 517–22. PMID 14756327.
http://www.ijdb.ehu.es/web/paper.php?doi=14756327.
244.^ Waters ER (2003). "Molecular adaptation and the origin of land plants". Mol. Phylogenet.
Evol. 29 (3): 456–63. doi:10.1016/j.ympev.2003.07.018. PMID 14615186.
245.^ Mayhew PJ (August 2007). "Why are there so many insect species? Perspectives from
fossils and phylogenies". Biol Rev Camb Philos Soc 82 (3): 425–54. doi:10.1111/j.1469-
185X.2007.00018.x. PMID 17624962.
246.^ Bull JJ, Wichman HA (2001). "Applied evolution". Annu Rev Ecol Syst 32: 183–217.
doi:10.1146/annurev.ecolsys.32.081501.114020.
247.^ Doebley JF, Gaut BS, Smith BD (2006). "The molecular genetics of crop domestication".
Cell 127 (7): 1309–21. doi:10.1016/j.cell.2006.12.006. PMID 17190597.
248.^ Jäckel C, Kast P, Hilvert D (2008). "Protein design by directed evolution". Annu Rev
Biophys 37: 153–73. doi:10.1146/annurev.biophys.37.032807.125832. PMID 18573077.
 249.^ Maher B. (April 2009). "Evolution: Biology's next top model?". Nature 458 (7239): 695–8.
doi:10.1038/458695a. PMID 19360058.
250.^ Borowsky R (January 2008). "Restoring sight in blind cavefish". Curr. Biol. 18 (1): R23–4.
doi:10.1016/j.cub.2007.11.023. PMID 18177707.
251.^ Gross JB, Borowsky R, Tabin CJ (January 2009). "A novel role for Mc1r in the parallel
evolution of depigmentation in independent populations of the cavefish Astyanax mexicanus". PLoS
Genet. 5 (1): e1000326. doi:10.1371/journal.pgen.1000326. PMID 19119422.
252.^ Yergeau DA, Cornell CN, Parker SK, Zhou Y, Detrich HW (July 2005). "bloodthirsty, an
RBCC/TRIM gene required for erythropoiesis in zebrafish". Dev. Biol. 283 (1): 97–112.
doi:10.1016/j.ydbio.2005.04.006. PMID 15890331.
253.^ Fraser AS (1958). "Monte Carlo analyses of genetic models". Nature 181 (4603): 208–9.
doi:10.1038/181208a0. PMID 13504138.
254.^ Rechenberg, Ingo (1973) (in German). Evolutionsstrategie – Optimierung technischer
Systeme nach Prinzipien der biologischen Evolution (PhD thesis) . Fromman-Holzboog.
255.^ Holland, John H. (1975). Adaptation in Natural and Artificial Systems. University of
Michigan Press. ISBN 0262581116.
256.^ Koza, John R. (1992). Genetic Programming. MIT Press. ISBN 0262111705.
257.^ Jamshidi M (2003). "Tools for intelligent control: fuzzy controllers, neural networks and
genetic algorithms". Philosophical transactions. Series A, Mathematical, physical, and engineering
sciences 361 (1809): 1781–808. doi:10.1098/rsta.2003.1225. PMID 12952685.
258.^ Browne, Janet (2003). Charles Darwin: The Power of Place. London: Pimlico. pp. 376–
379. ISBN 0-7126-6837-3.
ISBN 978-0684824710.
*For the scientific and social reception of evolution in the 19th and early 20th centuries, see:
Johnston, Ian C.. "History of Science: Origins of Evolutionary Theory". And Still We Evolve. Liberal
Studies Department, Malaspina University College. http://records.viu.ca/~johnstoi/darwin/sect3.htm.
Retrieved 2007-05-24.
*Bowler, PJ (2003). Evolution: The History of an Idea, Third Edition, Completely Revised and
Expanded. University of California Press. ISBN 978-0520236936.
*Zuckerkandl E (2006). "Intelligent design and biological complexity". Gene 385: 2–18.
doi:10.1016/j.gene.2006.03.025. PMID 17011142.
260.^ Ross, M.R. (2005). "Who Believes What? Clearing up Confusion over Intelligent Design
and Young-Earth Creationism" (PDF). Journal of Geoscience Education 53 (3): 319.
http://www.nagt.org/files/nagt/jge/abstracts/Ross_v53n3p319.pdf. Retrieved 2008-04-28.
261.^ Hameed, Salman (2008-12-12). "Science and Religion: Bracing for Islamic Creationism".
Science 322 (5908): 1637–1638. doi:10.1126/science.1163672. PMID 19074331.
http://helios.hampshire.edu/~sahCS/Hameed-Science-Creationism.pdf. Retrieved 2009.
262.^ Spergel D. N.; Scott, EC; Okamoto, S (2006). "Science communication. Public acceptance
of evolution". Science 313 (5788): 765–66. doi:10.1126/science.1126746. PMID 16902112.
263.^ Spergel, D. N.; Verde, L.; Peiris, H. V.; Komatsu, E.; Nolta, M. R.; Bennett, C. L.; Halpern,
M.; Hinshaw, G. et al. (2003). "First-Year Wilkinson Microwave Anisotropy Probe (WMAP)
Observations: Determination of Cosmological Parameters". The Astrophysical Journal Supplement
Series 148: 175–94. doi:10.1086/377226.
264.^ Wilde SA, Valley JW, Peck WH, Graham CM (2001). "Evidence from detrital zircons for
the existence of continental crust and oceans on the Earth 4.4 Gyr ago". Nature 409 (6817): 175–78.
doi:10.1038/35051550. PMID 11196637.
 265.^ Understanding Creationism after Kitzmiller 2007
266.^ On the history of eugenics and evolution, see Kevles, D (1998). In the Name of Eugenics:
Genetics and the Uses of Human Heredity. Harvard University Press. ISBN 978-0674445574.
267.^ Darwin strongly disagreed with attempts by Herbert Spencer and others to extrapolate
evolutionary ideas to all possible subjects; see Midgley, M (2004). The Myths we Live By. Routledge.
p. 62. ISBN 978-0415340779.
268.^ Allhoff F (2003). "Evolutionary ethics from Darwin to Moore". History and philosophy of the
life sciences 25 (1): 51–79. doi:10.1080/03919710312331272945. PMID 15293515.
269.^ Gowaty, Patricia Adair (1997). Feminism and evolutionary biology: boundaries,
intersections, and frontiers. London: Chapman & Hall. ISBN 0-412-07361-7.

[edit] Further reading

Introductory reading
• Carroll, S. (2005). Endless Forms Most Beautiful. New York: W.W. Norton. ISBN 0-393-
06016-0.
• Charlesworth, C.B. and Charlesworth, D. (2003). Evolution. Oxfordshire: Oxford University
Press. ISBN 0-192-80251-8.
• Dawkins, R. (2006). The Selfish Gene: 30th Anniversary Edition. Oxford University Press.
ISBN 0199291152.
• Gould, S.J. (1989). Wonderful Life: The Burgess Shale and the Nature of History . New York:
W.W. Norton. ISBN 0-393-30700-X.
• Jones, S. (2001). Almost Like a Whale: The Origin of Species Updated. (American title:
Darwin's Ghost). New York: Ballantine Books. ISBN 0-345-42277-5.
 • Mader, Sylvia S. (2007). Biology. Murray P. Pendarvis (9th ed.). McGraw Hill.
ISBN 9780073258393.
• Maynard Smith, J. (1993). The Theory of Evolution: Canto Edition. Cambridge University
Press. ISBN 0-521-45128-0.
• Pallen, M.J. (2009). The Rough Guide to Evolution. Rough Guides. ISBN 978-1-85828-946-
5.
• Smith, C.B. and Sullivan, C. (2007). The Top 10 Myths about Evolution. Prometheus Books.
ISBN 978-1-59102-479-8.
History of evolutionary thought
• Darwin, Charles (1859). On the Origin of Species (1st ed.). London: John Murray.
ISBN 0801413192. http://darwin-online.org.uk/content/frameset?
itemID=F373&viewtype=text&pageseq=1.
• Larson, E.J. (2004). Evolution: The Remarkable History of a Scientific Theory. New York:
Modern Library. ISBN 0-679-64288-9.
• Zimmer, C. (2001). Evolution: The Triumph of an Idea. London: HarperCollins. ISBN 0-060-
19906-7.
Advanced reading
• Barton, N.H., Briggs, D.E.G., Eisen, J.A., Goldstein, D.B. and Patel, N.H. (2007). Evolution.
Cold Spring Harbor Laboratory Press. ISBN 0-879-69684-2.
• Coyne, J.A. and Orr, H.A. (2004). Speciation. Sunderland: Sinauer Associates. ISBN 0-878-
93089-2.
 • Futuyma, D.J. (2005). Evolution. Sunderland: Sinauer Associates. ISBN 0-878-93187-2.
• Gould, S.J. (2002). The Structure of Evolutionary Theory. Cambridge: Belknap Press
(Harvard University Press). ISBN 0-674-00613-5.
• Maynard Smith, J. and Szathmáry, E. (1997). The Major Transitions in Evolution.
Oxfordshire: Oxford University Press. ISBN 0-198-50294-X.
• Mayr, E. (2001). What Evolution Is. New York: Basic Books. ISBN 0-465-04426-3.
• Olson, Wendy; Hall, Brian Keith (2003). Keywords and concepts in evolutionary
developmental biology. Cambridge: Harvard University Press. ISBN 0-674-02240-8.

[edit] External links

General information
• Everything you wanted to know about evolution by New Scientist
• Howstuffworks.com — How Evolution Works
• National Academies Evolution Resources
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and
Theories
• Understanding Evolution from University of California, Berkeley
• Evolution of Evolution - 150 Years of Darwin's "On the Origin of Species"
History of evolutionary thought
• The Complete Work of Charles Darwin Online
• Understanding Evolution: History, Theory, Evidence, and Implications
 On-line lectures
• What Genomes Can Tell Us About the Past – lecture by Sydney Brenner
• The Origin of Vertebrates – lecture by Marc Kirschner
• The Making of the Fittest – lecture by Sean B. Carroll

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Modularity · Phenotypic plasticity
concepts
 Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution"

Categories: Articles with separate introductions | Evolution | Biology theories | Evolutionary biology

• This page was last modified on 21 August 2010 at 03:16.

 W000

Évolution (biologie)
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir Évolution.

 Arbre phylogénétique hypothétique de tous les organismes vivants. L'arbre est basé sur des
séquences de l'ARNr 16S. À l'origine proposé par Carl Woese, il montre l'histoire évolutive supposée des
trois domaines du vivant (bactéries, archées et eucaryotes).
En biologie, l'évolution désigne la transformation des espèces vivantes au cours des générations.
Cette transformation peut aboutir à la formation de nouvelles espèces, et donc à une diversification des
formes de vie. Cette diversification depuis les premières formes est à l'origine de la biodiversité actuelle.
L’histoire de l'évolution de la vie peut ainsi être décrite sous forme d'un « arbre évolutif », ou arbre
phylogénétique.
L'idée d'évolution peut déjà se trouver chez certains philosophes de l'antiquité (grecs, romains) ou
bien musulmans, mais ce n'est qu'à partir du XIXe siècle que de véritables théories proposant une explication
du phénomène de l'évolution des espèces ont été développées. Si la théorie du transformisme de Lamarck a
ouvert la voie, la révolution évolutionniste est arrivée avec Charles Darwin et son ouvrage De l'origine des
espèces (1859) dans lequel deux grandes idées, appuyés par des faits, émergent : l'unité et la diversité du
vivant s'explique par l'évolution, et le moteur de l'évolution adaptative est la sélection naturelle. En profonde
contradiction avec les idées philosophiques et religieuses dominantes de l'époque, De l'origine des espèces
obtient un important écho et convainc rapidement la majorité des biologistes de la réalité de l'évolution[1].
Avec la théorie synthétique de l'évolution qui regroupe notamment les idées de Darwin avec celles
de Mendel, l'évolution fait l'objet d'un large consensus scientifique sur ses fondements et ses mécanismes
depuis le milieu du XXe siècle. Dans la biologie contemporaine, si l'idée que les espèces évoluent ne fait plus
aucun doute, les détails des mécanismes qui permettent d'expliquer cette évolution font toujours l'objet de
recherches et sont parfois au cœur de controverses scientifiques et médiatiques, comme celle ayant opposé
Stephen Jay Gould et Richard Dawkins sur l'intérêt d'introduire la notion d'équilibres ponctués.
L'évolution est causée, d'une part par la présence de variations parmi les traits héréditaires d'une
population d'individus (mutations), et d'autre part par divers mécanismes qui vont modifier la fréquence de
certains traits héréditaires. Parmi ces mécanismes, la sélection naturelle désigne la différence de
propagation entre les traits héréditaires causée par leur effet sur la survie et la reproduction des individus : si
un certain trait héréditaire favorise les chances de survie et la reproduction, il s'ensuit mécaniquement que la
fréquence de ce trait augmente d'une génération à l'autre. Dans une population de taille finie, un trait peut
également être propagé ou éliminé par le fait de fluctuations aléatoires (dérive génétique). À l'échelle des
temps géologiques, l'évolution conduit à des changements morphologiques, anatomiques, physiologiques et
comportementaux des espèces.
Si l'on veut retranscrire ces concepts en systématique, il faut considérer la théorie cladistique, selon
laquelle les grades évolutifs (qui induisent une vision de l'évolution aujourd'hui obsolète [réf. nécessaire]) ne
sont plus pris en compte, en faveur des clades[2].
Du fait, entre autres, de ses implications sur l'origine de l'humanité, l'évolution a été, et reste
toujours, mal comprise et/ou, parfois, mal admise hors de la communauté scientifique. Dans les sociétés
occidentales, la théorie de l'évolution se heurte à une vive opposition de la part de certains milieux religieux
fondamentalistes, notamment pour son incompatibilité avec une interprétation littérale de la Bible. Ses
détracteurs se basent sur des analyses pseudo-scientifiques ou religieuses pour contredire l'idée même
d'évolution des espèces ou la théorie de la sélection naturelle.
Sommaire
[masquer]
• 1 Histoire de la théorie de l'évolution
• 2 Les arguments en faveur de l'évolution.
• 2.1 Stratégie de raisonnement
• 2.2 Indices morphologiques
• 2.3 Indices moléculaires
• 2.4 Indices comportementaux
• 2.5 Un exemple d'évolution à échelle de temps
humaine : Podarcis sicula
• 3 Les critiques de la théorie synthétique de l'évolution
• 4 Méthodes d'étude de l'évolution
• 4.1 La paléobiologie
• 4.2 L'analyse comparative des caractères
• 4.3 La génétique des populations
• 4.4 La modélisation
• 4.5 L'expérimentation
• 5 Mécanismes de l'évolution
• 5.1 L'évolution des populations
• 5.1.1 L'apparition de nouveaux caractères
• 5.1.2 Variabilité des individus au sein des
Histoire de la théorie de l'évolution [modifier]
Article détaillé : Histoire de la pensée évolutionniste.
Bien que les hommes cherchaient l'origine de la diversité du vivant dès la période antique, l'idée
d'évolution, c'est-à-dire de modification des espèces au cours du temps, a mis du temps à s'imposer. Si
l'idée d'une évolution de la vie est déjà présente chez quelques philosophes grecs[3], l'un de ceux qui a le
plus marqué le monde occidental[4], Aristote, avait une conception fixiste du vivant, et cette vision est restée
prédominante dans la pensée occidentale jusqu'au XVIIIe siècle. L'influence des religions monothéistes
abrahamiques est prédominante dans la diffusion de ces idées fixistes, sous une nouvelle forme : le
créationnisme. En effet les récits bibliques, en particulier ceux de la Genèse, prônent que toutes les espèces
vivantes ont été créées telles quelles et de manière synchrone par Dieu et qu'elles sont parfaites donc
immuables ; de plus, l'homme occuperait une place à part dans le vivant puisqu'il serait à l'image de Dieu et
serait moralement supérieur à toutes les autres espèces[3].
Durant le Moyen Âge, les avancées scientifiques en Europe occidentale deviennent limitées par la
dominance du fondamentalisme chrétien, qui prône une interprétation littérale des textes sacrés[5]. Bien que
les autorités religieuses condamnent fermement toute idée scientifique remettant en cause les écrits
bibliques, l'idée d'évolution se retrouve chez certains savants comme Jérôme Cardan[6] et Giulio Cesare
Vanini[7]. Parallèlement, l'idée d'évolution apparait dans le monde musulman, et l'on trouve dès le IXe siècle
non seulement l'idée que les espèces évoluent au cours du temps, mais aussi une première théorie
cherchant à expliquer cette évolution[8]. Au XIIIe siècle, le philosophe Nasir ad-Din at-Tusi propose,
plusieurs siècles avant Charles Darwin, la sélection des meilleurs et l'adaptation des espèces à leur
environnement[9]. Cependant, ces idées n'ont eu qu'une faible popularité, y compris au sein du monde
musulman[réf. souhaitée].
 Au début du XVIIIe siècle, les idées fixistes alors prédominantes sont ébranlées par le
développement de la paléontologie et la découverte de fossiles de squelettes ne ressemblant à aucun
squelette actuel[10]. Pour concilier ces découvertes avec les textes bibliques, Georges Cuvier expose sa
théorie catastrophiste selon laquelle il y aurait eu une succession de créations divines entrecoupées
d'extinctions brutales au cours des temps géologiques[11]. Ils admet ainsi que les espèces terrestres n'ont
pas toujours été celles observées aujourd'hui, sans pour autant accepter l'évolution des espèces, et que les
6 000 ans estimés jusque là pour l'âge de la Terre sont trop courts pour y intégrer ces extinctions
successives[12].
Si l'idée d'évolution est réapparue au milieu du XVIIIe siècle avec Maupertuis et Buffon, la première
théorie véritablement scientifique considérant une évolution des espèces vivantes, le lamarckisme, est
fondée par le naturaliste Jean-Baptiste Lamarck.
Lamarck considère que les espèces peuvent se transformer selon deux principes :
1. La diversification, ou spécialisation, des êtres vivants en de multiples espèces, sous l'effet
des circonstances variées auxquelles ils sont confrontés dans des milieux variés et auxquelles ils
sont contraints de s'adapter en modifiant leur comportement ou leurs organes pour répondre à leurs
besoins, généralement désigné par « l'usage et le non-usage » ;
2. la complexification croissante de l'organisation des êtres vivants sous l'effet de la dynamique
interne propre à leur métabolisme.
Sa théorie a souvent été abusivement réduite à celle de la transmission des caractères acquis, qui
veut que les modifications acquises au cours de la vie d'un organisme soit héréditaires, mais en fait Lamarck
ne propose pas de théorie de l'hérédité de l'acquis (contrairement à ce que fera Darwin en 1868), il se
contente de reprendre les idées admises sur ce point depuis Aristote. Il n'en reste pas moins que Lamarck
est le premier à proposer une théorie expliquant les êtres vivant à partir de laquelle il tente de comprendre
l'évolution des espèces. La publication, en 1809, dans Philosophie zoologique, de sa théorie transformiste
entraine de virulents débats au sein de l'Académie des sciences car elle entre en contradiction avec les
idées en vigueur à l'époque et notamment le fixisme. Malgré de nombreuses critiques de la part des milieux
religieux et scientifique, et notamment de la part de Cuvier qui devient le principal opposant des
transformistes, les idées transformistes reçoivent une adhésion croissante à partir de 1825 et permettent de
rendre le débat naturaliste plus réceptif aux théories évolutionnistes[13] même si Lamarck ne verra jamais
ses travaux reconnus par la communauté scientifique[14].
Cependant, la première théorie satisfaisante permettant d'expliquer l'adaptation des espèces est
publiée en 1859 par Charles Darwin dans son livre De l'origine des espèces. Cette théorie, fondement de la
théorie actuelle de l'évolution, considère que, étant donné que tous les individus d'une espèce diffèrent au
moins légèrement, et que seule une partie de ces individus réussit à se reproduire, seuls les descendants
des individus les mieux adaptés à leur environnement participeront à la génération suivante. Ainsi, comme
les individus sélectionnés transmettent leurs caractères à leur descendance, les espèces évoluent et
s'adaptent en permanence à leur environnement. Il baptise du nom de sélection naturelle cette sélection des
individus les mieux adaptés[15]. Darwin n'avait cependant aucune idée du mécanisme permettant la
transmission des caractères[réf. souhaitée].
La découverte des lois de Mendel et de la génétique au début du XXe siècle bouleverse la
compréhension des mécanismes de l'évolution et donne naissance à la Théorie synthétique de l'évolution,
fondée entre autres par Theodosius Dobzhansky et Ernst Mayr. Cette théorie est une combinaison de la
théorie de la sélection naturelle proposée par Darwin et de la génétique mendellienne. Elle est à l'origine de
nouvelles méthodes dans l'étude de l'évolution, comme la génétique des populations ou la
modélisation[réf. nécessaire].
 À partir de ce moment, la biologie de l'évolution intègre toutes les autres disciplines de la biologie, et
cherche aussi bien à retracer l'histoire évolutive du vivant qu'à théoriser et prouver les mécanismes en jeu
lors de l'évolution des espèces. La fin du XXe siècle est ainsi très prolifique sur le plan théorique. Plus
récemment, l'étude de l'évolution profite du développement de l'informatique et de la biologie moléculaire, et
notamment du séquençage qui permet le développement de la phylogénie par un apport très important de
données[réf. nécessaire].
La biologie de l'évolution est aujourd'hui une composante majeure de la biologie qui nourrit aussi
bien qu'elle se nourrit de toutes les autres disciplines[réf. nécessaire].

Les arguments en faveur de l'évolution. [modifier]

Stratégie de raisonnement [modifier]

Si on arrive à établir un lien de parenté entre deux espèces différentes, alors cela veut dire qu'une
espèce ancestrale s'est transformée en, au moins, une de ces deux espèces. Il y a donc bien eu évolution.
Un lien de parenté entre espèces fossiles ou actuels peut être mis en évidence par le partage d'au
moins un caractère homologue, c'est-à-dire provenant d'un ancêtre. Ces indices de parenté sont décelables
au niveau de la morphologie, au niveau moléculaire et parfois même, pour des espèces très proches, au
niveau du comportement.
Utilisation des fossiles
 Archaefructus liaoningensis le plus ancien reste de fleur connu.
Il est en pratique impossible d'affirmer qu'une espèce fossile est l'ancêtre d'une espèce actuelle, car
il ne sera jamais garanti que l'espèce actuelle ne s'est pas différenciée à partir d'une autre espèce proche,
mais qui n'aurait pas été découverte. En effet, la conservation de restes d'espèces éteintes est un
événement relativement improbable surtout pour les périodes les plus anciennes. On peut seulement estimer
les liens de parenté, avec les autres espèces déjà connnues, actuelles ou fossiles. Par exemple le fossile de
fleur le plus ancien a été daté de 140 millions d'années. Cet organe est donc apparu sur Terre, il y a au
moins 140 millions d'années. Mais d'autres espèces proches, avec des fleurs, existaient aussi certainement
à cette époque. Personne n'est capable d'affirmer laquelle de ces espèces est l'ancêtre des plantes à fleur
actuelles. On ne cherchera que les relations de parenté, les relations d'ancêtre à descandant ne pouvant
jamais être reconstituées.
 L'âge d'une espèce fossile, en revanche, indique l'âge minimum d'apparition des caractères qu'elle
possède. Il est alors possible de reconstruire l'histoire de l'évolution, en plaçant sur une échelle des temps
l'apparition des différents caractères. Les fossiles nous indiquent que l'ordre d'apparition des innovations
évolutives est tout à fait en accord avec l'idée d'une évolution, qui dans un schéma général, part de
structures simples vers des structures plus complexes. C'est aussi en accord avec une origine aquatique des
êtres vivants.

Indices morphologiques [modifier]

Squelette de baleine. En c figure le vestige de bassin[16].

Les « mains » des tétrapodes ont un même plan d'organisation, ce qui traduit une homologie.
• les baleines, animaux adaptés à la vie aquatique gardent une trace de leurs ancêtres
quadrupèdes par la présence d'os vestigials correspondant au bassin (ceinture pelvienne) ;
• Il y a des vestiges de pattes chez certains serpents (boas)[17];
• En observant l'aile d'un oiseau ou d'une chauve-souris, on retrouve aisément la structure
osseuse du membre antérieur de tout tétrapode ;
• les défenses à croissance continue des éléphants sont en fait homologues des incisives des
autres mammifères, dont l'homme ;
• les appendices masticateurs des arthropodes sont à l'origine des appendices locomoteurs
réduits (il en va de même apparemment pour les Onychophores) ;
• les membres des tétrapodes proviennent des nageoires de poissons ;
• dans le monde végétal, la présence d'une double membrane autour des plastes et la
présence d'un ADN circulaire à l'intérieur de ceux-ci trahissent une origine endosymbiotique
procaryote.

Indices moléculaires [modifier]

• Le support de l'information héréditaire est toujours l'ADN pour l'ensemble du vivant ;
 • Le code génétique, code de correspondance entre l'ADN et les protéines est quasiment le
même chez tous les êtres vivants ;
• Le séquençage de l'ADN. fait apparaître de nombreuses régions étroitement proches donc
apparentés (gènes homologues: paralogues ou orthologue) qui codent des protéines aux fonctions
ou structures différentes mais assez proches (Exemple : les gènes qui codent les hémoglobines,
myoglobines...).

Indices comportementaux [modifier]

Chez certaines espèces de Lacertidés américains du genre Cnemidophorus, ou lézards à queue en
fouet, il n'existe plus que des femelles. Ces espèces pratiquent donc une reproduction asexuée. Cependant
des simulacres d'accouplements persistent : pour se reproduire une femelle monte sur une autre dans un
comportement similaire à celui des espèces sexuées. Ce comportement d'origine hormonale est à mettre en
relation avec une origine récente de ces espèces parthénogénétiques[18].
Un exemple d'évolution à échelle de temps humaine : Podarcis sicula [modifier]

Podarcis sicula. Des lézards des ruines déposés sur l'île de Prod Mrcaru en 1971 ont évolué en 36
ans de sorte à disposer d'un nouvel organe de digestion absent chez l'espèce d'origine : les valves cæcales.
Introduit en 1971 par l'équipe du professeur Eviatar Nevo sur l'île dalmate de Prod Mrcaru en mer
Adriatique, le lézard Podarcis sicula connu en France sous le nom de « lézard des ruines », y a été
abandonné à lui-même durant près de quatre décennies, l'accès à l'île ayant été interdit par les autorités
yougoslaves, puis par les conflits liés à l'éclatement de ce pays. En 2004, une équipe scientifique dirigée par
Duncan Irschick et Anthony Herrel put revenir sur l'île et découvrit que Podarcis sicula avait évolué en 36
ans, soit environ trente générations, de façon très significative. Le lézard a grandi, sa mâchoire est devenue
plus puissante, et surtout il a changé de régime alimentaire : d'insectivore il est devenu herbivore, et des
valves cæcales sont apparues au niveau des intestins, ce qui lui permet de digérer les herbes... Cette
découverte confirme, s'il en était encore besoin, que l'évolution n'est pas une théorie parmi d'autres, mais un
phénomène biologique concrètement observable, et pas seulement chez les virus, les bactéries ou les
espèces domestiquées[19].

Les critiques de la théorie synthétique de l'évolution [modifier]

Article détaillé : Historique des critiques des théories de l'évolution.
Ceux-ci peuvent être rapidement classés en diverses origines :
• Critiques idéologiques.
• Critiques scientifiques (notamment par rapport aux fossiles).
• Critiques religieuses (créationnisme et intelligent design).

Méthodes d'étude de l'évolution [modifier]

La paléobiologie [modifier]
Articles détaillés : Paléontologie et Paléogénétique.
La paléobiologie, étude de la vie des temps passés, permet de reconstituer l'histoire des êtres
vivants. Cette histoire donne aussi des indices sur les mécanismes évolutifs en jeu dans l'évolution des
espèces. La paléontologie s'occupe plus particulièrement des restes fossiles des êtres vivants. La
paléogénétique, science récente, s'intéresse au matériel génétique ayant survécu jusqu'à aujourd'hui[20].
Ces deux approches sont limitées par la dégradation du matériel biologique au cours du temps. Ainsi, les
informations issues des restes sont d'autant plus rares que l'être vivant concerné est ancien. De plus,
certaines conditions sont plus propices que d'autres à la conservation du matériel biologique. Ainsi, les
environnements anoxiques ou très froids entravent la dégradation des restes. Les restes vivants sont donc
lacunaires et sont bien souvent insuffisants pour retracer l'histoire évolutive du vivant.

L'analyse comparative des caractères [modifier]

Articles détaillés : Génétique évolutive du développement et Phylogénie.
Tous les êtres vivants actuels étant issus d'un même ancêtre commun, ils partagent des
caractéristiques héritées de cet ancêtre. L'analyse des ressemblances entre êtres vivants donne de
nombreuses informations sur leurs relations de parenté, et permet de retracer l'histoire évolutive des
espèces. La phylogénie est la discipline scientifique qui cherche à retracer les relations entre êtres vivants
actuels et fossiles à partir de l'analyse comparative des caractères morphologiques, physiologiques ou
moléculaires. L'analyse comparative permet de retracer l'histoire évolutive des différents caractères dans les
lignées du vivant. L'évolution des caractères ne suit pas nécessairement celle des espèces, certains
caractères (dits convergents) peuvent être apparus plusieurs fois de manière indépendante dans différentes
lignées.
L'évolution des caractères et des lignées peut être associée à des évènements géologiques ou
biologiques marquant l'histoire de la Terre, ce qui permet de proposer des hypothèses sur les mécanismes à
l'origine de l'évolution des espèces.
La nature des caractères pouvant être analysés est extrêmement diverse, et il peut s'agir aussi bien
de caractères morphologiques (taille, forme ou volume de différentes structures), anatomiques (structure,
organisation des organes), tissulaires, cellulaires ou moléculaires (séquences protéiques ou nucléiques).
Ces différents caractères apportent des informations diverses et souvent complémentaires. Actuellement, les
caractères moléculaires (en particulier les séquences d'ADN) sont privilégiées, du fait de leur universalité, de
leur fiabilité et du faible coût des technologies associées. Ils ne peuvent cependant pas être utilisés lors de
l'étude de fossiles pour lesquels seuls les caractères morphologiques sont en général informatifs.

La génétique des populations [modifier]

Article détaillé : Génétique des populations.

La modélisation [modifier]
La modélisation en biologie de l'évolution se base sur les mécanismes de l'évolution mis en évidence
pour mettre en place des modèles théoriques. Ces modèles peuvent produire des résultats qui dépendent
des hypothèses de départ de ce modèle, ces résultats pouvant être comparés à des données réellement
observées. On peut ainsi tester la capacité du modèle à refléter la réalité, et, dans une certaine mesure, la
validité de la théorie sous-jacente à ce modèle.
Les modèles dépendent souvent de paramètres, lesquels ne peuvent pas toujours être déterminés a
priori. La modélisation permet de comparer les résultats du modèles et ceux de la réalité pour de
nombreuses valeurs différentes de ces paramètres, et ainsi déterminer quelles sont les combinaisons de
paramètres qui permettent au modèle décrire au mieux la réalité. Ces paramètres correspondent souvent à
des paramètres biologiques, et on peut ainsi estimer à partir du modèle certains paramètres biologiques
difficile à mesurer. La justesse de l'estimation de ces paramètres dépend cependant de la validité du modèle,
laquelle est parfois difficile à tester.
 La modélisation permet enfin de prédire certaines évolutions à venir, en utilisant les données
actuelles comme données de départ du modèle.

L'expérimentation [modifier]
Article détaillé : Évolution expérimentale.
L'évolution expérimentale est la branche de la biologie qui étudie l'évolution par de réelles
expériences, à l'inverse de l'étude comparative des caractères, qui ne fait que regarder l'état actuel des êtres
vivants. Les expériences consistent généralement en l'isolement d'une ou plusieurs espèces dans un milieu
biologique contrôlé. On laisse alors ces espèces évoluer pendant un certain temps, en appliquant
éventuellement des changements contrôlés de conditions environnementales. On compare enfin certaines
caractéristiques des espèces avant et après la période d'évolution.
L'évolution expérimentale permet non seulement d'observer l'évolution en cours, mais aussi de
vérifier certaines prédictions énoncées dans le cadre de la théorie de l'évolution, et tester l'importance
relative de différents mécanismes évolutifs.
L'évolution expérimentale ne peut étudier que des caractères évoluant rapidement, et se limite donc
à des organismes se reproduisant rapidement, notamment des virus ou des unicellulaires, mais aussi
certains organismes à génération plus longue comme la drosophile ou certains rongeurs.
Un exemple : l'expérience de Luria et Delbrück.

Mécanismes de l'évolution [modifier]

Article détaillé : Théorie synthétique de l'évolution.
L'évolution des populations [modifier]

L'évolution des caractères dans les populations: diversité, sélection et transmission

Parce que les individus d'une population possèdent des caractères héritables différents, et que seule
une partie de ces individus accède à la reproduction, les caractères les plus adaptés à l'environnement sont
préférentiellement conservés par la sélection naturelle. De plus, le hasard de la reproduction sexuée rend
partiellement aléatoire les caractères qui seront transmis, par effet de dérive génétique. Ainsi, la proportion
des différents caractères d'une population varie d'une génération à l'autre, conduisant à l'évolution des
populations.

L'apparition de nouveaux caractères [modifier]

Cela se produit par mutation et recombinaison génétique, ou remaniement chromosomique. Mais
cela ne se déroule que dans un individu, pas dans l'espèce entière. Il faut, pour que ce nouveau caractère se
répande, l'effet de la sélection naturelle et/ou de la dérive génétique.

Variabilité des individus au sein des populations [modifier]

Articles détaillés : Diversité génétique, Mutation et Reproduction sexuée.
Tous les individus d'une espèces sont uniques et diffèrent les uns des autres. Ces différences sont
observables à toutes les échelles, du point de vue morphologique jusqu'à l'échelle moléculaire. Cette
diversité des populations a deux origines principales: les individus sont dissemblables parce qu'ils ne
possèdent pas la même information génétique et parce qu'ils ont subi des influences environnementales
différentes.
La diversité génétique se manifeste par des variations locales de la séquence d'ADN, formant
différents variants de la même séquence appelés allèles. Cette variabilité a plusieurs origines. Des allèles
peuvent être formés spontanément par mutation de la séquence d'ADN. Par ailleurs, la reproduction sexuée
contribue à la diversité génétique des populations de deux manières: d'une part, la recombinaison génétique
permet de diversifier les combinaisons d'allèles réunies sur un même chromosome. D'autre part, une partie
du génome de chaque parent est sélectionnée aléatoirement pour former un nouvel individu, dont le génome
est par conséquent unique.
 La diversité issue de l'environnement s'acquiert tout au long de l'histoire de l'individu, depuis la
formation des gamètes jusqu'à sa mort. L'environnement étant unique à chaque endroit et à chaque moment,
il exerce des effets unique sur chaque individu, et ce à toutes les échelles, de la morphologie jusqu'à la
biologie moléculaire. Ainsi, deux individus possédant la même information génétique (c'est par exemple le
cas pour les vrais jumeaux) sont tout de même différents. Ils peuvent notamment avoir une organisation et
une expression différente de l'information génétique.

L'hérédité [modifier]
Articles détaillés : Hérédité, ADN, Reproduction, Réplication de l'ADN et Épigénétique.
Les êtres vivants sont capables de se reproduire, transmettant ainsi une partie de leurs caractères à
leurs descendants. On distingue la reproduction asexuée, ne faisant intervenir qu'un individu, de la
reproduction sexuée pendant laquelle deux individus mettent en commun une partie de leur matériel
génétique, formant ainsi un individu génétiquement unique.
mitochondries, n'est transmis que par une partie des individus de l'espèce (les femelles chez les
mammifères).

La transmission des caractères acquis, une hypothèse non totalement rejetée [modifier]
Article détaillé : Transmission des caractères acquis.
La théorie synthétique de l'évolution, paradigme dominant actuel, se fonde sur un déterminisme
génétique intégral et écarte donc toute transmission héréditaire de caractères acquis au cours de la vie de
l'individu. Néanmoins de plus en plus de travaux scientifiques remettent en cause ce modèle et rétablissent
pour partie l'idée d'une transmission héréditaire de caractères acquis que défendait le lamarckisme[21].
Tout d'abord, certains caractères dits épigénétiques concernent la structure et l'organisation des
génomes sont transmis par les parents en même temps que les molécules d'acide nucléique elles-mêmes.
De plus, la mère fournit l'environnement cytoplasmique de la cellule-oeuf du descendant, et transmet ainsi un
certain nombre de caractéristiques cellulaires à l'enfant. Des modifications épigénétiques conservées dans la
lignée germinale sont désormais décrites chez plusieurs espèces. Chez les plantes il existe une corrélation
entre le niveau d'expression d'un gène et sa méthylation. Pareillement, chez les mammifères nous
témoignons de la méthylation d'une séquence transposable qui est insérée à proximité d'un gène particulier.
Le degré de méthylation d'un transposon pouvant enfin moduler l'expression du gène dans lequel il s'est
inséré[22]. L'étude de l'épigénétique, longtemps délaissée, connait un grand essor depuis la fin du
séquençage de nombreux génomes, dont celui de l'homme.
Ainsi, Une étude de 2009 du MIT affirme mettre en évidence une hérédité de certains caractères
acquis chez des rongeurs[23]. Par ailleurs, l’obésité serait non pas uniquement un effet direct touchant les
individus atteints eux-mêmes mais également un effet transgénérationnel. Des données chez l'homme et
chez l'animal semblent montrer que les effets d'une sous-alimentation subies par des individus pourraient en
effet être transmises aux descendants.

La dérive génétique [modifier]

Article détaillé : dérive génétique.
 Simulation informatique de l'évolution de la fréquence d'un allèle neutre au cours du temps dans une
population de 10 (en haut) ou 100 individus (en bas). Chaque courbe représente une simulation différente,
les différences illustrant l'effet du hasard (dérive génétique). Les fluctuations de la fréquence de l'allèle sont
plus importantes lorsque la population est de taille réduite, et la fixation (fréquence égale à 1) ou la perte
(fréquence égale à 0) d'un allèle est alors plus rapide.
Lors de la reproduction sexuée, la transmission des caractères (notamment des allèles) comporte
une grande part de hasard due à la recombinaison homologue, et au brassage génétique. Ainsi, on observe
une variation aléatoire des fréquences alléliques d'une génération à l'autre, appelée dérive génétique. La
dérive génétique génère donc une composante aléatoire dans l'évolution des populations. Ainsi, deux
populations d'une même espèce n'échangeant pas de matériel génétique vont diverger jusqu'à former, si le
temps d'isolement génétique est suffisant, deux espèces différentes. La dérive génétique est donc un des
moteurs de la spéciation.
L'effet de la dérive génétique est particulièrement visible lorsqu'un faible nombre d'individus est à
l'origine d'une population beaucoup plus nombreuse. C'est le cas lorsque se forme un goulot d'étranglement
c'est-à-dire qu'une population est décimée et se reconstitue, ou lorsque quelques individus d'une population
migrent pour aller coloniser un nouvel espace et former une nouvelle population (effet fondateur). Lorsqu'un
tel évènement se produit, un allèle même faiblement représenté dans la population de départ peut se
retrouver en forte proportion dans la population nouvellement formée sous le simple effet d'un hasard dans le
tirage des individus à l'origine de la nouvelle population. Inversement, un allèle fortement représenté peut ne
pas être tiré, et disparaît de la nouvelle population. Par ailleurs, la formation d'une nouvelle population à
partir d'un faible nombre d'individu a pour effet d'augmenter la consanguinité dans la population et augmente
le pourcentage d'homozygotie, ce qui fragilise la population.
 La sélection naturelle [modifier]
Article détaillé : Sélection naturelle.
Dans la très grande majorité des espèces, le nombre de cellules-œuf produits est bien plus grand
que le nombre d'individus arrivant à l'âge de la maturité sexuelle et parmi ceux-ci, une partie seulement
accède à la reproduction. Ainsi, seule une partie des individus formés se reproduit à la génération suivante. Il
existe donc une sélection des individus perpétuant l'espèce, seuls les individus n'étant pas éliminé par les
conditions environnementales pouvant se reproduire. Cette sélection a été baptisée sélection naturelle.
Comme il existe une variabilité au sein des espèces, les individus possédant des caractères
différents, et qu'une partie de ces caractères sont héréditaires, les caractères permettant à l'individu de
survivre et de mieux se reproduire seront préférentiellement transmis à la descendance, par rapport aux
autres caractères. Ainsi la proportion des caractères au sein des espèces évolue au cours du temps.
La sélection naturelle peut prendre des formes très variées. La sélection utilitaire est une élimination
des individus les moins capables de survivre et les moins féconds, alors que la sélection sexuelle conserve
préférentiellement les individus les plus aptes à rencontrer un partenaire sexuel. Bien que ces sélections
soient complémentaires, on observe souvent des conflits, chaque forme de sélection pouvant favoriser
l'évolution d'un caractère dans un sens différent.
Il est parfois observé une sélection d'individus qui favorisent la survie ou la reproduction d'individus
qui leurs sont ou non apparentés, comme c'est le cas chez les insectes eusociaux ou lorsqu'un individu se
sacrifie pour permettre la survie de son groupe ou de sa descendance. En sociobiologie, ces comportements
altruistes s'expliquent notamment par les théories controversées de la sélection de parentèle, de la sélection
de groupe et de l'altruisme réciproque. La sélection de parentèle prédit qu'il peut être plus avantageux pour
un individu de favoriser beaucoup la reproduction d'un individu apparenté (donc avec lequel il partage des
caractères) que de se reproduire un peu ou pas du tout, la sélection de groupe repose sur le même principe
mais du point de vue du groupe et pourrait expliquer certains actes chez l'homme comme les guerres ou la
xénophobie, l'altruisme réciproque se penche sur les cas d'altruisme entre individus non-apparentés et induit
une contribution réciproque dont l'aide donnée en retour peut être différé dans le temps.
Enfin, la sélection artificielle n'est qu'une forme de sélection naturelle exercée par l'homme.

Conséquences évolutives [modifier]

Adaptation des espèces [modifier]

Articles détaillés : Adaptation et Neutralisme.
En conséquence de la sélection naturelle, les espèces conservent préférentiellement les caractères
les plus adaptés à leur environnement, et sont donc de mieux en mieux adaptées à leur environnement. Les
pressions de sélection en jeux dans cette adaptation sont nombreuses et concernent tous les aspects de
l'environnement, des contraintes physiques jusqu'aux espèces biologiques interagissantes.
L'adaptation de plusieurs espèces différentes sous l'effet des mêmes pressions environnementales
peut conduire à l'apparition répétée et indépendante du même caractère adaptatif chez ces espèces, par un
phénomène de convergence évolutive. Par exemple, chez les mammifères les cétacés et les siréniens ont
tout deux développé des nageoires, de manière indépendante. L'évolution de ces nageoires montre une
adaptation convergente à la vie aquatique.
 Cependant, l'effet de la sélection naturelle est réduit par celui de la dérive génétique. Ainsi, un
caractère avantageux pourra ne pas être sélectionné à cause de l'inertie donnée par la dérive.

Apparition et disparition des espèces [modifier]

Articles détaillés : Spéciation et Extinction des espèces.
L'évolution d'une population sous l'effet du hasard et des contraintes environnementales peut aboutir
à la disparition de la population et éventuellement de l'espèce à laquelle elle appartient. Inversement, deux
populations peuvent s'individualiser au sein d'une même espèce jusqu'à former deux espèces distinctes par
un processus nommé spéciation.

Controverses sur les mécanismes de l'évolution [modifier]

Cet article ne cite pas suffisamment ses sources (décembre 2009).
Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
bas de page. (Modifier l'article)
L'évolution et ses mécanismes sont encore largement étudiés aujourd'hui, et de nombreux points sur
les mécanismes de l'évolution ne sont pas éclaircis. Certaines questions déjà soulevées par Charles Darwin
n'ont d'ailleurs toujours pas de réponse certaine.
Une des grandes questions de la théorie de l'évolution est l'origine des rangs taxinomiques
supérieurs à celui de l'espèce. En outre, la manière dont sont apparus les 33 embranchements animaux,
issus de l'explosion cambrienne, pose encore problème. Ainsi, la théorie gradualiste estime que les
changements interviennent de manière progressive au cours de l'évolution, alors que la théorie des
équilibres ponctués défend qu'il existe des sauts évolutifs majeurs.
La transmission des caractères acquis, complètement délaissée depuis la découverte des lois de
l'hérédité, est réactualisée par la découverte des phénomènes épigénétiques. Dès lors, l'importance de cette
transmission de caractères non hérités des parents dans l'évolution des espèces doit se poser. Cependant,
notre connaissance des mécanismes épigénétiques est encore trop faible pour pouvoir répondre à cette
question. En outre, peu d'études sur le rôle de l'épigénétique dans l'évolution ont été réalisées à l'heure
actuelle.
Il a été longtemps admis que l'évolution s'accompagnait d'un accroissement de la complexité des
êtres vivants. Cependant, cette idée, largement influencée par l'anthropocentrisme, est fortement débattue
aujourd'hui. La complexité n'ayant pas de définition précise à l'heure actuelle, il est difficile de vérifier une
éventuelle augmentation de complexité. Par ailleurs, lorsque cette idée est admise, les origines de cette
augmentation de complexité sont, elles aussi, source de controverse. En fait, tout cela à déjà été clairement
expliqué par Lamarck.

Histoire évolutive du vivant [modifier]

L'origine de la Vie se situerait vers - 3.8 milliards d'années. Il s'agissait probablement d'organismes
procaryotes unicellulaires. On suppose un ancêtre unique à tous les êtres vivants (LUCA). À partir de cet
ancêtre se sont diversifiées les différentes formes de Vie.
Articles détaillés : Histoire évolutive du vivant et Arbre phylogénétique du vivant.
Évolution et sociétés humaines [modifier]

Évolution et agriculture [modifier]

Article détaillé : Sélection artificielle.
L'homme a su très vite utiliser la variabilité des populations à son profit : l'évolution dirigée par
l'homme, ou sélection artificielle, à cause de la sélection par les éleveurs et les cultivateurs, se produit
depuis des millénaires. Il avait été remarqué depuis longtemps que les animaux d'élevage héritaient, dans
une certaine mesure, de caractéristiques de leurs parents et nul n'aurait songé à utiliser ses bêtes les plus
malingres pour la reproduction. D'ailleurs, Darwin utilise de nombreuses observations issues de la sélection
des plantes et des animaux en agriculture pour étayer ses idées. Ainsi, l'homme peut créer une sélection dite
artificielle sur son environnement, volontairement pour des raisons économiques, ou involontairement via la
pression de chasse, cueillette ou pêche)[24].

Évolution et informatique [modifier]

Article détaillé : Algorithme évolutionniste.
L'efficacité du processus de sélection naturelle a inspiré la création d'algorithmes évolutionnistes
(comme les algorithmes génétiques) en informatique. Ces algorithmes heuristiques modélisent plusieurs
caractéristiques de l'évolution biologique (en particulier les mutations et les recombinaisons) pour trouver
une solution satisfaisante à un problème trop complexe pour être abordé par d'autres méthodes.
Eugénisme [modifier]
Articles détaillés : Darwinisme social et Eugénisme.
Article détaillé : Évolutionnisme (anthropologie).
La pensée évolutionniste s'est notamment propagée au sein de l'anthropologie évolutionniste au
XIXe siècle. Pour les anthropologues de cette époque, l'espèce humaine ne fait qu'une, et donc, chaque
société suit la même évolution, qui commence à l'état de « primitif » pour arriver jusqu'au modèle de la
civilisation occidentale. Cette théorie a été très fortement remise en question. En effet, elle ne correspond
pas à la réalité historique observée (les civilisations suivent des « chemins » divergents, ne poursuivent pas
les mêmes « objectifs », et la civilisation occidentale, qui devrait pourtant constituer le stade ultime de
l'évolution, continue pourtant à vivre de profondes mutations.) et est douteuse d'un point de vue éthique
(considérant notre société occidentale comme l'aboutissement ultime de la civilisation). À l'inverse de ce qui
était pratiqué jusqu'au milieu du XXe siècle, les approches modernes de l'anthropologie évolutionniste
privilégient une méthodologie précise (confrontant des sources multiples, s'inspirant des outils d'analyse
quantitative des sciences sociales, tentant de se départir de l'ethnocentrisme) et s'appuie sur des théories
plus élaborées que l'évolutionnisme simpliste des débuts. Théories inspirées non seulement par la biologie
de l'évolution moderne mais aussi par la modélisation mathématique et informatique et parfois enrichies par
les connaissances contemporaines en psychologie.

La psychologie évolutionniste [modifier]

Article détaillé : Psychologie évolutionniste.
évolutionniste. Même si Darwin avait déjà émis l'idée que la sélection naturelle a pu façonner aussi bien des
caractères anatomiques que psychologiques, cette discipline s'est véritablement formalisée au début des
années 1990 dans le cadre conceptuel des sciences cognitives. Depuis, la psychologie évolutionniste est au
centre d'une intense controverse scientifique qui tient à de multiples raisons : difficulté méthodologique à
établir une histoire évolutive des comportements qui ne sont pas des objets matériels, résistance
intellectuelle à envisager l'esprit humain comme en partie déterminé par l'évolution, utilisation simpliste et
abusive des théories évolutionnistes, médiatisation et déformation auprès du grand public des
problématiques scientifiques... Dans le milieu scientifique toutefois, la psychologie évolutionniste fait
désormais partie des paradigmes scientifiques valides.

Evolutionnisme et religions [modifier]

La théorie évolutionniste est-elle compatible avec la croyance en Dieu ? En fait, Ernst Mayr dit à ce
sujet : « Il me semble évident que Darwin a perdu la foi un an sinon deux, avant de formuler sa théorie de la
sélection naturelle (sur laquelle il a sans doute travaillé plus de dix ans). Par conséquent, il n'est pas fondé
d'avancer que la biologie et l'adhésion à la théorie de la sélection naturelle risquent de vous éloigner de
Dieu.[25]»
Le biologiste Richard Dawkins, dans son ouvrage Pour en finir avec Dieu (2008), pense que la
sélection naturelle est « supérieure » à l'« hypothèse de Dieu » qu'il qualifie d'« improbabilité statistique », et
défend l'athéisme.
Le biologiste Kenneth R. Miller (en) estime que la pensée évolutionniste n'est pas forcément
incompatible avec la foi en un Dieu[26]. Pour lui les écrits de la Bible sont des métaphores.
 L'évolution est encore aujourd'hui rejetée par certains milieux religieux, tenants du créationnisme,
surtout protestants.
La position de l'Église Catholique sur ce sujet est plus nuancée, tout en maintenant l'innerance de la
Bible[27], une microévolution au sein des espèces semble aujourd'hui « plus qu'une hypothèse ». Elle
déclare que Dieu est le seul créateur, qu'Il a créé le monde par amour, mais que l'esprit ne peut pas être le
fruit d'une évolution de la matière[28].

Aspects politiques et judiciaires [modifier]

Les polémiques ont débordé, depuis les années 1990, le simple cadre du débat public, notamment
aux États-Unis.
Dans certains États, les tenants du créationnisme ont essayé de rendre obligatoire son
enseignement dans les écoles publiques, en tant que « théorie scientifique concurrente » de celle de
l'évolution. Cependant ces mesures ont été déclarés anticonstitutionnelles vis-à-vis du premier amendement
sur la liberté d'expression, du fait du caractère religieux de cette théorie. Devant ces tentatives, des
scientifiques ont ironiquement demandé à ce que soit aussi enseigné le pastafarisme (qui a été inventé à
cette occasion).
Un nouveau concept est apparu dans la mouvance créationniste, baptisé dessein intelligent
(« Intelligent Design »), qui affirme que « certaines caractéristiques de l'Univers et du monde vivant sont
mieux expliquées par une cause intelligente, plutôt que par des processus aléatoires tels que la sélection
naturelle[29] ». Cette thèse est présentée comme une théorie appuyée par des travaux scientifiques, et ne
nie pas l'existence de tout phénomène évolutif. La justice américaine, s'appuyant sur les travaux
scientifiques, a cependant jugé (voir Kitzmiller v. Dover Area School) que cette thèse était de nature
religieuse et non scientifique, et que les promoteurs de l’ Intelligent Design n'explicitaient pas cette « cause
intelligente » afin de contourner le problème juridique et d'échapper au qualificatif religieux. D'autres groupes
utilisent les arguments de l’Intelligent Design, avec diverses attributions pour la « cause intelligente », par
exemple des extraterrestres.

Notes et références [modifier]

1. ↑ Darwin n'utilise pas le mot évolution dans son œuvre, puisque ce terme n'est introduit que
dans les années 1870. Cf. Gould (1997) : 33-37, Laurent (2001) : 17.
2. ↑ W. Hennig, Phylogenetic Systematics, Illinois University Press, 1966, traduit par D. Dwight
Davis & R. Zangerl.
3. ↑ a et b Barbara Cassin & al., L'animal dans l'Antiquité [lire en ligne [archive]], Centre National
de Recherche Scientifique, éd. Vrin, 1997, 618 p., (ISBN 2711613232).
4. ↑ Jean-Philippe Omotunde, Platon et Aristote : les deux piliers de la pensée
occidentale [archive], Institut Africamaat, (page consultée le 4 juillet 2008).
5. ↑ Jean Chaline, Quoi de neuf depuis Darwin ?, éd. Ellipses, 2006 (ISBN 978-2-7298-3100-
4).
6. ↑ Jérôme Cardan définit une théorie de l'évolution dans son ouvrage De Subtilitate Rerum
en 1551.
7. ↑ Giulio Cesare Vanini est brûlé vif en 1619 pour avoir notamment déclaré que l'homme et le
singe pouvaient être des parents.
8. ↑ Mehmet Bayrakdar (The Islamic Quarterly Third Quarter, 1983). "Al-Jahiz And the Rise of
Biological Evolutionism [archive]", Londres.
 9. ↑ Farid Alakbarli, « A 13th-Century Darwin? », dans Azerbaijan International, vol. 9.2, 2001,
p. 48-49
10.↑ Laurent Dubois, [pdf] Histoire de la paléontologie, Darwin et Théorie de
l'Evolution [archive], Géopolis.fr, (page consultée le 4 juillet 2008).
11.↑ Lapierre, S., « Éléments de théorie de l'évolution [archive] » sur
http://www.colvir.net/prof/serge.lapierre/index.html [archive]. Consulté le 19 octobre 2008
12.↑ Serge Lapierre, Éléments de théorie de l'évolution [archive], Collège de Bois de Boulogne
- Département de philosophie, (page consultée le 25 octobre 2008).
13.↑ Hélène Blais, « Lamarck, genèse et enjeux du transformisme, 1770-1830 », La Revue
pour l’histoire du CNRS [lire en ligne [archive]], n°7 - Novembre 2002, mis en ligne le 6 mars 2006.
Consulté le 7 juillet 2008.
14.↑ Benoît Virole, Le voyage intérieur de Charles Darwin: essai sur la genèse psychologique
d'une œuvre scientifique [lire en ligne [archive]], p. 124-126, éd. des archives contemporaines, 2000,
144 p. (ISBN 9057090171)
15.↑ Darwin, C., De l'Origine des espèces, Flammarion, 1859, 1997
16.↑ L. Bejder, B.K. Hall, « Limbs in whales and limblessness in other vertebrates: mechanisms
of evolutionary and developmental transformation and loss », dans Evol. Dev., vol. 6, no 4, Nov.-Dev.
2002, p. 445-58.
17.↑ (en)Vestigials Organs, A Snake—With Legs ! American Museum of Natural
History [archive] Site du muséum d'histoire naturelle américain.
18.↑ Serotonergic modulation of male-like pseudocopulatory behavior in the parthenogenetic
whiptail lizard, Cnemidophorus uniparens Brian George Dias et David Crews Hormones and
Behavior Volume 50, Issue 3, Septembre 2006, 401-409.
19.↑ Voir site : [pdf](en)PNAS, vol. 105, n°12, pages 4792-4795 [archive] (25 mars 2008).
 20.↑ Eva-Maria Geigt, « L'émergence de la paléogénétique », dans Biofutur, no 164, Février
1997, p. 28-34 (ISSN 0294-3506)
21.↑ La théorie de l'évolution en évolution ? Hominidés.com [archive]
22.↑ Claudine Junien, « Obésité et diabète de type 2 : L'hypothèse de la transmission
épigénétique », dans Cahiers de nutrition et de diététique, vol. 37, no 4, 2002, p. 261-272
(ISSN 0007-9960) [ texte intégral [archive] (page consultée le 11/03/2009) ]
23.↑ http://www.technologyreview.com/biomedicine/22061/ [archive]
24.↑ Chris T. Darimonta et al. ; Human predators outpace other agents of trait change in the
wild ; Ed : Gretchen C. Daily, Stanford University, Stanford, CA, PNAS, approuvé le 21 nov 2008
(reçu pour relecture le 15 septembre 2008) (Lire l'article [archive])
25.↑ Biologie Campbell, De Boeck Université ISBN 2-8041-2084-8 p.417
26.↑ Kenneth Miller À la recherche du Dieu de Darwin, édition Sciences et quête de sens, 2000
27.↑
http://www.vatican.va/roman_curia/congregations/cfaith/pcb_documents/rc_con_cfaith_doc_194801
16_fonti-pentateuco_fr.html [archive]
28.↑ http://www.hominides.com/html/theories/jean_paul_evolution.php [archive]
29.↑ « The theory of intelligent design [...] holds that certain features of the universe and of
living things are best explained by an intelligent cause rather than an undirected process such as
natural selection » — Intelligent Design Network, Inc. [archive]
 Voir aussi [modifier]
Ouvrages sur le sujet [modifier]
Lorsqu'il y a deux dates, la première est
celle de la première parution, dans la langue
d'origine.
• Brondex, F. (1999) Évolution :
synthèse des faits et théories, Dunod.
• Buican, D. (1989) La Révolution de
l'évolution, PUF.
• Buican, D. (1997) L'Évolution et les
théories évolutionnistes, Masson.
• Buican, D. (2008) L'odyssée de
l'évolution, Ellipses
• Chapouthier, G. (2001) L'homme,
ce singe en mosaïque, Odile Jacob
• Combes, C. (2006) Darwin,
dessine-moi les hommes
• Darwin, C. (1997, éd. or. 1859)
L'Origine des espèces, Flammarion.
• Charles Darwin. Origines - Lettres
choisies 1828-1859 (2009), introduction et
• Jacob, F. (1981) Le Jeu des
possibles, Fayard.
• Kropotkine, P (2001, éd. or.
1902) L'entraide : un facteur de
l'évolution, Ecosociété.
• Laurent, G. (2001). La
Naissance du transformisme. Lamarck
entre Linné et Darwin, Vuibert (Paris) et
ADAPT (Paris) : 151 p. (ISBN 2-7117-
5348-4)
• Dominique Lecourt, (1992, 3e
éd. « Quadrige » 1998), L’Amérique
entre la Bible et Darwin, suivi de
Intelligent design : science, morale et
politique, PUF.
• Lecourt, D. dir. (1999, 4e réed.
« Quadrige » 2006), Dictionnaire
d’histoire et philosophie des sciences,
PUF.
• Le Guyader, H., dir. (1998)
édition française dirigée par Dominique
Lecourt, préface S. J. Gould, éditions
Bayard, (ISBN 978-2-227-47843-5).
• David, P. & Samadi, S. (2000) La
Théorie de l'évolution, Flammarion.
• Dawkins, R. (1982) The Extended
Phenotype, Oxford University Press.
• Dawkins, R. (1986, 1989)
L'Horloger aveugle, Éditions Robert Laffont.
• Dawkins, R. (1996) Climbing Mount
Improbable, Norton (anglophone).
• Dawkins, R. (1976, 1996) Le Gène
égoïste, Odile Jacob.
• Dennett, D. (2000) Darwin est-il
dangereux ?, Odile Jacob.
• Devillers, C. & Tintant, H. (1996)
Questions sur la théorie de l'évolution,
PUF.
• Dorléans, P. (2003) Il était une fois
l'évolution, Ellipses.
• Futuyma, D.J (1997) Evolutionary
Biology, Sinauer Associates.
• Gould, S. J. (1982) Le Pouce du
L'évolution, Belin/Pour la Science.
• Le Guyader, H. et Lecointre, G.,
Classification phylogénétique du vivant,
Belin (Paris) : 560 p. (ISBN 2-7011-
4273-3)
• Lehman, J.-P. (1973) Les
preuves paléontologiques de l'évolution,
PUF.
• Lodé, T. (2006) La guerre des
sexes chez les animaux Odile Jacob,
Paris. (ISBN 2-7381-1901-8)
• Marchand, D. (2002) Les
merveilles de l'évolution, P.U. Dijon.
• Mayr, E. (1989) The Growth of
Biological Thought: Diversity, Evolution
and Inheritance, Ed. Cambridge, Harvard
University Press — traduit en français
sous le titre de Histoire de la biologie.
Diversité, évolution et hérédité, Fayard
(1989) : 894 p. (ISBN 2213018944).
• Mayr, E. (2004) What makes
biology unique? Considerations on the
Autonomy of a Scientific Discipline, Ed.
New York, Cambridge University Press —
panda, Grasset.
• Gould, S. J. (1991) La Vie est belle,
Le Seuil.
• Gould, S. J. (1997) L'Éventail du
vivant, Le Seuil.
• Gould, S. J. (1997). Darwin et les
grandes énigmes de la vie. Réflexions sur
l'histoire naturelle. 1, S 43, Seuil (Paris),
collection Point Science : 311 p.
• Gould, S. J. (2000) Et Dieu dit :
Que Darwin soit ! : Science et religion,
enfin la paix ?, préface de D. Lecourt, Le
Seuil.
• Gould, S. J. (2002) The structure of
evolutionary theory, Harvard University
Press (anglophone).
• Grasset P. P. (1973) L'Évolution du
vivant, matériaux pour une théorie
transformiste, Albin Michel.
• Grimoult, C. (2000) Histoire de
l'évolutionnisme contemporain en France
(1945-1995), Genève, Droz.
• Grimoult, C.(2001) L'évolution
biologique en France. Une révolution
traduit en français sous le titre de Ernst
Après Darwin. La biologie, une science
pas comme les autres, Dunod (2006) :
237 p. (ISBN 2-10-049560-7).
• Pichot, A. (1993) Histoire de la
notion de vie, éd. Gallimard, coll. TEL.
• Ridley (1998) Evolution
Biologique, Ed. De Boeck (traduction
française).
• Tort, P. (1996) Dictionnaire du
darwinisme et de l’évolution, Ed. Paris,
PUF, 3 vol., 5 000 p. Ouvrage couronné
par l’Académie des Sciences.
• Wright, R. (1995) L’Animal
Moral, Michalon.
• Zimmer, K. (2001) Evolution :
the triumph of an idea, Harper Collins
(anglophone)
scientifique, politique et culturelle, Genève,
Droz.
Articles connexes [modifier]
• Adaptation • Ève • Mémétique
• Arbre mitochondriale • Panspermie
phylogénétique du • Évolution • Photo-guide
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History of evolutionary thought

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This article is about the history of evolutionary thought in biology. For the history of evolutionary
thought in the social sciences, see Sociocultural evolution. For the history of religious discussions, see
History of the creation-evolution controversy.
 The Tree of Life as depicted by Ernst Haeckel in The Evolution of Man (1879) illustrates the 19th-
century view that evolution was a progressive process leading towards man.
Evolutionary thought, the conception that species change over time, has roots in antiquity, in the
ideas of the ancient Greeks, Romans, and Chinese as well as in medieval Islamic science. However, until the
18th century, Western biological thinking was dominated by essentialism, the belief that every species has
essential characteristics that are unalterable. This began to change during the Enlightenment when
evolutionary cosmology and the mechanical philosophy spread from the physical sciences to natural history.
Naturalists began to focus on the variability of species; the emergence of paleontology with the concept of
extinction further undermined the static view of nature. In the early 19th century, Jean-Baptiste Lamarck
proposed his theory of the transmutation of species, the first fully formed scientific theory of evolution.
In 1858, Charles Darwin and Alfred Russel Wallace published a new evolutionary theory that was
explained in detail in Darwin's On the Origin of Species (1859). Unlike Lamarck, Darwin proposed common
descent and a branching tree of life. The theory was based on the idea of natural selection, and it
synthesized a broad range of evidence from animal husbandry, biogeography, geology, morphology, and
embryology.
The debate over Darwin's work led to the rapid acceptance of the general concept of evolution, but
the specific mechanism he proposed, natural selection, was not widely accepted until it was revived by
developments in biology that occurred during 1920s through the 1940s. Before that time most biologists
argued that other factors were responsible for evolution. Alternatives to natural selection suggested during
the eclipse of Darwinism included inheritance of acquired characteristics (neo-Lamarckism), an innate drive
for change (orthogenesis), and sudden large mutations (saltationism). The synthesis of natural selection with
Mendelian genetics during the 1920s and 1930s founded the new discipline of population genetics.
Throughout the 1930s and 1940s, population genetics became integrated with other biological fields,
resulting in a widely applicable theory of evolution that encompassed much of biology—the modern
evolutionary synthesis.
Following the establishment of evolutionary biology, studies of mutation and variation in natural
populations, combined with biogeography and systematics, led to sophisticated mathematical and causal
models of evolution. Paleontology and comparative anatomy allowed more detailed reconstructions of the
history of life. After the rise of molecular genetics in the 1950s, the field of molecular evolution developed,
based on protein sequences and immunological tests, and later incorporating RNA and DNA studies. The
gene-centered view of evolution rose to prominence in the 1960s, followed by the neutral theory of molecular
evolution, sparking debates over adaptationism, the units of selection, and the relative importance of genetic
drift versus natural selection. In the late 20th century, DNA sequencing led to molecular phylogenetics and
the reorganization of the tree of life into the three-domain system. In addition, the newly recognized factors of
symbiogenesis and horizontal gene transfer introduced yet more complexity into evolutionary history.
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Contents
[hide]
• 1 Antiquity
• 1.1 Greeks
• 1.2 Chinese
• 1.3 Romans
• 1.4 Augustine of Hippo
• 2 Middle Ages
• 2.1 Islamic philosophy and the struggle for
existence
• 2.2 Christian philosophy and the great chain of
being
• 2.3 Thomas Aquinas on creation and natural
processes
• 3 Renaissance and Enlightenment
• 4 Early 19th century
• 4.1 Paleontology and geology
• 4.2 Transmutation of species
• 4.3 Anticipations of natural selection
• 4.4 Natural selection
• 5 1859–1930s: Darwin and his legacy
[edit] Antiquity
[edit] Greeks
Several ancient Greek philosophers discussed ideas that involved change in living organisms over
time. Anaximander (c.610–546 BC) proposed that the first animals lived in water and animals that live on land
were generated from them.[1] Empedocles (c. 490–430 BC) wrote of a non-supernatural origin for living
things,[2] suggesting that adaptation did not require an organizer or final cause. Aristotle summarized his
idea: "Wherever then all the parts came about just what they would have been if they had come to be for an
end, such things survived, being organized spontaneously in a fitting way; whereas those which grew
otherwise perished and continue to perish ..." although Aristotle himself rejected this view.[3]
 Plato (left) and Aristotle (right), a detail of The School of Athens
Plato (c. 428–348 BC) was, in the words of biologist and historian Ernst Mayr, "the great antihero of
evolutionism",[4] as he established the philosophy of essentialism, which he called the Theory of Forms. This
theory holds that objects observed in the real world are only reflections of a limited number of essences
(eide). Variation merely results from an imperfect reflection of these constant essences. In his Timaeus, Plato
set forth the idea that the Demiurge had created the cosmos and everything in it because, being good, and
hence, "... free from jealousy, He desired that all things should be as like Himself as they could be". The
creator created all conceivable forms of life, since "... without them the universe will be incomplete, for it will
not contain every kind of animal which it ought to contain, if it is to be perfect". This "plenitude principle"—the
idea that all potential forms of life are essential to a perfect creation—greatly influenced Christian thought.[5]
Aristotle (384–322 BC), one of the most influential of the Greek philosophers, is the earliest natural
historian whose work has been preserved in any real detail. His writings on biology resulted from his
research into natural history on and around the isle of Lesbos, and have survived in the form of four books,
usually known by their Latin names, De anima (on the essence of life), Historia animalium (inquiries about
animals), De generatione animalium (reproduction), and De partibus animalium (anatomy). Aristotle's works
contain some remarkably astute observations and interpretations—along with sundry myths and mistakes—
reflecting the uneven state of knowledge during his time.[6] However, for Charles Singer, "Nothing is more
remarkable than [Aristotle's] efforts to [exhibit] the relationships of living things as a scala naturæ."[6] This
scala naturæ, described in Historia animalium, classified organisms in relation to a hierarchical "Ladder of
Life" or "Chain of Being", placing them according to their complexity of structure and function, with organisms
that showed greater vitality and ability to move described as "higher organisms".[5]

[edit] Chinese
Ideas on evolution were expressed by ancient Chinese thinkers such as Zhuangzi (Chuang Tzu), a
Taoist philosopher who lived around the 4th century BC. According to Joseph Needham, Taoism explicitly
denies the fixity of biological species and Taoist philosophers speculated that species had developed
differing attributes in response to differing environments.[7] Humans, nature and the heavens are seen as
existing in a state of "constant transformation" known as the Tao, in contrast with the more static view of
nature typical of Western thought.[8]
[edit] Romans
Titus Lucretius Carus (d. 50 BC), the Roman philosopher and atomist, wrote the poem On the Nature
of Things (De rerum natura), which provides the best surviving explanation of the ideas of the Greek
Epicurean philosophers. It describes the development of the cosmos, the Earth, living things, and human
society through purely naturalistic mechanisms, without any reference to supernatural involvement. On the
Nature of things would influence the cosmological and evolutionary speculations of philosophers and
scientists during and after the Renaissance.[9][10]

[edit] Augustine of Hippo

In line with earlier Greek thought, the 4th century bishop and theologian, St. Augustine of Hippo,
wrote that the creation story in Genesis should not be read too literally. In his book De Genesi ad litteram
("On The Literal Interpretation of Genesis"), he stated that in some cases new creatures may have come
about through the "decomposition" of earlier forms of life.[11] For Augustine, "plant, fowl and animal life are
not perfect ... but created in a state of potentiality", unlike what he considered the theologically perfect forms
of angels, the firmament and the human soul.[12] Augustine's idea 'that forms of life had been transformed
"slowly over time"' prompted Father Giuseppe Tanzella-Nitti, Professor of Theology at the Pontifical Santa
Croce University in Rome, to claim that Augustine had suggested a form of evolution.[13][14]

[edit] Middle Ages

[edit] Islamic philosophy and the struggle for existence
See also: Early Islamic philosophy and Science in medieval Islam
 Although Greek and Roman evolutionary ideas died out in Europe after the fall of the Roman Empire,
they were not lost to Islamic philosophers and scientists. In the Islamic Golden Age of the 8th to the 13th
centuries, philosophers explored ideas about natural history. These ideas included transmutation from non-
living to living: "from mineral to plant, from plant to animal, and from animal to man".[15]
The first Muslim biologist and philosopher to speculate in detail about natural history was the Afro-
Arab writer al-Jahiz in the 9th century. In the Book of Animals, he considered the effects of the environment
on an animal's chances for survival, and described the struggle for existence.[16] Al-Jahiz also wrote
descriptions of food chains.[17] Al-Jahiz speculated on the influence of the environment on animals and
considered the effects of the environment on the likelihood of an animal to survive. For example, Al-Jahiz's
wrote in his Book of Animals:
All animals, in short, can not exist without food, neither can the hunting animal escape being
hunted in his turn. Every weak animal devours those weaker than itself. Strong animals cannot
escape being devoured by other animals stronger than they.[16]
[edit] Christian philosophy and the great chain of being
 Drawing of the great chain of being from Rhetorica Christiana (1579) by Diego Valades
During the Early Middle Ages, Greek classical learning was all but lost to the West. However, contact
with the Islamic world, where Greek manuscripts were preserved and expanded, soon led to a massive spate
of Latin translations in the 12th century. Europeans were re-introduced to the works of Plato and Aristotle, as
well as to Islamic thought. Christian thinkers of the scholastic school, in particular Abelard and Thomas
Aquinas, combined Aristotelian classification with Plato's ideas of the goodness of God, and of all potential
life forms being present in a perfect creation, to organize all inanimate, animate, and spiritual beings into a
huge interconnected system: the scala naturæ, or great chain of being.[5][18]
Within this system, everything that existed could be placed in order, from "lowest" to "highest", with
Hell at the bottom and God at the top—below God, an angelic hierarchy marked by the orbits of the planets,
mankind in an intermediate position, and worms the lowest of the animals. As the universe was ultimately
perfect, the great chain was also perfect. There were no empty links in the chain, and no link was
represented by more than one species. Therefore no species could ever move from one position to another.
Thus, in this Christianized version of Plato's perfect universe, species could never change, but remained
forever fixed, in accordance with the text of Genesis. For humans to forget their position was seen as sinful,
whether they behaved like lower animals or aspired to a higher station than was given them by their Creator.
[5]
Creatures on adjacent steps were expected to closely resemble each other, an idea expressed in the
saying: natura non facit saltum ("nature does not make leaps").[5] This basic concept of the great chain of
being greatly influenced the thinking of Western civilization for centuries (and still has an influence today). It
formed a part of the argument from design presented by natural theology. As a classification system, it
became the major organizing principle and foundation of the emerging science of biology in the 17th and
18th centuries.[5]
[edit] Thomas Aquinas on creation and natural processes
While the development of the great chain of being and the argument from design by Christian
theologians contributed to the view that the natural world fit into an unchanging designed hierarchy, some
theologians were more open to the possibility that the world might have developed through natural
processes. Thomas Aquinas went even farther than Augustine of Hippo in arguing that scriptural texts like
Genesis should not be interpreted in a literal way that conflicted with or constrained what natural
philosophers learned about the workings of the natural world. He felt that the autonomy of nature was a sign
of God's goodness and that there was no conflict between the concept of a divinely created universe, and the
idea that the universe may have evolved over time through natural mechanisms.[19] However, Aquinas
disputed the views of those like the ancient Greek philosopher Empodocles who held that such natural
processes showed that the universe could have developed without an underlying purpose. Rather holding
that:
Hence, it is clear that nature is nothing but a certain kind of art, i.e., the divine art, impressed
upon things, by which these things are moved to a determinate end. It is as if the shipbuilder
were able to give to timbers that by which they would move themselves to take the form of a
ship.[20]

[edit] Renaissance and Enlightenment

Main article: Evolutionary ideas of the Renaissance and Enlightenment
 Pierre Belon compared the skeletons of birds and humans in his Book of Birds (1555).
In the first half of the 17th century, René Descartes's mechanical philosophy encouraged the use of
the metaphor of the universe as a machine, a concept that would come to characterise the scientific
revolution.[21] Between 1650 and 1800, some evolutionist theories supported the view that the universe,
including life on Earth, had developed mechanically, entirely without divine guidance. In contrast, most
contemporary theories of evolution, such of those of Gottfried Leibniz and J. G. Herder, held that evolution
was a fundamentally spiritual process.[22] In 1751, Pierre Louis Maupertuis veered toward more materialist
ground. He wrote of natural modifications occurring during reproduction and accumulating over the course of
many generations, producing races and even new species, a description that anticipated in general terms the
concept of natural selection.[23]
 Later in the 18th century, the French philosopher G. L. L. Buffon, one of the leading 18th century
naturalists, suggested that what most people referred to as species were really just well-marked varieties,
modified from an original form by environmental factors. For example, he believed that lions, tigers, leopards
and house cats might all have a common ancestor. He further speculated that the 200 or so species of
mammals then known might have descended from as few as 38 original animal forms. Buffon's evolutionary
ideas were limited; he believed each of the original forms had arisen through spontaneous generation and
that each was shaped by "internal moulds" that limited the amount of change. Buffon's works, Natural History
and The Epochs of Nature, containing well developed theories about a completely materialistic origin for the
Earth and his ideas questioning the fixity of species, were extremely influential.[24][25]
Between 1767 and 1792, James Burnett, Lord Monboddo included in his writings not only the
concept that man had descended from primates, but also that, in response to the environment, creatures had
found methods of transforming their characteristics over long time intervals.[26] Charles Darwin's
grandfather, Erasmus Darwin, published Zoönomia in 1796, which suggested that "all warm-blooded animals
have arisen from one living filament".[27] In his 1802 poem Temple of Nature, he described the rise of life
from minute organisms living in mud to all of its modern diversity.[28]
[edit] Early 19th century
 Diagram of the geologic timescale from an 1861 book by Richard Owen showing the appearance of
major animal types

[edit] Paleontology and geology

See also: History of paleontology
In 1796, Georges Cuvier published his findings on the differences between living elephants and
those found in the fossil record. His analysis demonstrated that mammoths and mastodons were distinct
species different from any living animal, effectively ending a long-running debate over whether the extinction
of a species was possible.[29] In 1788, James Hutton described gradual geological processes operating
continuously over deep time.[30] In the 1790s William Smith began the process of ordering rock strata by
examining fossils in the layers while he worked on his geologic map of England. Independently, in 1811,
Georges Cuvier and Alexandre Brongniart published an influential study of the geologic history of the region
around Paris, based on the stratigraphic succession of rock layers. These works helped establish the
antiquity of the Earth.[31] Cuvier advocated catastrophism to explain the patterns of extinction and faunal
succession revealed by the fossil record.
Knowledge of the fossil record continued to advance rapidly during the first few decades of the 19th
century. By the 1840s, the outlines of the geologic timescale were becoming clear, and in 1841 John Phillips
named three major eras, based on the predominant fauna of each: the Paleozoic, dominated by marine
invertebrates and fish, the Mesozoic, the age of reptiles, and the current Cenozoic age of mammals. This
progressive picture of the history of life was accepted even by conservative English geologists like Adam
Sedgwick and William Buckland; however, like Cuvier, they attributed the progression to repeated
catastrophic episodes of extinction followed by new episodes of creation.[32] Unlike Cuvier, Buckland and
some other advocates of natural theology among British geologists made efforts to explicitly link the last
catastrophic episode proposed by Cuvier to the biblical flood.[33][34]
From 1830 to 1833, Charles Lyell published his multi-volume work Principles of Geology, which,
building on Hutton's ideas, advocated a uniformitarian alternative to the catastrophic theory of geology. Lyell
claimed that, rather than being the products of cataclysmic (and possibly supernatural) events, the geologic
features of the Earth are better explained as the result of the same gradual geologic forces observable in the
present day—but acting over immensely long periods of time. Although Lyell opposed evolutionary ideas
(even questioning the consensus that the fossil record demonstrates a true progression), his concept that the
Earth was shaped by forces working gradually over an extended period, and the immense age of the Earth
assumed by his theories, would strongly influence future evolutionary thinkers such as Charles Darwin.[35]

[edit] Transmutation of species

Main article: Transmutation of species
 Diagram from Vestiges of the Natural History of Creation (1844) by Robert Chambers shows a model
of development where fish (F), reptiles (R), and birds (B) represent branches from a path leading to
mammals (M).
Jean-Baptiste Lamarck proposed, in his Philosophie Zoologique of 1809, a theory of the
transmutation of species. Lamarck did not believe that all living things shared a common ancestor but rather
that simple forms of life were created continuously by spontaneous generation. He also believed that an
innate life force drove species to become more complex over time, advancing up a linear ladder of
complexity that was related to the great chain of being. Lamarck recognized that species were adapted to
their environment. He explained this by saying that the same innate force driving increasing complexity
caused the organs of an animal (or a plant) to change based on the use or disuse of those organs, just as
muscles are affected by exercise. He argued that these changes would be inherited by the next generation
and produce slow adaptation to the environment. It was this secondary mechanism of adaptation through the
inheritance of acquired characteristics that would become known as Lamarckism and would influence
discussions of evolution into the 20th century.[36][37]
A radical British school of comparative anatomy that included the anatomist Robert Grant was closely
in touch with Lamarck's French school of Transformationism. One of the French scientists who influenced
Grant was the anatomist Étienne Geoffroy Saint-Hilaire, whose ideas on the unity of various animal body
plans and the homology of certain anatomical structures would be widely influential and lead to intense
debate with his colleague Georges Cuvier. Grant became an authority on the anatomy and reproduction of
marine invertebrates. He developed Lamarck's and Erasmus Darwin's ideas of transmutation and
evolutionism, and investigated homology, even proposing that plants and animals had a common
evolutionary starting point. As a young student Charles Darwin joined Grant in investigations of the life cycle
of marine animals. In 1826 an anonymous paper, probably written by Robert Jameson, praised Lamarck for
explaining how higher animals had "evolved" from the simplest worms; this was the first use of the word
"evolved" in a modern sense.[38][39]
In 1844, the Scottish publisher Robert Chambers anonymously published an extremely controversial
but widely read book entitled Vestiges of the Natural History of Creation. This book proposed an evolutionary
scenario for the origins of the Solar System and life on Earth. It claimed that the fossil record showed a
progressive ascent of animals with current animals being branches off a main line that leads progressively to
humanity. It implied that the transmutations lead to the unfolding of a preordained plan that had been woven
into the laws that governed the universe. In this sense it was less completely materialistic than the ideas of
radicals like Robert Grant, but its implication that humans were only the last step in the ascent of animal life
incensed many conservative thinkers. The high profile of the public debate over Vestiges, with its depiction of
evolution as a progressive process, would greatly influence the perception of Darwin's theory a decade later.
[40][41]
Ideas about the transmutation of species were associated with the radical materialism of the
Enlightenment and were attacked by more conservative thinkers. Georges Cuvier attacked the ideas of
Lamarck and Geoffroy Saint-Hilaire, agreeing with Aristotle that species were immutable. Cuvier believed
that the individual parts of an animal were too closely correlated with one another to allow for one part of the
anatomy to change in isolation from the others, and argued that the fossil record showed patterns of
catastrophic extinctions followed by re-population, rather than gradual change over time. He also noted that
drawings of animals and animal mummies from Egypt, which were thousands of years old, showed no signs
of change when compared with modern animals. The strength of Cuvier's arguments and his scientific
reputation helped keep transmutational ideas out of the mainstream for decades.[42]
 This 1847 diagram by Richard Owen shows his conceptual archetype for all vertebrates.
In Britain the philosophy of natural theology remained influential. William Paley's 1802 book Natural
Theology with its famous watchmaker analogy had been written at least in part as a response to the
transmutational ideas of Erasmus Darwin.[43] Geologists influenced by natural theology, such as Buckland
and Sedgwick, made a regular practice of attacking the evolutionary ideas of Lamarck, Grant, and The
Vestiges of the Natural History of Creation.[44][45] Although the geologist Charles Lyell opposed scriptural
geology, he also believed in the immutability of species, and in his Principles of Geology (1830–1833), he
criticized Lamarck's theories of development.[35] Idealists such as Louis Agassiz and Richard Owen believed
that each species was fixed and unchangeable because it represented an idea in the mind of the creator.
They believed that relationships between species could be discerned from developmental patterns in
embryology, as well as in the fossil record, but that these relationships represented an underlying pattern of
divine thought, with progressive creation leading to increasing complexity and culminating in humanity. Owen
developed the idea of "archetypes" in the Divine mind that would produce a sequence of species related by
anatomical homologies, such as vertebrate limbs. Owen led a public campaign that successfully
marginalized Robert Grant in the scientific community. Darwin would make good use of the homologies
analyzed by Owen in his own theory, but the harsh treatment of Grant, and the controversy surrounding
Vestiges, showed him the need to ensure that his own ideas were scientifically sound.[39][46][47]

[edit] Anticipations of natural selection

Several writers anticipated aspects of Darwin's theory, and in the third edition of On the Origin of
Species published in 1861 Darwin named those he knew about in an introductory appendix, An Historical
Sketch of the Recent Progress of Opinion on the Origin of Species , which he expanded in later editions.[48]
In 1813, William Charles Wells read before the Royal Society essays assuming that there had been
evolution of humans, and recognising the principle of natural selection. Charles Darwin and Alfred Russel
Wallace were unaware of this work when they jointly published the theory in 1858, but Darwin later
acknowledged that Wells had recognised the principle before them, writing that the paper "An Account of a
White Female, part of whose Skin resembles that of a Negro" was published in 1818, and "he distinctly
recognises the principle of natural selection, and this is the first recognition which has been indicated; but he
applies it only to the races of man, and to certain characters alone."[49] When Darwin was developing his
theory, he was influenced by Augustin de Candolle's natural system of classification, which laid emphasis on
the war between competing species.[50][51]
Patrick Matthew wrote in the obscure book Naval Timber & Arboriculture (1831) of "continual
balancing of life to circumstance. ... [The] progeny of the same parents, under great differences of
circumstance, might, in several generations, even become distinct species, incapable of co-
reproduction."[52] Charles Darwin discovered this work after the initial publication of the Origin. In the brief
historical sketch that Darwin included in the 3rd edition he says "Unfortunately the view was given by Mr.
Matthew very briefly in an Appendix to a work on a different subject ... He clearly saw, however, the full force
of the principle of natural selection."[53]
 It is possible to look through the history of biology from the ancient Greeks onwards and discover
anticipations of almost all of Darwin's key ideas. However, as historian of science Peter J. Bowler says,
"Through a combination of bold theorizing and comprehensive evaluation, Darwin came up with a concept of
evolution that was unique for the time." Bowler goes on to say that simple priority alone is not enough to
secure a place in the history of science; someone has to develop an idea and convince others of its
importance to have a real impact.[54]
T. H. Huxley said in his essay on the reception of the Origin of Species:
The suggestion that new species may result from the selective action of external conditions
upon the variations from their specific type which individuals present and which we call
spontaneous because we are ignorant of their causation is as wholly unknown to the historian of
scientific ideas as it was to biological specialists before 1858. But that suggestion is the central
idea of the Origin of Species, and contains the quintessence of Darwinism.[55]
 Darwin's first sketch of an evolutionary tree from his First Notebook on Transmutation of Species
(1837)

[edit] Natural selection

Main articles: Inception of Darwin's theory, Development of Darwin's theory, and Publication of
Darwin's theory
 The biogeographical patterns Charles Darwin observed in places such as the Galapagos islands
during the voyage of the Beagle caused him to doubt the fixity of species, and in 1837 Darwin started the first
of a series of secret notebooks on transmutation. Darwin's observations led him to view transmutation as a
process of divergence and branching, rather than the ladder-like progression envisioned by Lamarck and
others. In 1838 he read the new 6th edition of An Essay on the Principle of Population, written in the late
1700s by Thomas Malthus. Malthus' idea of population growth leading to a struggle for survival combined
with Darwin's knowledge on how breeders selected traits, led to the inception of Darwin's theory of natural
selection. Darwin did not publish his ideas on evolution for 20 years. However he did share them with certain
other naturalists and friends, starting with Joseph Hooker, with whom he discussed his unpublished 1844
essay on natural selection. During this period he used the time he could spare from his other scientific work
to slowly refine his ideas and, aware of the intense controversy around transmutation, amass evidence to
support them. In September 1854 he began full time work on writing his book on natural selection.[47][56][57]
Unlike Darwin, Alfred Russel Wallace, influenced by the book Vestiges of the Natural History of
Creation, already suspected that transmutation of species occurred when he began his career as a naturalist.
By 1855 his biogeographical observations during his field work in South America and the Malay Archipelago
made him confident enough in a branching pattern of evolution to publish a paper stating that every species
originated in close proximity to an already existing closely allied species. Like Darwin, it was Wallace's
consideration of how the ideas of Malthus might apply to animal populations that led him to conclusions very
similar to those reached by Darwin about the role of natural selection. In February 1858 Wallace, unaware of
Darwin's unpublished ideas, composed his thoughts into an essay and mailed them to Darwin, asking for his
opinion. The result was the joint publication in July of an extract from Darwin's 1844 essay along with
Wallace's letter. Darwin also began work on a short abstract summarising his theory, which he would publish
in 1859 as On the Origin of Species.[58]
 Diagram by O.C. Marsh of the evolution of horse feet and teeth over time as reproduced in T.H
Huxley's 1876 book Professor Huxley in America
[edit] 1859–1930s: Darwin and his legacy
See also: Reaction to Darwin's theory
By the 1850s whether or not species evolved was a subject of intense debate, with prominent
scientists arguing both sides of the issue.[59] However, it was the publication of Charles Darwin's On the
Origin of Species (1859) that fundamentally transformed the discussion over biological origins.[60] Darwin
argued that his branching version of evolution explained a wealth of facts in biogeography, anatomy,
embryology, and other fields of biology. He also provided the first cogent mechanism by which evolutionary
change could persist: his theory of natural selection.[61]
One of the first and most important naturalists to be convinced by Origin of the reality of evolution
was the British anatomist Thomas Henry Huxley. Huxley recognized that unlike the earlier transmutational
ideas of Lamarck and Vestiges, Darwin's theory provided a mechanism for evolution without supernatural
involvement, even if Huxley himself was not completely convinced that natural selection was the key
evolutionary mechanism. Huxley would make advocacy of evolution a cornerstone of the program of the X
Club to reform and professionalise science by displacing natural theology with naturalism and to end the
domination of British natural science by the clergy. By the early 1870s in English-speaking countries, thanks
partly to these efforts, evolution had become the mainstream scientific explanation for the origin of species.
[61] In his campaign for public and scientific acceptance of Darwin's theory, Huxley made extensive use of
new evidence for evolution from paleontology. This included evidence that birds had evolved from reptiles,
including the discovery of Archaeopteryx in Europe, and a number of fossils of primitive birds with teeth found
in North America. Another important line of evidence was the finding of fossils that helped trace the evolution
of the horse from its small five-toed ancestors.[62] However, acceptance of evolution among scientists in
non-English speaking nations such as France, and the countries of southern Europe and Latin America was
slower. An exception to this was Germany, where both August Weismann and Ernst Haeckel championed
this idea: Haeckel used evolution to challenge the established tradition of metaphysical idealism in German
biology, much as Huxley used it to challenge natural theology in Britain.[63] Haeckel and other German
scientists would take the lead in launching an ambitious programme to reconstruct the evolutionary history of
life based on morphology and embryology.[64]
Darwin's theory succeeded in profoundly altering scientific opinion regarding the development of life
and in producing a small philosophical revolution.[65] However, this theory could not explain several critical
components of the evolutionary process. Specifically, Darwin was unable to explain the source of variation in
traits within a species, and could not identify a mechanism that could pass traits faithfully from one
generation to the next. Darwin's hypothesis of pangenesis, while relying in part on the inheritance of acquired
characteristics, proved to be useful for statistical models of evolution that were developed by his cousin
Francis Galton and the "biometric" school of evolutionary thought. However, this idea proved to be of little
use to other biologists.[66]
[edit] Application to humans

This illustration was the frontispiece of Thomas Henry Huxley's book Evidence as to Man's Place in
Nature (1863).
Charles Darwin was aware of the severe reaction in some parts of the scientific community against
the suggestion made in Vestiges of the Natural History of Creation that humans had arisen from animals by a
process of transmutation. Therefore he almost completely ignored the topic of human evolution in The Origin
of Species. Despite this precaution, the issue featured prominently in the debate that followed the book's
publication. For most of the first half of the 19th century, the scientific community believed that, although
geology had shown that the Earth and life were very old, human beings had appeared suddenly just a few
thousand years before the present. However, a series of archaeological discoveries in the 1840s and 1850s
showed stone tools associated with the remains of extinct animals. By the early 1860s, as summarized in
Charles Lyell's 1863 book Geological Evidences of the Antiquity of Man , it had become widely accepted that
humans had existed during a prehistoric period – which stretched many thousands of years before the start of
written history. This view of human history was more compatible with an evolutionary origin for humanity than
was the older view. On the other hand, at that time there was no fossil evidence to demonstrate human
evolution. The only human fossils found before the discovery of Java man in the 1890s were either of
anatomically modern humans or of Neanderthals that were too close, especially in the critical characteristic
of cranial capacity, to modern humans for them to be convincing intermediates between humans and other
primates.[67]
Therefore the debate that immediately followed the publication of The Origin of Species centered on
the similarities and differences between humans and modern apes. Carolus Linnaeus had been criticised in
the 18th century for grouping humans and apes together as primates in his ground breaking classification
system.[68] Richard Owen vigorously defended the classification suggested by Cuvier and Johann Friedrich
Blumenbach that placed humans in a separate order from any of the other mammals, which by the early 19th
century had become the orthodox view. On the other hand, Thomas Henry Huxley sought to demonstrate a
close anatomical relationship between humans and apes. In one famous incident, Huxley showed that Owen
was mistaken in claiming that the brains of gorillas lacked a structure present in human brains. Huxley
summarized his argument in his highly influential 1863 book Evidence as to Man's Place in Nature. Another
viewpoint was advocated by Charles Lyell and Alfred Russel Wallace. They agreed that humans shared a
common ancestor with apes, but questioned whether any purely materialistic mechanism could account for
all the differences between humans and apes, especially some aspects of the human mind.[67]
In 1871, Darwin published The Descent of Man, and Selection in Relation to Sex , which contained
his views on human evolution. Darwin argued that the differences between the human mind and the minds of
the higher animals were a matter of degree rather than of kind. For example, he viewed morality as a natural
outgrowth of instincts that were beneficial to animals living in social groups. He argued that all the differences
between humans and apes were explained by a combination of the selective pressures that came from our
ancestors moving from the trees to the plains, and sexual selection. The debate over human origins, and
over the degree of human uniqueness continued well into the 20th century.[67]

[edit] Alternatives to natural selection

Main article: The eclipse of Darwinism
 This photo from Henry Fairfield Osborn's 1918 book Origin and Evolution of Life shows models
depicting the evolution of Titanothere horns over time, which Osborn claimed was an example of an
orthogenic trend in evolution.
The concept of evolution was widely accepted in scientific circles within a few years of the publication
of Origin, but the acceptance of natural selection as its driving mechanism was much less widespread. The
four major alternatives to natural selection in the late 19th century were theistic evolution, neo-Lamarckism,
orthogenesis, and saltationism. Theistic evolution (a term promoted by Darwin's greatest American advocate
Asa Gray) was the idea that God intervened in the process of evolution to guide it in such a way that the
living world could still be considered to be designed. However, this idea gradually fell out of favor among
scientists, as they became more and more committed to the idea of methodological naturalism and came to
believe that direct appeals to supernatural involvement were scientifically unproductive. By 1900, theistic
evolution had largely disappeared from professional scientific discussions, although it retained a strong
popular following.[69][70]
In the late 19th century, the term neo-Lamarckism came to be associated with the position of
naturalists who viewed the inheritance of acquired characteristics as the most important evolutionary
mechanism. Advocates of this position included the British writer and Darwin critic Samuel Butler, the
German biologist Ernst Haeckel, and the American paleontologist Edward Drinker Cope. They considered
Lamarckism to be philosophically superior to Darwin's idea of selection acting on random variation. Cope
looked for, and thought he found, patterns of linear progression in the fossil record. Inheritance of acquired
characteristics was part of Haeckel's recapitulation theory of evolution, which held that the embryological
development of an organism repeats its evolutionary history.[69][70] Critics of neo-Lamarckism, such as the
German biologist August Weismann and Alfred Russel Wallace, pointed out that no one had ever produced
solid evidence for the inheritance of acquired characteristics. Despite these criticisms, neo-Lamarckism
remained the most popular alternative to natural selection at the end of the 19th century, and would remain
the position of some naturalists well into the 20th century.[69][70]
Orthogenesis was the hypothesis that life has an innate tendency to change, in a unilinear fashion,
towards ever-greater perfection. It had a significant following in the 19th century, and its proponents included
the Russian biologist Leo Berg and the American paleontologist Henry Fairfield Osborn. Orthogenesis was
popular among some paleontologists, who believed that the fossil record showed a gradual and constant
unidirectional change. Saltationism was the idea that new species arise as a result of large mutations. It was
seen as a much faster alternative to the Darwinian concept of a gradual process of small random variations
being acted on by natural selection, and was popular with early geneticists such as Hugo de Vries, William
Bateson, and early in his career, T. H. Morgan. It became the basis of the mutation theory of evolution.[69]
[70]
 Diagram from T.H. Morgan's 1919 book The Physical Basis of Heredity, showing the sex-linked
inheritance of the white-eyed mutation in Drosophila melanogaster

[edit] Mendelian genetics, biometrics, and mutation

The so-called rediscovery of Gregor Mendel's laws of inheritance in 1900 ignited a fierce debate
between two camps of biologists. In one camp were the Mendelians, who were focused on discrete variations
and the laws of inheritance. They were led by William Bateson (who coined the word genetics) and Hugo de
Vries (who coined the word mutation). Their opponents were the biometricians, who were interested in the
continuous variation of characteristics within populations. Their leaders, Karl Pearson and Walter Frank
Raphael Weldon, followed in the tradition of Francis Galton, who had focused on measurement and statistical
analysis of variation within a population. The biometricians rejected Mendelian genetics on the basis that
discrete units of heredity, such as genes, could not explain the continuous range of variation seen in real
populations. Weldon's work with crabs and snails provided evidence that selection pressure from the
environment could shift the range of variation in wild populations, but the Mendelians maintained that the
variations measured by biometricians were too insignificant to account for the evolution of new species.[71]
[72]
When T. H. Morgan began experimenting with breeding the fruit fly Drosophila melanogaster, he was
a saltationist who hoped to demonstrate that a new species could be created in the lab by mutation alone.
Instead, the work at his lab between 1910 and 1915 reconfirmed Mendelian genetics and provided solid
experimental evidence linking it to chromosomal inheritance. His work also demonstrated that most
mutations had relatively small effects, such as a change in eye color, and that rather than creating a new
species in a single step, mutations served to increase variation within the existing population.[71][72]
[edit] 1920s–1940s
See also: Modern evolutionary synthesis

Biston betularia f. typica is the white-bodied form of the peppered moth.

Biston betularia f. carbonaria is the black-bodied form of the peppered moth.

[edit] Population genetics
The Mendelian and biometrician models were eventually reconciled with the development of
population genetics. A key step was the work of the British biologist and statistician R.A. Fisher. In a series of
papers starting in 1918 and culminating in his 1930 book The Genetical Theory of Natural Selection, Fisher
showed that the continuous variation measured by the biometricians could be produced by the combined
action of many discrete genes, and that natural selection could change gene frequencies in a population,
resulting in evolution. In a series of papers beginning in 1924, another British geneticist, J.B.S. Haldane,
applied statistical analysis to real-world examples of natural selection, such as the evolution of industrial
melanism in peppered moths, and showed that natural selection worked at an even faster rate than Fisher
assumed.[73][74]
The American biologist Sewall Wright, who had a background in animal breeding experiments,
focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated
populations that exhibited genetic drift. In 1932, Wright introduced the concept of an adaptive landscape and
argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive
peak, allowing natural selection to drive it towards different adaptive peaks. The work of Fisher, Haldane and
Wright founded the discipline of population genetics. This integrated natural selection with Mendelian
genetics, which was the critical first step in developing a unified theory of how evolution worked.[73][74]

[edit] Modern evolutionary synthesis

In the first few decades of the 20th century, most field naturalists continued to believe that
Lamarckian and orthogenic mechanisms of evolution provided the best explanation for the complexity they
observed in the living world. But as the field of genetics continued to develop, those views became less
tenable.[75] Theodosius Dobzhansky, a postdoctoral worker in T. H. Morgan's lab, had been influenced by
the work on genetic diversity by Russian geneticists such as Sergei Chetverikov. He helped to bridge the
divide between the foundations of microevolution developed by the population geneticists and the patterns of
macroevolution observed by field biologists, with his 1937 book Genetics and the Origin of Species.
Dobzhansky examined the genetic diversity of wild populations and showed that, contrary to the assumptions
of the population geneticists, these populations had large amounts of genetic diversity, with marked
differences between sub-populations. The book also took the highly mathematical work of the population
geneticists and put it into a more accessible form. In Great Britain E.B. Ford, the pioneer of ecological
genetics, continued throughout the 1930s and 1940s to demonstrate the power of selection due to ecological
factors including the ability to maintain genetic diversity through genetic polymorphisms such as human
blood types. Ford's work would contribute to a shift in emphasis during the course of the modern synthesis
towards natural selection over genetic drift.[73][74][76][77]
Evolutionary biologist Ernst Mayr was influenced by the work of the German biologist Bernhard
Rensch showing the influence of local environmental factors on the geographic distribution of sub-species
and closely related species. Mayr followed up on Dobzhansky's work with the 1942 book Systematics and
the Origin of Species, which emphasized the importance of allopatric speciation in the formation of new
species. This form of speciation occurs when the geographical isolation of a sub-population is followed by the
development of mechanisms for reproductive isolation. Mayr also formulated the biological species concept
that defined a species as a group of interbreeding or potentially interbreeding populations that were
reproductively isolated from all other populations.[73][74][78]
In the 1944 book Tempo and Mode in Evolution, George Gaylord Simpson showed that the fossil
record was consistent with the irregular non-directional pattern predicted by the developing evolutionary
synthesis, and that the linear trends that earlier paleontologists had claimed supported orthogenesis and
neo-Lamarckism did not hold up to closer examination. In 1950, G. Ledyard Stebbins published Variation
and Evolution in Plants, which helped to integrate botany into the synthesis. The emerging cross-disciplinary
consensus on the workings of evolution would be known as the modern evolutionary synthesis. It received its
name from the book Evolution: The Modern Synthesis by Julian Huxley.[73][74]
The evolutionary synthesis provided a conceptual core—in particular, natural selection and Mendelian
population genetics—that tied together many, but not all, biological disciplines. It helped establish the
legitimacy of evolutionary biology, a primarily historical science, in a scientific climate that favored
experimental methods over historical ones.[79] The synthesis also resulted in a considerable narrowing of
the range of mainstream evolutionary thought (what Stephen Jay Gould called the "hardening of the
synthesis"): by the 1950s, natural selection acting on genetic variation was virtually the only acceptable
mechanism of evolutionary change (panselectionism), and macroevolution was simply considered the result
of extensive microevolution.[80][81]

[edit] 1940s–1960s: Molecular biology and evolution

Main article: History of molecular evolution
The middle decades of the 20th century saw the rise of molecular biology, and with it an
understanding of the chemical nature of genes as sequences of DNA and their relationship, through the
genetic code, to protein sequences. At the same time, increasingly powerful techniques for analyzing
proteins, such as protein electrophoresis and sequencing, brought biochemical phenomena into realm of the
synthetic theory of evolution. In the early 1960s, biochemists Linus Pauling and Emile Zuckerkandl proposed
the molecular clock hypothesis: that sequence differences between homologous proteins could be used to
calculate the time since two species diverged. By 1969, Motoo Kimura and others provided a theoretical
basis for the molecular clock, arguing that—at the molecular level at least—most genetic mutations are neither
harmful nor helpful and that genetic drift, rather than natural selection, is responsible for a large portion of
genetic change: the neutral theory of molecular evolution.[82] Studies of protein differences within species
also brought molecular data to bear on population genetics by providing estimates of the level of
heterozygosity in natural populations.[83]
From the early 1960s, molecular biology was increasingly seen as a threat to the traditional core of
evolutionary biology. Established evolutionary biologists—particularly Ernst Mayr, Theodosius Dobzhansky
and G. G. Simpson, three of the architects of the modern synthesis—were extremely skeptical of molecular
approaches, especially when it came to the connection (or lack thereof) to natural selection. The molecular
clock hypothesis and the neutral theory were particularly controversial, spawning the neutralist-selectionist
debate over the relative importance of drift and selection, which continued into the 1980s without a clear
resolution.[84][85]

[edit] Late 20th century

[edit] Gene-centered view
In the mid-1960s, George C. Williams strongly critiqued explanations of adaptations worded in terms
of "survival of the species" (group selection arguments). Such explanations were largely replaced by a gene-
centered view of evolution, epitomized by the kin selection arguments of W. D. Hamilton, George R. Price
and John Maynard Smith.[86] This viewpoint would be summarized and popularized in the influential 1976
book The Selfish Gene by Richard Dawkins.[87] Models of the period showed that group selection was
severely limited in its strength; though newer models do admit the possibility of significant multi-level
selection.[88]
 In 1973, Leigh Van Valen proposed the term "Red Queen", which he took from Through the Looking-
Glass by Lewis Carroll, to describe a scenario where a species involved in one or more evolutionary arms
races would have to constantly change just to keep pace with the species with which it was co-evolving.
Hamilton, Williams and others suggested that this idea might explain the evolution of sexual reproduction: the
increased genetic diversity caused by sexual reproduction would help maintain resistance against rapidly
evolving parasites, thus making sexual reproduction common, despite the tremendous cost from the gene-
centric point of view of a system where only half of an organism's genome is passed on during reproduction.
[89][90] The gene-centric view has also led to an increased interest in Darwin's old idea of sexual selection,
[91] and more recently in topics such as sexual conflict and intragenomic conflict.

[edit] Sociobiology
W. D. Hamilton's work on kin selection contributed to the emergence of the discipline of sociobiology.
The existence of altruistic behaviors has been a difficult problem for evolutionary theorists from the
beginning.[92] Significant progress was made in 1964 when Hamilton formulated the inequality in kin
selection known as Hamilton's rule, which showed how eusociality in insects (the existence of sterile worker
classes) and many other examples of altruistic behavior could have evolved through kin selection. Other
theories followed, some derived from game theory, such as reciprocal altruism.[93] In 1975, E.O. Wilson
published the influential and highly controversial book Sociobiology: The New Synthesis which claimed
evolutionary theory could help explain many aspects of animal, including human, behavior. Critics of
sociobiology, including Stephen Jay Gould and Richard Lewontin, claimed that sociobiology greatly
overstated the degree to which complex human behaviors could be determined by genetic factors. They also
claimed that the theories of sociobiologists often reflected their own ideological biases. Despite these
criticisms, work has continued in sociobiology and the related discipline of evolutionary psychology, including
work on other aspects of the altruism problem.[94][95]
 A phylogenetic tree showing the three-domain system. Eukaryotes are colored red, Archaea green,
and Bacteria blue.

[edit] Evolutionary paths and processes

One of the most prominent debates arising during the 1970s was over the theory of punctuated
equilibrium. Niles Eldredge and Stephen Jay Gould proposed that there was a pattern of fossil species that
remained largely unchanged for long periods (what they termed stasis), interspersed with relatively brief
periods of rapid change during speciation.[96][97] Improvements in sequencing methods resulted in a large
increase of sequenced genomes, allowing the testing and refining of evolutionary theories using this huge
amount of genome data.[98] Comparisons between these genomes provide insights into the molecular
mechanisms of speciation and adaptation.[99][100] These genomic analyses have produced fundamental
changes in the understanding of the evolutionary history of life, such as the proposal of the three-domain
system by Carl Woese.[101] Advances in computational hardware and software allow the testing and
extrapolation of increasingly advanced evolutionary models and the development of the field of systems
biology.[102] One of the results has been an exchange of ideas between theories of biological evolution and
the field of computer science known as evolutionary computation, which attempts to mimic biological
evolution for the purpose of developing new computer algorithms. Discoveries in biotechnology now allow the
modification of entire genomes, advancing evolutionary studies to the level where future experiments may
involve the creation of entirely synthetic organisms.[103]

[edit] Microbiology, horizontal gene transfer, and endosymbiosis

Main article: Horizontal gene transfer
Microbiology was largely ignored by early evolutionary theory. This was due to the paucity of
morphological traits and the lack of a species concept in microbiology, particularly amongst prokaryotes.[104]
Now, evolutionary researchers are taking advantage of their improved understanding of microbial physiology
and ecology, produced by the comparative ease of microbial genomics, to explore the taxonomy and
evolution of these organisms.[105] These studies are revealing unanticipated levels of diversity amongst
microbes.[106][107]
One particularly important outcome from studies on microbial evolution was the discovery in Japan of
horizontal gene transfer in 1959.[108] This transfer of genetic material between different species of bacteria
was first recognized because it played a major role in the spread of antibiotic resistance.[109] More recently,
as knowledge of genomes has continued to expand, it has been suggested that lateral transfer of genetic
material has played an important role in the evolution of all organisms.[110] These high levels of horizontal
gene transfer have led to suggestions that the family tree of today's organisms, the so-called "tree of life", is
more similar to an interconnected web or net.[111][112]
 Indeed, as part of the endosymbiotic theory for the origin of organelles, a form of horizontal gene
transfer has been a critical step in the evolution of eukaryotes such as fungi, plants, and animals.[113][114]
The endosymbiotic theory holds that organelles within the cells of eukorytes such as mitochondria and
chloroplasts, had descended from independent bacteria that came to live symbiotically within other cells. It
had been suggested in the late 19th century when similarities between mitochondria and bacteria were
noted, but largely dismissed until it was revived and championed by Lynn Margulis in the 1960s and 70s;
Margulis was able to make use of new evidence that such organelles had their own DNA that was inherited
independently from that in the cell's nucleus.[115]

[edit] Evolutionary developmental biology

Main article: Evolutionary developmental biology
In the 1980s and 1990s the tenets of the modern evolutionary synthesis came under increasing
scrutiny. There was a renewal of structuralist themes in evolutionary biology in the work of biologists such as
Brian Goodwin and Stuart Kauffman, which incorporated ideas from cybernetics and systems theory, and
emphasized the self-organizing processes of development as factors directing the course of evolution. The
evolutionary biologist Stephen Jay Gould revived earlier ideas of heterochrony, alterations in the relative
rates of developmental processes over the course of evolution, to account for the generation of novel forms,
and, with the evolutionary biologist Richard Lewontin, wrote an influential paper in 1979 suggesting that a
change in one biological structure, or even a structural novelty, could arise incidentally as an accidental
result of selection on another structure, rather than through direct selection for that particular adaptation.
They called such incidental structural changes "spandrels" after an architectural feature.[116] Later, Gould
and Vrba discussed the acquisition of new functions by novel structures arising in this fashion, calling them
"exaptations".[117]
 Molecular data regarding the mechanisms underlying development accumulated rapidly during the
1980s and '90s. It became clear that the diversity of animal morphology was not the result of different sets of
proteins regulating the development of different animals, but from changes in the deployment of a small set
of proteins that were common to all animals.[118] These proteins became known as the "developmental
toolkit".[119] Such perspectives influenced the disciplines of phylogenetics, paleontology and comparative
developmental biology, and spawned the new discipline of evolutionary developmental biology.[120]
More recent work in this field by Mary Jane West-Eberhard has emphasized phenotypic and
developmental plasticity.[121] It has been suggested, for example, that the rapid emergence of basic animal
body plans in the Cambrian explosion was due in part to changes in the environment acting on inherent
material properties of cell aggregates, such as differential cell adhesion and biochemical oscillation. The
resulting forms were later stabilized by natural selection.[122] Experimental and theoretical research on
these and related ideas have been presented in the multi-authored volume Origination of Organismal Form.

[edit] 21st century

[edit] Epigenetic inheritance
Main article: Epigenetics
been no genetic changes. This shows that in some cases non genetic changes to an organism can be
inherited and it has been suggested that such inheritance can help with adaptation to local conditions and
affect evolution.[123][124] Some have suggested that in some cases a form of Lamarckian evolution may
occur.[125]

[edit] Unconventional evolutionary theory

[edit] Omega Point
Pierre Teilhard de Chardin's metaphysical Omega Point Theory describes the gradual development
of the universe from subatomic particles to human society, which he viewed as its final stage and goal.

[edit] Gaia hypothesis

Teilhard de Chardin's ideas have been seen as being connected to the more specific Gaia theory by
James Lovelock, who proposed that the living and nonliving parts of Earth can be viewed as a complex
interacting system with similarities to a single organism.[126] The Gaia hypothesis has also been viewed by
Lynn Margulis[127] and others as an extension of endosymbiosis and exosymbiosis.[128] This modified
hypothesis postulates that all living things have a regulatory effect on the Earth's environment that promotes
life overall.
[edit] Transhumanism
Futurists have often viewed scientific and technological progress as a continuation of biological
evolution. Among these, transhumanists often view such technological evolution itself as a goal in their
philosophy, possibly in the form of a technological singularity.

[edit] See also

Evolutionary biology portal

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Faith and rationality
• Galápagos Islands
• The Voyage of the Beagle

[edit] Notes
1. ^ Couprie, Dirk L.. "Anaximander". Internet Encyclopedia of Philosophy.
http://www.iep.utm.edu/anaximan/#H8. Retrieved 2010-02-27.
 2. ^ Campbell, Gordon. "Empedocles". Internet Encyclopedia of Philosophy.
http://www.iep.utm.edu/e/empedocl.htm#H4. Retrieved 2008-07-15.
3. ^ Hardie, R.P.; R. K. Gaye. "Physics by Aristotle".
http://classics.mit.edu/Aristotle/physics.2.ii.html. Retrieved 2008-07-15.
4. ^ Mayr 1982, p. 304
5. ^ a b c d e f Johnston, Ian (1999). "Section Three: The Origins of Evolutionary Theory". ... And
Still We Evolve: A Handbook on the History of Modern Science . Liberal Studies Department,
Malaspina University College. http://records.viu.ca/~johnstoi/darwin/sect3.htm. Retrieved 2007-08-
11.
6. ^ a b Singer 1931
7. ^ Needham & Ronan 1995, p. 101
8. ^ Miller, James (January 8, 2008). "Daoism and Nature" (PDF). Royal Asiatic Society.
http://www.jamesmiller.ca/RAS%20lecture%20on%20daoism%20and%20nature.pdf. Retrieved
2008-07-15.
9. ^ Sedley, David (August 4, 2004). "Lucretius". Stanford Encyclopedia of Philosophy.
http://plato.stanford.edu/entries/lucretius/. Retrieved 2008-07-24.
10.^ Simpson, David (2006). "Lucretius". The Internet Encyclopedia of Philosophy.
http://www.iep.utm.edu/l/lucretiu.htm. Retrieved 2008-07-24.
11.^ Augustine 1982, pp. 89–90
12.^ Gill 2005, p. 251
13.^ Owen, Richard (2009-02-11). "Vatican buries the hatchet with Charles Darwin". London:
Times Online. http://www.timesonline.co.uk/tol/comment/faith/article5705331.ece. Retrieved 2009-
02-12.
 14.^ Irvine, Chris (2009-02-12). "The Vatican claims Darwin's theory of evolution is compatible
with Christianity". London: Telegraph.co.uk.
http://www.telegraph.co.uk/news/newstopics/religion/4588289/The-Vatican-claims-Darwins-theory-
of-evolution-is-compatible-with-Christianity.html. Retrieved 2009-02-12.
15.^ "Medieval and Renaissance Concepts of Evolution and Paleontology". University of
California Museum of Paleontology. http://www.ucmp.berkeley.edu/history/medieval.html. Retrieved
11-3-2010.
16.^ a b Conway Zirkle (1941). Natural Selection before the "Origin of Species", Proceedings of
the American Philosophical Society 84 (1), pp. 71–123.
17.^ Frank N. Egerton, "A History of the Ecological Sciences, Part 6: Arabic Language Science –
Origins and Zoological", Bulletin of the Ecological Society of America , April 2002: 142–146 [143]
18.^ Lovejoy 1936, pp. 67–80
19.^ Carroll, William E.. "Creation, Evolution, and Thomas Aquinas". Catholic Education
Resource Center. http://www.catholiceducation.org/articles/science/sc0035.html. Retrieved 2010-03-
20.
20.^ Thomas Aquinas. "Physica Book 2, Lecture 14". Fathers of the English Dominican
Province. http://dhspriory.org/thomas/Physics2.htm#14. Retrieved 2010-03-23.
21.^ Bowler 2003, pp. 33–38
22.^ Schelling, System of Transcendental Idealism, 1800
23.^ Bowler 2003, pp. 73–75
24.^ Bowler 2003, pp. 75–80
25.^ Larson 2004, pp. 14–15
26.^ Henderson 2000
27.^ Darwin 1818, Vol I section XXXIX
 28.^ Darwin 1825, p. 15
29.^ Larson 2004, p. 7
30.^ American Museum of Natural History (2000). "James Hutton: The Founder of Modern
Geology". Earth: Inside and Out.
http://www.amnh.org/education/resources/rfl/web/essaybooks/earth/p_hutton.html. "we find no
vestige of a beginning, no prospect of an end."
31.^ Bowler 2003, p. 113
32.^ Larson 2004, pp. 29–38
33.^ Bowler 2003, pp. 115–116
34.^ "Darwin and design: historical essay". Darwin Correspondence Project.
http://www.darwinproject.ac.uk/content/view/110/104/. Retrieved 2008-01-17.
35.^ a b Bowler 2003, pp. 129–134
36.^ Bowler 2003, pp. 86–94
37.^ Larson 2004, pp. 38–41
38.^ Desmon & Moore 1993, p. 40
39.^ a b Bowler 2003, pp. 120–129
40.^ Bowler 2003, pp. 134–138
41.^ Bowler & Morus 2005, pp. 142–143
42.^ Larson 2004, pp. 5–24
43.^ Bowler 2003, pp. 103–104
44.^ Larson 2004, pp. 37–38
45.^ Bowler 2003, p. 138
46.^ Larson 2004, pp. 42–46
 47.^ a b van Wyhe, John (27 March 2007). "Mind the gap: Did Darwin avoid publishing his theory
for many years?". pp. 177–205. doi:10.1098/rsnr.2006.0171. http://darwin-
online.org.uk/content/frameset?viewtype=text&itemID=A544&pageseq=1. Retrieved 2009-11-17.
48.^ Darwin 1861, p. xiii
49.^ Darwin 1866, p. xiv
50.^ Bowler 2003, p. 151
51.^ Darwin 1859, pp. 62
52.^ Matthew, Patrick (1860). "Nature's law of selection. Gardeners' Chronicle and Agricultural
Gazette". The Complete Works of Charles Darwin Online. http://darwin-
online.org.uk/content/frameset?itemID=A143&viewtype=text&pageseq=1. Retrieved 2007-11-01.
53.^ Darwin 1861, p. xiv
54.^ Bowler 2003, p. 158
55.^ Huxley, Thomas Henry (1895). "The Reception of the Origin of Species". Project
Gutenberg. http://infomotions.com/etexts/gutenberg/dirs/etext00/oroos10.htm. Retrieved 2007-11-
02.
56.^ Bowler & Morus 2005, pp. 129–149
57.^ Larson 2004, pp. 55–71
58.^ Bowler 2003, pp. 173–176
59.^ Larson 2004, p. 50
60.^ The centrality of Origin of Species in the rise of widespread evolutionary thinking has been
has long been accepted by historians of science. However, some scholars have recently begun to
challenge this idea. James A. Secord, in his study of the impact of Vestiges of the Natural History of
Creation, argues that in some ways Vestiges had as much or more impact than Origin, at least into
the 1880s. Focusing so much on Darwin and Origin, he argues, "obliterates decades of labor by
teachers, theologians, technicians, printers, editors, and other researchers, whose work has made
evolutionary debates so significant during the past two centuries." Secord 2000, pp. 515–518
61.^ a b Larson 2004, pp. 79–111
62.^ Larson 2004, pp. 139–40
63.^ Larson 2004, pp. 109–110
64.^ Bowler 2003, pp. 190–191
65.^ Bowler 2003, pp. 177–223
66.^ Larson 2004, pp. 121–123, 152–157
67.^ a b c Bowler 2003, pp. 207–216
68.^ Bowler 2003, pp. 49–51
69.^ a b c d Larson 2004, pp. 105–129
70.^ a b c d Bowler 2003, pp. 196–253
71.^ a b Bowler 2003, pp. 256–273
72.^ a b Larson 2004, pp. 153–174
73.^ a b c d e Bowler 2003, pp. 325–339
74.^ a b c d e Larson 2004, pp. 221–243
75.^ Mayr & Provine 1998, pp. 295–298, 416
76.^ Mayr, E§year=1988. Towards a new philosophy of biology: observations of an evolutionist .
Harvard University Press. p. 402.
77.^ Mayr & Provine 1998, pp. 338–341
78.^ Mayr & Provine 1998, pp. 33–34
79.^ Smocovitis 1996, pp. 97–188
80.^ Sapp 2003, pp. 152–156
 81.^ Gould, Stephen Jay (1983). "The hardening of the modern synthesis". in Marjorie Grene.
Dimensions of Darwinism. Cambridge University Press.
http://www.stephenjaygould.org/library/gould_synthesis.html.
82.^ Dietrich, Michael R. (1994-03-01). "The origins of the neutral theory of molecular evolution".
Journal of the History of Biology 27 (1): 21–59. doi:10.1007/BF01058626. PMID 11639258.
83.^ Powell, Jeffrey R (1994) [1994]. "Molecular techniques in population genetics: A brief
history". in B. Schierwater, B. Streit, G. P. Wagner, and R. De Salle (eds.). Molecular Ecology and
Evolution: Approaches and Applications. Birkhäuser Verlag. pp. 131–156. ISBN 3-7643-2942-4.
84.^ Dietrich, Michael R. (1998-03-01). "Paradox and Persuasion: Negotiating the Place of
Molecular Evolution within Evolutionary Biology". Journal of the History of Biology 31 (1): 85–111.
doi:10.1023/A:1004257523100. PMID 11619919.
85.^ Hagen, JB (1999). "Naturalists, Molecular Biologists, and the Challenges of Molecular
Evolution". Journal of the History of Biology 32 (2): 321–341. doi:10.1023/A:1004660202226.
PMID 11624208.
86.^ Mayr E (1997). "The objects of selection". Proc. Natl. Acad. Sci. U.S.A. 94 (6): 2091–94.
doi:10.1073/pnas.94.6.2091. PMID 9122151. PMC 33654.
http://www.pnas.org/cgi/content/full/94/6/2091.
87.^ Bowler 2003, p. 361
88.^ Gould SJ (1998). "Gulliver's further travels: the necessity and difficulty of a hierarchical
theory of selection". Philos. Trans. R. Soc. Lond., B, Biol. Sci. 353 (1366): 307–14.
doi:10.1098/rstb.1998.0211. PMID 9533127. PMC 1692213.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=9533127.
89.^ Larson 2004, p. 279
90.^ Bowler 2003, p. 358
 91.^ Bowler 2003, pp. 358–359
92.^ Sachs J (2006). "Cooperation within and among species". J. Evol. Biol. 19 (5): 1415–8;
discussion 1426–36. doi:10.1111/j.1420-9101.2006.01152.x. PMID 16910971.
93.^ Nowak M (2006). "Five rules for the evolution of cooperation". Science 314 (5805): 1560–
63. doi:10.1126/science.1133755. PMID 17158317.
94.^ Larson 2004, pp. 270–278
95.^ Bowler 2003, pp. 359–361
96.^ Niles Eldredge and Stephen Jay Gould, 1972. "Punctuated equilibria: an alternative to
phyletic gradualism" In T.J.M. Schopf, ed., Models in Paleobiology. San Francisco: Freeman Cooper.
pp. 82–115. Reprinted in N. Eldredge Time frames. Princeton: Princeton Univ. Press. 1985
97.^ Gould SJ (1994). "Tempo and mode in the macroevolutionary reconstruction of Darwinism".
Proc. Natl. Acad. Sci. U.S.A. 91 (15): 6764–71. doi:10.1073/pnas.91.15.6764. PMID 8041695.
PMC 44281. http://www.pnas.org/cgi/reprint/91/15/6764.
98.^ Pollock DD, Eisen JA, Doggett NA, Cummings MP (1 December 2000). "A case for
evolutionary genomics and the comprehensive examination of sequence biodiversity". Mol. Biol.
Evol. 17 (12): 1776–88. PMID 11110893. http://mbe.oxfordjournals.org/cgi/content/full/17/12/1776.
99.^ Koonin EV (2005). "Orthologs, paralogs, and evolutionary genomics". Annu. Rev. Genet.
39: 309–38. doi:10.1146/annurev.genet.39.073003.114725. PMID 16285863.
100.^ Hegarty MJ, Hiscock SJ (2005). "Hybrid speciation in plants: new insights from molecular
studies". New Phytol. 165 (2): 411–23. doi:10.1111/j.1469-8137.2004.01253.x. PMID 15720652.
101.^ Woese C, Kandler O, Wheelis M (1990). "Towards a natural system of organisms:
proposal for the domains Archaea, Bacteria, and Eucarya". Proc Natl Acad Sci USA 87 (12): 4576–
79. doi:10.1073/pnas.87.12.4576. PMID 2112744. PMC 54159.
http://www.pnas.org/cgi/reprint/87/12/4576.
 102.^ Medina M (2005). "Genomes, phylogeny, and evolutionary systems biology". Proc. Natl.
Acad. Sci. U.S.A. 102 Suppl 1: 6630–5. doi:10.1073/pnas.0501984102. PMID 15851668.
PMC 1131869. http://www.pubmedcentral.nih.gov/articlerender.fcgi?
tool=pubmed&pubmedid=15851668.
103.^ Benner SA, Sismour AM (2005). "Synthetic biology". Nat. Rev. Genet. 6 (7): 533–43.
doi:10.1038/nrg1637. PMID 15995697.
104.^ Gevers D, Cohan FM, Lawrence JG, et al. (2005). "Opinion: Re-evaluating prokaryotic
species". Nat. Rev. Microbiol. 3 (9): 733–9. doi:10.1038/nrmicro1236. PMID 16138101.
105.^ Coenye T, Gevers D, Van de Peer Y, Vandamme P, Swings J (2005). "Towards a
prokaryotic genomic taxonomy". FEMS Microbiol. Rev. 29 (2): 147–67.
doi:10.1016/j.femsre.2004.11.004. PMID 15808739.
106.^ Whitman W, Coleman D, Wiebe W (1998). "Prokaryotes: the unseen majority". Proc Natl
Acad Sci USA 95 (12): 6578–83. doi:10.1073/pnas.95.12.6578. PMID 9618454. PMC 33863.
http://www.pnas.org/cgi/content/full/95/12/6578.
107.^ Schloss P, Handelsman J (2004). "Status of the microbial census". Microbiol Mol Biol Rev
68 (4): 686–91. doi:10.1128/MMBR.68.4.686-691.2004. PMID 15590780. PMC 539005.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=15590780#r6.
108.^ Ochiai K, Yamanaka T, Kimura K Sawada O (1959). "Inheritance of drug resistance (and
its transfer) between Shigella strains and Between Shigella and E.coli strains". Hihon Iji Shimpor
1861: 34. (in Japanese)
109.^ "Lateral gene transfer and the nature of bacterial innovation" (PDF). Nature Vol 405, May
18, 2000. http://www.stat.rice.edu/~mathbio/Ochman2000.pdf. Retrieved 2007-09-01.
 110.^ de la Cruz F, Davies J (2000). "Horizontal gene transfer and the origin of species: lessons
from bacteria". Trends Microbiol. 8 (3): 128–33. doi:10.1016/S0966-842X(00)01703-0.
PMID 10707066.
111.^ Kunin V, Goldovsky L, Darzentas N, Ouzounis CA (2005). "The net of life: reconstructing
the microbial phylogenetic network". Genome Res. 15 (7): 954–9. doi:10.1101/gr.3666505.
PMID 15965028. PMC 1172039. http://www.genome.org/cgi/pmidlookup?
view=long&pmid=15965028.
112.^ Doolittle WF, Bapteste E (February 2007). "Pattern pluralism and the Tree of Life
hypothesis". Proc. Natl. Acad. Sci. U.S.A. 104 (7): 2043–9. doi:10.1073/pnas.0610699104.
PMID 17261804. PMC 1892968. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=17261804.
113.^ Poole A, Penny D (2007). "Evaluating hypotheses for the origin of eukaryotes". Bioessays
29 (1): 74–84. doi:10.1002/bies.20516. PMID 17187354.
114.^ Dyall S, Brown M, Johnson P (2004). "Ancient invasions: from endosymbionts to
organelles". Science 304 (5668): 253–7. doi:10.1126/science.1094884. PMID 15073369.
115.^ "Endosymbiosis: Lynn Margulis". University of California Berkely.
http://evolution.berkeley.edu/evolibrary/article/_0_0/history_24. Retrieved 2010-02-20.
116.^ Gould SJ (1997). "The exaptive excellence of spandrels as a term and prototype". Proc.
Natl. Acad. Sci. U.S.A. 94 (20): 10750–5. doi:10.1073/pnas.94.20.10750. PMID 11038582.
PMC 23474. http://www.pubmedcentral.nih.gov/articlerender.fcgi?
tool=pubmed&pubmedid=11038582.
117.^ Gould SJ and Vrba ES (1982). "Exaptation—a missing term in the science of form".
Paleobiology 8 (1): 4–15.
 118.^ True JR, Carroll SB (2002). "Gene co-option in physiological and morphological
evolution". Annu. Rev. Cell Dev. Biol. 18: 53–80. doi:10.1146/annurev.cellbio.18.020402.140619.
PMID 12142278.
119.^ Cañestro C, Yokoi H, Postlethwait JH (2007). "Evolutionary developmental biology and
genomics". Nat Rev Genet 8 (12): 932–942. doi:10.1038/nrg2226. PMID 18007650.
120.^ Baguñà J, Garcia-Fernàndez J (2003). "Evo-Devo: the long and winding road". Int. J. Dev.
Biol. 47 (7–8): 705–13. PMID 14756346. http://www.ijdb.ehu.es/web/paper.php?doi=14756346.
*Gilbert SF (2003). "The morphogenesis of evolutionary developmental biology". Int. J. Dev. Biol. 47
(7–8): 467–77. PMID 14756322.
121.^ West-Eberhard, M-J (2003). Developmental Plasticity and Evolution. Oxford University
Press.
122.^ Newman SA, Müller GB (2000). "Epigenetic mechanisms of character origination". J. Exp.
Zool. B Mol. Develop. Evol. 288 (4): 304–17. doi:10.1002/1097-010X(20001215)288:4<304::AID-
JEZ3>3.0.CO;2-G. PMID 11144279.
123.^ Roberts, Christina. "Epigenetics and Evolution". South Florida University.
http://web.me.com/christinalrichards/Portfolio/Epigenetics.html. Retrieved 2010-02-21.
124.^ Rapp, Ryan; Wendell, Jonathan. "Epigenetics and Plant Evolution". New Phytologist.
http://www3.interscience.wiley.com/journal/118647390/abstract. Retrieved 2010-02-21.
125.^ Singer, Emily (2009). "A Comeback for Lamarckian Evolution?". MIT Technology Review.
http://www.technologyreview.com/biomedicine/22061/. Retrieved 2010-02-21.
126.^ Lovelock J (2003). "Gaia: the living Earth". Nature 426 (6968): 769–70.
doi:10.1038/426769a. PMID 14685210.
127.^ Margulis, Lynn (1995). "Gaia Is a Tough Bitch". The Third Culture.
http://www.edge.org/documents/ThirdCulture/n-Ch.7.html. Retrieved 2007-09-30.
 128.^ Fox, Robin (2004). "Symbiogenesis.". Journal of the royal society of medicine 97 (12):
559. doi:10.1258/jrsm.97.12.559. PMID 15574850. PMC 1079665.
http://jrsm.rsmjournals.com/cgi/content/full/97/12/559.

[edit] References
• Augustine (1982). The Literal Meaning of Genesis. trans. John Hammond Taylor. The
Newman Press. ISBN 0809103265, 9780809103263. http://books.google.com/?id=_s0kIgD0nCcC.
• Bowler, Peter J. (2003). Evolution: The History of an Idea (3rd ed.). University of California
Press. ISBN 0-52023693-9.
• Bowler, Peter J.; Morus, Iwan Rhys (2005). Making Modern Science. The University of
Chicago Press. ISBN 0-226-06861-7.
• Darlington, Cyril (1959). Darwin's place in history. Blackwell, Oxford, p85.
• Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life (1st ed.). John Murray, London.
• Darwin, Charles (1861). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life (3rd ed.). John Murray, London.
• Darwin, Charles (1866). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life (4th ed.). John Murray, London.
• Darwin, Charles (1872). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life (6th ed.). John Murray, London.
• Darwin, Erasmus (1825). The Temple of Nature, or The Origin of Society. Jones & Company.
http://books.google.com/?id=oAl9y-0FSJQC&dq=Erasmus+Darwin+Temple.
 • Darwin, Erasmus (1818). Zoonomia. http://www.gutenberg.org/files/15707/15707-h/15707-
h.htm.
• Desmond, Adrian; Moore, James (1994). Darwin: The Life of a Tormented Evolutionist. W.
W. Norton & Company. ISBN 0393311503.
• Gill, Meredith Jane (2005). Augustine in the Italian Renaissance: Art and Philosophy from
Petrarch to Michelangelo. Cambridge University Press. ISBN 0521832144, ISBN 9780521832144.
http://books.google.com/?id=AksE4jKI8yQC.
• Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Belknap Press of Harvard
University Press. ISBN 0-674-00613-5.
• Henderson, Jan-Andrew (2000). The Emperor's Kilt: The Two Secret Histories of Scotland .
Mainstream Publishing.
• Larson, Edward J. (2004). Evolution: The Remarkable History of a Scientific Theory . Modern
Library. ISBN 0-679-64288-9.
• Lovejoy, Arthur (1936). The Great Chain of Being: A Study of the History of an Idea . Harvard
University Press. ISBN 0-674-36153-9.
• Mayr, Ernst (1982). The Growth of Biological Thought: Diversity, Evolution, and Inheritance .
The Belknap Press of Harvard University Press. ISBN 0-674-36445-7.
• Mayr, Ernst; Provine, W. B., eds (1998). The Evolutionary Synthesis: Perspectives on the
Unification of Biology. Harvard University Press. ISBN 0-674-27225-0.
• Needham, Joseph; Ronan, Colin Alistair (1995). The Shorter Science and Civilisation in
China: An Abridgement of Joseph Needham's Original Text, Vol. 1. Cambridge University Press.
ISBN 0521292867.
 • Sapp, Jan (2003). Genesis: The Evolution of Biology. Oxford University Press. ISBN 0-19-
515618-8.
• Secord, James A. (2000). Victorian Sensation: The Extraordinary Publication, Reception,
and Secret Authorship of Vestiges of the Natural History of Creation . University of Chicago Press.
ISBN 0-226-74410-8.
• Singer, Charles (1931). A Short History of Biology. Clarendon Press.
• Smocovitis, Vassiliki Betty (1996). Unifying Biology: The Evolutionary Synthesis and
Evolutionary Biology. Princeton University Press. ISBN 0-691-03343-9.
• This article incorporates text from a publication now in the public domain: Chisholm,
Hugh, ed (1911). Encyclopædia Britannica (Eleventh ed.). Cambridge University Press.

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 W000

Histoire de la pensée évolutionniste

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

L’Histoire de la pensée évolutionniste raconte la succession des idées qui ont permis de comprendre
le mécanisme de l'évolution des espèces.
Sommaire
[masquer]
• 1 Dans l'Antiquité
• 2 Moyen Âge
• 2.1 Dans le monde musulman
• 3 Les théories au XVIIIe siècle
• 3.1 Le Fixisme: Catastrophisme et Gradualisme
• 3.2 Le Transformisme
• 4 Le XIXe siècle et la révolution darwinienne
• 5 Le XXe siècle: La théorie synthétique de l'évolution
• 5.1 La découverte des mécanismes de l'hérédité
• 5.2 La théorie de la récapitulation
• 5.3 La naissance de la théorie synthétique de
l'évolution
• 5.4 Développements de la théorie depuis les
années 1950
• 5.5 L'étude de l'évolution aujourd'hui
• 6 Notes et références
• 7 Voir aussi
• 7.1 Articles connexes
• 7.2 Bibliographie
Dans l'Antiquité [modifier]
Ce sont chez les philosophes grecs qu'on trouve les plus anciennes traces d'explication de la
diversité biologique que la Terre porte. Le présocratique Anaximandre, élève de Thalès de Milet, concevait
déjà l'homme comme étant précédemment un poisson ou un animal proche de celui-ci. Il avança ainsi l’idée
selon laquelle les hommes avaient dû passer une partie de cette transition dans la bouche de gros poissons
pour se protéger du climat jusqu’à ce qu’ils puissent regagner l’air libre et perdre leurs écailles[1]. La vie vient
de l'eau pour Anaximandre.
Démocrite considère la vie comme un assemblage heureux d'éléments qui a pu perdurer de façon
viable.
Platon inscrit la vie dans deux mondes. Un monde réel, idéal impalpable et un monde illusoire perçu
par les sens. Il a une vision statique de la biocénose que l'on qualifiera de fixiste.
Son disciple Aristote classe le vivant sur une échelle de complexité : la Scala naturæ l'échelle de la
vie. Pour Aristote, l'ensemble du vivant est pourvu d'une psyché (anima en latin). Il existe un continuum
hiérarchisé allant de la plante à l'animal. Capable de sensation, de désir, de mouvement, celui-ci reste
néanmoins inférieur à l'homme, qui possède l'apanage de la pensée. Mais comme Platon, Aristote défend
une vision fixiste du monde vivant.
La culture judéo-chrétienne, notamment dans les récits de l'Ancien Testament, appuie les visions
fixistes des philosophes grecs. Selon le récit de la Genèse, Dieu a créé tout le vivant de la Terre, y compris
l'homme qui est à son image. C'est d'ailleurs pour découvrir cet ordre divin que Carl von Linné le père de la
taxinomie a mis en place la classification binomiale.
Moyen Âge [modifier]
Dans le monde musulman [modifier]
Après la chute de l'Empire romain, les idées évolutionnistes continuèrent à être exposées par les
savants et philosophes musulmans au Moyen Âge durant l'âge d'or de la civilisation islamique, alors que les
théories anciennes de l'évolution et de la sélection naturelle[réf. nécessaire] étaient largement enseignés
dans écoles islamiques[2],[3]. Le savant, historien et philosophe John William Draper a parlé au XIXe siècle
de la théorie mahométane de l'évolution[4].
Le premier naturaliste et philosophe musulman à développer une théorie de l'évolution fut le
zoologiste Al-Jahiz (776-868) au IXe siècle[5],[6]. Dans son Livre des Animaux, il dresse une anthologie
animalière où est évoquée une évolution articulée selon trois mécanismes principaux : la lutte pour
l’existence, la transformation d’espèces vivantes et l’influence de l’environnement naturel[réf. nécessaire]. Il
marque ainsi l’unité de la nature et les rapports entre divers groupes d’êtres vivants.
À sa suite, pendant le Xe siècle, plusieurs penseurs musulmans reprennent ses idées sur l'évolution
des êtres vivants, comme Ali ibn Abbas al-Majusi ou Nasir ad-Din at-Tusi. Selon Sigrid Hunke (1913-1999),
Ali ibn Abbas al-Majusi (?-982 ou 994) a expliqué l'origine des espèces par la voie de la sélection naturelle 9
siècles avant Darwin[7]. D'après Réda Benkirane, cette pensée naturaliste décrivant une évolution globale
impliquant le minéral, le végétal et l’animal se retrouve entre autres chez le philosophe et historien iranien
Ibn Miskawayh (932-1030)[8]
Au Xe siècle, les Frères de la pureté (Ikhwan al-Safa ) décrivent dans une section de l'Épître des
frères de la pureté la création des mondes et l'évolution par strates de la vie avec des détails qui auraient
impressionné Darwin[9]. On y trouve l'idée d'évolution à partir de la matière, laquelle se transforme en
vapeur, puis en eau, en minéraux, en plantes, en animaux, en singes et enfin en hommes[10],[11]. Ainsi les
groupes d’êtres parcourent dans l’engendrement de leurs formes définitives une évolution qui va du simple
au complexe, passant par les quatre éléments (feu, terre, air, eau), les quatre natures (chaud, froid, sec,
humide) et leurs combinaisons poursuivent encore la différenciation en règnes minéral, végétal et animal et
précisent indéfiniment la spéciation du vivant[8].
L'épitre explique comment se déroule la manifestation par couches successives, ou stratifiées à
partir du royaume minéral. Selon lui les entités minérales les plus développées vivent plus bas dans le
royaume minéral jusqu'à ses plus hautes strates pour se mélanger imperceptiblement dans la strate
supérieure du règne végétal. Il explique aussi l'existence de contacts entre les règnes animal et végétal ; et
jusqu'au plus haut niveau du règne animal, dont le point culminant serait l'Homme. Les plus évolués seraient
les hommes placés dans les hautes sphères, debout entre les anges et les animaux, pour servir sur la Terre
comme lieutenants de Dieu.
Par la suite, Nasir ad-Din at-Tusi (1201-1274) suggère la sélection des meilleurs et l'adaptation des
espèces pour l'évolution environ 6 siècles avant Charles Darwin. Il utilise pour expliquer les transformations
des espèces, le mot takâmul, qui signifie en arabe « perfectionnement ». Selon Tusi, ce sont les
transformations de l'environnement qui poussent les espèces à évoluer ; ainsi ce seraient les espèces dont
les individus sont les plus diversifiés en formes qui s'adapteraient le mieux aux changements.
Tusi écrira ainsi : « "...l'équilibre (originel) a été endommagé, et les contrastes essentiels ont commencé à
apparaitre à l'intérieur de ce monde très tôt. Par conséquent, quelques substances ont commencé à se
développer plus rapidement et à s'améliorer plus que les autres." » et encore : « "Les organismes qui
peuvent gagner les nouveaux dispositifs plus rapidement sont plus variables. En conséquence, elles
gagnent des avantages par rapport à d'autres créatures." [12] »
Farid Alakbarov étudie en détail ce domaine dans son livre intitulé : Nasiraddin Tusinin takamul
gorushlari[13].
Enfin, l’historien maghrébin Ibn Khaldoun (1338-1405) recourt aux notions d’ordre, de structure, de
plan, de rapports entre les êtres et des permutations réciproques, de progrès graduel de la création et de
continuum des êtres vivants. Il suggère également la transformation progressive et organisée du minéral
vers le végétal, l'animal, le singe et finalement l'Homme[14].
Il écrit ainsi que : « "le plan humain est atteint à partir du monde des singes ( qirada)."[15] »
Si ces écrits n'ont pas fait condamner leurs auteurs par les autorités islamiques[16], ils n'ont eu
cependant que peu d'écho.

Les théories au XVIIIe siècle [modifier]

Le Fixisme: Catastrophisme et Gradualisme [modifier]
Au XVIIIe siècle, les idées fixistes sont partagées par un grand nombre de scientifiques dont fait
partie Carl von Linné, le père de la taxinomie, qui met en place la nomenclature binomiale des espèces,
toujours en vigueur, afin de mieux comprendre cet ordre divin[17]. Contrairement à la scala naturæ, sa
classification repose sur une hiérarchie de catégories de plus en plus précises regroupant les espèces
semblables, mais Linné n'établit aucun lien de parenté entre celles-ci et effectue plutôt un lien avec leur plan
de création[18]. En s'inspirant des travaux d'anatomie de Nicolaes Tulp et surtout d'Edward Tyson, Linné
constate que le chimpanzé présente plus de caractères communs avec l'homme qu'avec les autres singes et
amène ainsi l'homme et le chimpanzé a cohabiter pour la première fois dans l'ordre des Primates en 1758
[[19]].
 L'étude des fossiles a également permis d'alimenter l'idée d'évolution[18],[20]. Georges Cuvier,
fondateur de la paléontologie, remarque que plus une strate est profonde, plus les fossiles en son sein se
démarquent des espèces actuelles et que certaines espèces apparaissent tandis que d'autres disparaissent.
Fixiste et farouche opposant au transformisme de Lamarck, Cuvier devient le principal partisan du
catastrophisme, théorie selon laquelle il y aurait eu plusieurs créations entrecoupées de catastrophes
planétaires et qui permet de concilier la présence de fossiles d'espèces éteintes avec les récits bibliques[21].
À l'époque, cette thèse s'oppose aux idées défendues par la thèse du gradualisme, établie en 1795 par
James Hutton, puis par son complément l'uniformitarisme de Charles Lyell, qui postulent au contraire que les
processus géologiques qui se sont exercés dans le passé lointain sont lents et graduels et s'exercent encore
actuellement de la même façon et à la même vitesse[22]. Selon ce principe, les changements géologiques
se sont déroulés sur une bien plus longue période que les 6000 ans accordées à la Terre par les
théologiens. Le principe du gradualisme sur l'évolution géologique sera ensuite repris par les évolutionnistes,
dont Darwin, pour expliquer l'évolution biologique.

Le Transformisme [modifier]
Article détaillé : Transformisme.
Un certain nombre de penseurs avaient affirmé ou suggéré l'idée d'une évolution des espèces, par
exemple Jérôme Cardan dans De Subtilitate (1550)[23], ou Giambattista della Porta dans son livre De Magia
Naturale[23] qui inspira Francis Bacon[23]. Selon certains auteurs, dont Kohlbrugge, Jean Bodin aurait été
l'un des premiers « évolutionnistes », avec son livre Heptaplomeres, qui ne fut publié qu'en 1841 en raison
de son caractère scandaleux[23]. D'autres (dont Franck Bourdier) considèrent qu'il n'y a évoqué, tout au
plus, que la génération spontanée[23]. Lucilio Vanini, qui fut brûlé vif, reprit quelques thèses de Cardan[23].
On peut encore citer Paolo Sarpi[23], ou encore, de façon plus étrange, aussi bien Cyrano de Bergerac que
le jésuite Athanase Kircher[23].
Si l'idée d'évolution commence à s'affirmer au milieu du XVIIIe siècle[réf. nécessaire] avec
Maupertuis (Essai sur la transformation des corps organisés, 1754[23]) et Buffon, la première théorie
scientifique rendant compte de l'évolution des espèces dans le temps est attribué au naturaliste français
Jean-Baptiste Lamarck. La publication, en 1809, dans Philosophie zoologique, de sa théorie transformiste
tournant résolument le dos au fixisme entraîne de virulents débats devant l'Académie des sciences.
Dans cet ouvrage, complété et corrigé par l'introduction de son Histoire naturelle des animaux sans
vertèbres en 1815, Lamarck tente de comprendre les êtres vivants en tant que phénomènes physiques.
Cette démarche matérialiste et mécaniste s'oppose au vitalisme encore dominant à l'époque. Il expose une
théorie des êtres vivants expliquant l'évolution des espèces à l'aide de deux tendances opposées:
• La complexification croissante de l'organisation des êtres vivants sous l'effet de la
dynamique interne propre à leur métabolisme ; (a compléter)
• La diversification, ou spécialisation, des êtres vivants sous l'effet des circonstances variées
auxquelles ils sont confrontés et auxquelles ils s'adaptent en modifiant leur habitudes de manière à
répondre à leurs besoins. C'est le principe de l'usage et du non-usage, qui veut qu'un organe se
développe ou s'atrophie selon son degré d'utilisation.
Lamarck, comme nombre de ses prédécesseurs depuis Aristote, admet la transmission à la
descendance de caractères acquis au cours de la vie d'un individu. Il est le premier évolutionniste moderne
et on lui doit une des premières formulations des relations de parenté entre les grands groupes
d'organismes, notamment les invertébrés.
 Malgré de nombreuses critiques de la part des milieux religieux et scientifique, les idées
transformistes reçoivent une adhésion croissante à partir de 1825 et ont permis de rendre le débat
naturaliste plus réceptif aux théories évolutionnistes [24].

Le XIXe siècle et la révolution darwinienne [modifier]

En 1859, Charles Darwin, naturaliste anglais, publie De l'origine des espèces. Il y reprend les idées
de Lamarck tout en les critiquant et en les modifiant. Darwin ajoute surtout une foule de preuves en faveur de
l'idée d'évolution (par transformation graduelle) et propose pour la première fois le mécanisme de la
sélection naturelle ; mais il n'y remet pas en cause l'idée de l'hérédité des caractères acquis, citant même
dans L'Origine des espèces les « effets cumulatifs du dressage » sur une lignée de chiens pointers.
Darwin propose d'ailleurs un modèle pour la transmission des caractères acquis sous le nom
« d’hypothèse de la pangenèse » dans Les variations des animaux et des plantes sous l’effet de la
domestication (1868). Son modèle ressemble à celui qu’avait proposé Maupertuis dans sa Vénus physique
(1745) et son Système de la Nature, hormis l’utilisation de la récente théorie cellulaire ; sans reconnaître
pour autant l’origine de ses idées. Darwin introduit aussi l'idée d'une sélection sexuelle qui permet
notamment d'expliquer les cas étonnant de dimorphisme sexuel comme les couleurs chatoyantes des mâles
dans certaines espèces. Enfin, Darwin insiste sur le fait que l'espèce humaine est elle aussi le produit de
l'évolution par sélection naturelle et sexuelle dans son ouvrage La Descendance de l'homme et la sélection
sexuelle.

Le XXe siècle: La théorie synthétique de l'évolution [modifier]

Article détaillé : Théorie synthétique de l'évolution.
La découverte des mécanismes de l'hérédité [modifier]
À la fin du XIXe siècle, le moine autrichien Gregor Mendel découvre les lois de la génétique avec ses
expériences sur les pois. En proposant un mécanisme pour l'hérédité, c'est-à-dire la transmission (au moins
partielle) des caractères d'un individu à ses descendants. Toutefois, ses travaux ne seront pas diffusés et les
découvertes de Mendel seront oubliées jusqu'à ce que des biologistes (Hugo de Vries, Carl Correns et Erich
von Tschermak) au début du XXe siècle, les redécouvrent. Les lois de l'hérédité mendélienne apportent alors
un soutien crucial aux idées de Darwin.

La théorie de la récapitulation [modifier]

Selon la théorie de la récapitulation, dont la formulation est due à Ernst Haeckel (début du
XXe siècle), chaque être vivant, au cours de son développement, récapitulerait l'histoire évolutive de son
groupe. La « série des embryons » et la « série des ancêtres », qui aboutissent toutes deux au même
individu, devraient être identiques. Sans être totalement fausse, cette théorie n'est que partiellement exacte
et n'est justifiée par aucun mécanisme évolutif reconnu. Au contraire même, la néoténie (arrêt prématuré du
développement maintenant des caractéristiques juvéniles chez l'adulte) est un processus évolutif fréquent,
que l'on retrouve jusque chez les primates ; on admet par exemple que l'être humain ressemble plus à un
jeune chimpanzé qu'à un singe adulte. Aldous Huxley a utilisé cette observation dans son roman Jouvence.
En 1910, le biologiste de Vries découvre les mutations génétiques. Ces modifications aléatoires du
code génétique permettent d'expliquer la variabilité naturelle des caractères individuels, terrain sur lequel se
fait la sélection darwinienne. Tout au long du XXe siècle, la génétique émergente viendra étayer les idées de
Darwin. Dans les années 1930, des biologistes comme Thomas Hunt Morgan font beaucoup progresser la
génétique, notamment grâce à l'étude des chromosomes de la mouche drosophile.
 Ces mêmes années, le statisticien et généticien Ronald Fisher apporte d'importants développement
mathématiques à la théorie de l'évolution en lien avec les mécanismes de sélection naturelle mais surtout
sexuelle fournissant par exemple une explication au fait que les membres des deux sexes, mâles et
femelles, sont aussi nombreux dans la plupart des espèces. Les résultats de Fisher donneront naissance à
ce qu'on appellera la génétique des populations.

La naissance de la théorie synthétique de l'évolution [modifier]

Dans les années 1940, quelques pionniers (Theodosius Dobzhansky, Ernst Mayr, George Gaylord
Simpson et Julian Huxley) fondent la théorie synthétique de l'évolution (TSE). Comme son nom l'indique,
celle-ci est destinée à synthétiser et à englober dans une vision d'ensemble les données accumulées
séparément par la génétique, la biologie et la paléontologie.
Simultanément, un embryologiste autrichien, Richard Goldschmidt, propose sa théorie du monstre
prometteur.

Développements de la théorie depuis les années 1950 [modifier]

Au cours des années 1960, William Hamilton poursuivra les travaux de Fisher. Il contribuera par
exemple à la théorie de la reine rouge expliquant l'avantage de la reproduction sexuée en ce qu'elle permet
de résister de façon plus efficace aux parasites.
En 1975, le biologiste Amotz Zahavi formulera le principe du handicap qui permet d'expliquer
certaines formes impressionnantes de dimorphisme sexuel. Ce principe explique le fait que dans certaines
espèces les mâles (en général) présentent des caractères exubérants car ce faisant ils montrent qu'ils sont
capable de survivre malgré le handicap qu'inflige une telle exubérance (ils sont par exemple plus visibles des
prédateurs). Les femelles ont donc intérêt à choisir pour leur progéniture un père qui a le handicap le plus
fort et qui est donc le plus capable de dépasser ce handicap.
Au cours des années 1980, la théorie darwinienne se verra développée sous l'angle mathématique
et statistique par des biologistes comme John Maynard Smith. Ce dernier s'inspirant de la théorie des jeux
développée par le mathématicien John Nash, introduira le concept de stratégie évolutionnairement stable qui
désigne le fait que lorsque plusieurs populations sont en compétition, celles qui dominent les autres vont se
répandre tandis que celles qui sont dominées vont disparaitre.
Maynard Smith collaborera aussi avec le biochimiste Eörs Szathmáry dans un travail de synthèse
sur les transitions majeurs dans l'histoire de la vie intitulé comme l'apparition du code génétique, des
eucaryotes, des êtres multicellulaires, ou encore l'apparition des sociétés humaines.

L'étude de l'évolution aujourd'hui [modifier]

Enfin, en privilégiant l'approche simultanée des organismes, certains auteurs comme Leigh Van
Valen, William Rice ou encore en France le biologiste Thierry Lodé[25] insistent sur les processus de
coévolution et réfutent certains points de la théorie néodarwinienne[26]. Dans sa théorie de la reine rouge,
Van Valen postule que l'évolution résulte des interactions entre organismes et se poursuit perpétuellement,
tandis que les théories du conflit de Rice et Lodé proposent que l'évolution résulte principalement des
interactions antagonistes, et notamment du conflit sexuel.
Notes et références [modifier]
1. ↑ Plutarque mentionne également cette théorie d’Anaximandre où les humains naissaient à
l’intérieur des poissons, se nourrissant tels des requins, et que lorsqu’ils étaient en mesure de se
défendre, ils étaient rejetés sur le rivage pour vivre sur la terre ferme.
2. ↑ Selon Réda Benkirane, en islam, l’évolution et la contingence sont inscrites au cœur
même de la révélation coranique. Ainsi selon la tradition islamique (Sunna), la raison coranique telle
qu’elle s’est elle-même révélée à l’être adamique est donc éminemment évolutive et non linéaire. Par
ailleurs, même en partant d’une création divine du monde, la conception islamique où Dieu travaille
continuellement à sa création, souligne que l’évolution biologique ne crée pas d’impasse
métaphysique particulière aux musulmans. Le Créateur y reste incommensurable, inconnaissable,
quand « Il n’engendre pas et n’est pas engendré » et que « Nul n’est égal à Lui. » (Coran : 112, 1-4).
La notion théologique de renouvellement de la création (tajdid al-khalq) est ici spécifique de la
tradition islamique. C’est parce que « tout ce qui est sur terre est périssable » que le processus de
création est en fait une re-création permanente (« Chaque jour, Il est à la tâche », (Cor. LV : 29).
Création divine et évolution biologique ne s’excluent donc pas l’une l’autre.
3. ↑ Plusieurs savants soutiennent une interprétation moderne du Coran à ce sujet dont :
Sayyid Qutb (1906-1966) dans Fî Dhilâl'il Qur'ân, Elmalılı Muhammed Hamdi Yazır (1877-1942) dans
son exégèse du Coran intitulé Hak dîni Kur'ân dili et le Dr. Maurice Bucaille (1920-?)...dans son livre
intitulé L’homme, d’où vient-il ? La réponse de la science et des écritures saintes. Qui sont accusés
par les occidentaux de concordisme.
4. ↑ (en) John William Draper, History of the Conflict between Religion and
Science [archive], D. Appleton and Co., New York, 1881, 8e éd., 373 p., p. 188
 5. ↑ Conway Zirkle (1941). Natural Selection before the "Origin of Species", Proceedings of the
American Philosophical Society 84 (1), p. 71-123.
6. ↑ Mehmet Bayrakdar (Third Quarter, 1983). "Al-Jahiz And the Rise of Biological
Evolutionism", The Islamic Quarterly. Londres. [1] [archive]
7. ↑ [2] [archive]Le Soleil d'Allah brille sur l'Occident, Espace Libre, éd. Albin Michel, 1997,
p.158.
8. ↑ a et b Réda Benkirane, Créationnisme en islam, une aberration, La Recherche n⁰ 417, mars
2008 texte en ligne [archive]
9. ↑ [3] [archive] Source material for this article includes: A History of Islamic Philosophy by
Majid Fakhry, 1970; Arabic Thought and Its Place in History by DeLacy O'Leary, 1922; and by the
same author, How Greek Science Passed to the Arabs, 1948. (From Sunrise magazine, April & May,
1973. Copyright © 1973 by Theosophical University Press)
10.↑ Muhammad Hamidullah and Afzal Iqbal (1993), The Emergence of Islam: Lectures on the
Development of Islamic World-view, Intellectual Tradition and Polity , p. 143-144. Islamic Research
Institute, Islamabad.
11.↑ Eloise Hart, Pages of Medieval Mideastern History. (cf. Isma'ili, Yezidi, Sufi, The Brethren
Of Purity [archive], Ismaili Heritage Society)
12.↑ Akhlag Nasiri, ou Nasirean Ethics traduit vers l'anglais par G.M. Wickens et édité par
George Allen & Unwin en 1964.
13.↑ [4] [archive] : [Dr. Farid Alakbarov first became interested in Tusi's views on evolution in
1986, when he began his research of Arabic texts at Baku's Institute of Manuscripts. In 2000, he
wrote a booklet entitled (Azeri) (The Evolutionary Views by Nasiraddin Tusi), known as
"Evolutsionniye vzglyadi Nasiraddina Tusi" in Russian. He also presented a paper on the subject at
Azerbaijan International University's Conference Devoted to the 800th Jubilee of Nasiraddin Tusi. Dr.
Alakbarov writes a regular feature column in AI related to medieval medicine. His previous articles
are available at AZER.com in English; or at AZERI.org, in the Azeri Latin script.
14.↑ (fr) Ibn Khaldoun, Les Prolégomènes, éd. Institut de France, Paris, 1863, p.
229 [archive]
15.↑ (fr) Ibn Khaldoun, Les Prolégomènes, éd. Institut de France, Paris, 1863, p.
229 [archive]
16.↑ Plusieurs savants soutiennent une interprétation moderne du Coran à ce sujet dont :
Sayyid Qutb (1906-1966) dans Fî Dhilâl'il Qur'ân, Elmalılı Muhammed Hamdi Yazır (1877-1942) dans
son exégèse du Coran intitulé Hak dîni Kur'ân dili et le Dr. Maurice Bucaille (1920-?). Qui sont
accusés par les occidentaux de concordisme.
17.↑ [pdf] Thierry Hoquet, Les fondements de la botanique, Linné et la classification des
plantes [archive], Revue Le Banquet, n°23, 2006.
18.↑ a et b Neil Campbell & al., Biologie, éd. De Boeck, 2007, p. 476.
19.↑ Pascal Picq, Lucy et l'obscurantisme, éd. Odile Jacob, 2007.
20.↑ Laurent Dubois, [pdf] Histoire de la paléontologie, Darwin et Théorie de
l'Évolution [archive], Géopolis.fr, (page consultée le 4 juillet 2008).
21.↑ Serge Lapierre, Éléments de théorie de l'évolution [archive], Collège de Bois de Boulogne
- Département de philosophie, (page consultée le 5 juillet 2008).
22.↑ Une brève histoire de la géologie [archive], Université de Liège, mise à jour le 1 novembre
2007 (page consultée le 7 juillet 2008).
23.↑ a, b, c, d, e, f, g, h et i Bourdier Franck. « Trois siècles d'hypothèses sur l'origine et la
transformation des êtres vivants (1550-1859) [archive]. ». In: Revue d'histoire des sciences et de
leurs applications. 1960, Tome 13 n°1. Lamarck et Darwin. À l'occasion du Centenaire de "L'Origine
des espèces". pp. 1-44.
 24.↑ Hélène Blais, « Lamarck, genèse et enjeux du transformisme, 1770-1830 », La Revue
pour l’histoire du CNRS [lire en ligne [archive]], n°7 - novembre 2002, mis en ligne le 6 mars 2006.
Consulté le 8 juin 2010.
25.↑ Thierry Lodé "La guerre des sexes chez les animaux, une histoire naturelle de la sexualité"
2006, Eds Odile Jacob, Paris ISBN 2-7381-1901-8
26.↑ J Lassègue, 2009, Compte-Rendu CNRS École polytechnique [5] [archive]

Voir aussi [modifier]

Articles connexes [modifier]
• La Planète des singes
• Jean-Baptiste de Lamarck

Bibliographie [modifier]
• Jean Chaline, Quoi de neuf depuis Darwin ? (2006), Ellipses, Paris. (Historique des théories
de l'évolution, présentation de l'évo-dévo)

Liens externes [modifier]

• (fr) Evolution, de l'origine de la vie aux origines de l'homme, site CNRS/sagascience

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Modern evolutionary synthesis

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The modern evolutionary synthesis is also referred to as the new synthesis, the modern synthesis,
the evolutionary synthesis and the neo-darwinian synthesis. It is a union of ideas from several biological
specialties which provides a widely accepted account of evolution. The synthesis has been accepted by
nearly all working biologists.[1] The synthesis was produced over a decade (1936–1947). The previous
development of population genetics (1918–1932) was a stimulus, as it showed that Mendelian genetics was
consistent with natural selection and gradual evolution. The synthesis is still, to a large extent, the current
paradigm in evolutionary biology.[2]
Julian Huxley invented the term, when he produced his book, Evolution: The Modern Synthesis
(1942). Other major figures in the modern synthesis include R. A. Fisher, Theodosius Dobzhansky, J.B.S.
Haldane, Sewall Wright, E.B. Ford, Ernst Mayr, Bernhard Rensch, Sergei Chetverikov, George Gaylord
Simpson, and G. Ledyard Stebbins.
The modern synthesis solved difficulties and confusions caused by the specialisation and poor
communication between biologists in the early years of the 20th century. Discoveries of early geneticists
were difficult to reconcile with gradual evolution and the mechanism of natural selection. The synthesis
reconciled the two schools of thought, while providing evidence that studies of populations in the field were
crucial to evolutionary theory. It drew together ideas from several branches of biology that had become
separated, particularly genetics, cytology, systematics, botany, morphology, ecology and paleontology.
Contents
[hide]
• 1 Summary of the modern synthesis
• 2 Developments leading up to the synthesis
• 2.1 1859–1899
• 2.2 1900–1915
• 2.3 The foundation of population genetics
• 3 The modern synthesis
• 4 Further advances
• 5 After the synthesis
• 5.1 Understanding of Earth history
• 5.2 Symbiotic origin of eukaryotic cell structures
• 5.3 Trees of life
• 5.4 Evo-devo
• 5.5 Fossil discoveries
• 6 See also
• 7 Footnotes
• 8 References
• 9 External links
[edit] Summary of the modern synthesis
The modern synthesis bridged the gap between experimental geneticists and naturalists, and
between both and palaeontologists. It states that:[3][4][5]
1. All evolutionary phenomena can be explained in a way consistent with known genetic
mechanisms and the observational evidence of naturalists.
2. Evolution is gradual: small genetic changes, recombination ordered by natural selection.
Discontinuities amongst species (or other taxa) are explained as originating gradually through
geographical separation and extinction (not saltation).
3. Natural selection is by far the main mechanism of change; even slight advantages are
important when continued. The object of selection is the phenotype in its surrounding environment.
4. The role of genetic drift is equivocal. Though strongly supported initially by Dobzhansky, it
was downgraded later as results from ecological genetics were obtained.
5. Thinking in terms of populations, rather than individuals, is primary: the genetic diversity
existing in natural populations is a key factor in evolution. The strength of natural selection in the wild
is greater than previously expected; the effect of ecological factors such as niche occupation and the
significance of barriers to gene flow are all important.
6. In palaeontology, the ability to explain historical observations by extrapolation from
microevolution to macroevolution is proposed. Historical contingency means explanations at different
levels may exist. Gradualism does not mean constant rate of change.
The idea that speciation occurs after populations are reproductively isolated has been much debated.
In plants, polyploidy must be included in any view of speciation. Formulations such as 'evolution consists
primarily of changes in the frequencies of alleles between one generation and another' were proposed rather
later. The traditional view is that developmental biology ('evo-devo') played little part in the synthesis,[6] but
an account of Gavin de Beer's work by Stephen J. Gould suggests he may be an exception.[7]

[edit] Developments leading up to the synthesis

See also: History of evolutionary thought

[edit] 1859–1899
Charles Darwin's The Origin of Species was successful in convincing most biologists that evolution
had occurred, but was less successful in convincing them that natural selection was its primary mechanism.
In the 19th and early 20th centuries, variations of Lamarckism, orthogenesis ('progressive' evolution), and
saltationism (evolution by jumps) were discussed as alternatives.[8] Also, Darwin did not offer a precise
explanation of how new species arise. As part of the disagreement about whether natural selection alone
was sufficient to explain speciation, George Romanes coined the term neo-Darwinism to refer to the version
of evolution advocated by Alfred Russel Wallace and August Weismann with its heavy dependence on
natural selection.[9] Weismann and Wallace rejected the Lamarckian idea of inheritance of acquired
characteristics, something that Darwin had not ruled out.[10]
Weismann's idea was that the relationship between the hereditary material, which he called the germ
plasm (de: Keimplasma), and the rest of the body (the soma) was a one-way relationship: the germ-plasm
formed the body, but the body did not influence the germ-plasm, except indirectly in its participation in a
population subject to natural selection. Weismann was translated into English, and though he was influential,
it took many years for the full significance of his work to be appreciated.[11] Later, after the completion of the
modern synthesis, the term neo-Darwinism came to be associated with its core concept: evolution, driven by
natural selection acting on variation produced by genetic mutation, and genetic recombination (chromosomal
crossovers).[9]

[edit] 1900–1915
Gregor Mendel's work was re-discovered by Hugo de Vries and Carl Correns in 1900. News of this
reached William Bateson in England, who reported on the paper during a presentation to the Royal
Horticultural Society in May 1900.[12] It showed that the contributions of each parent retained their integrity
rather than blending with the contribution of the other parent. This reinforced a division of thought, which was
already present in the 1890s.[13] The two schools were:
• Saltationism (large mutations or jumps), favored by early Mendelians who viewed hard
inheritance as incompatible with natural selection[14]
• Biometric school: led by Karl Pearson and Walter Weldon, argued vigorously against it,
saying that empirical evidence indicated that variation was continuous in most organisms, not
discrete as Mendelism predicted.
The relevance of Mendelism to evolution was unclear and hotly debated, especially by Bateson, who
opposed the biometric ideas of his former teacher Weldon. Many scientists believed the two theories
substantially contradicted each other.[15] This debate between the biometricians and the Mendelians
continued for some 20 years and was only solved by the development of population genetics.
T. H. Morgan began his career in genetics as a saltationist, and started out trying to demonstrate that
mutations could produce new species in fruit flies. However, the experimental work at his lab with Drosophila
melanogaster, which helped establish the link between Mendelian genetics and the chromosomal theory of
inheritance, demonstrated that rather than creating new species in a single step, mutations increased the
genetic variation in the population.[16]

[edit] The foundation of population genetics

The first step towards the synthesis was the development of population genetics. R.A. Fisher, J.B.S.
Haldane, and Sewall Wright provided critical contributions. In 1918, Fisher produced the paper "The
Correlation Between Relatives on the Supposition of Mendelian Inheritance",[17] which showed how the
continuous variation measured by the biometricians could be the result of the action of many discrete genetic
loci. In this and subsequent papers culminating in his 1930 book The Genetical Theory of Natural Selection,
Fisher was able to show how Mendelian genetics was, contrary to the thinking of many early geneticists,
completely consistent with the idea of evolution driven by natural selection.[18] During the 1920s, a series of
papers by J.B.S. Haldane applied mathematical analysis to real world examples of natural selection such as
the evolution of industrial melanism in peppered moths.[18] Haldane established that natural selection could
work in the real world at a faster rate than even Fisher had assumed.[19]
Sewall Wright focused on combinations of genes that interacted as complexes, and the effects of
inbreeding on small relatively isolated populations, which could exhibit genetic drift. In a 1932 paper he
introduced the concept of an adaptive landscape in which phenomena such as cross breeding and genetic
drift in small populations could push them away from adaptive peaks, which would in turn allow natural
selection to push them towards new adaptive peaks.[18] Wright's model would appeal to field naturalists
such as Theodosius Dobzhansky and Ernst Mayr who were becoming aware of the importance of
geographical isolation in real world populations.[19] The work of Fisher, Haldane and Wright founded the
discipline of population genetics. This is the precursor of the modern synthesis, which is an even broader
coalition of ideas.[18][19][20]
[edit] The modern synthesis
Theodosius Dobzhansky, an Ukrainian emigrant, who had been a postdoctoral worker in Morgan's
fruit fly lab, was one of the first to apply genetics to natural populations. He worked mostly with Drosophila
pseudoobscura. He says pointedly: "Russia has a variety of climates from the Arctic to sub-tropical...
Exclusively laboratory workers who neither possess nor wish to have any knowledge of living beings in
nature were and are in a minority".[21] Not surprisingly, there were other Russian geneticists with similar
ideas, though for some time their work was known to only a few in the West. His 1937 work Genetics and the
Origin of Species was a key step in bridging the gap between population geneticists and field naturalists. It
presented the conclusions reached by Fisher, Haldane, and especially Wright in their highly mathematical
papers in a form that was easily accessible to others. It also emphasized that real world populations had far
more genetic variability than the early population geneticists had assumed in their models, and that
genetically distinct sub-populations were important. Dobzhansky argued that natural selection worked to
maintain genetic diversity as well as driving change. Dobzhansky had been influenced by his exposure in the
1920s to the work of a Russian geneticist named Sergei Chetverikov who had looked at the role of recessive
genes in maintaining a reservoir of genetic variability in a population before his work was shut down by the
rise of Lysenkoism in the Soviet Union.[18][19]
Edmund Brisco Ford's work complemented that of Dobzhansky. It was as a result of Ford's work, as
well as his own, that Dobzhansky changed the emphasis in the third edition of his famous text from drift to
selection.[22] Ford was an experimental naturalist who wanted to test natural selection in nature. He virtually
invented the field of research known as ecological genetics. His work on natural selection in wild populations
of butterflies and moths was the first to show that predictions made by R.A. Fisher were correct. He was the
first to describe and define genetic polymorphism, and to predict that human blood group polymorphisms
might be maintained in the population by providing some protection against disease.[23]
 Ernst Mayr's key contribution to the synthesis was Systematics and the Origin of Species, published
in 1942. Mayr emphasized the importance of allopatric speciation, where geographically isolated sub-
populations diverge so far that reproductive isolation occurs. He was sceptical of the reality of sympatric
speciation believing that geographical isolation was a prerequisite for building up intrinsic (reproductive)
isolating mechanisms. Mayr also introduced the biological species concept that defined a species as a group
of interbreeding or potentially interbreeding populations that were reproductively isolated from all other
populations.[18][19][24] Before he left Germany for the United States in 1930, Mayr had been influenced by
the work of German biologist Bernhard Rensch. In the 1920s Rensch, who like Mayr did field work in
Indonesia, analyzed the geographic distribution of polytypic species and complexes of closely related
species paying particular attention to how variations between different populations correlated with local
environmental factors such as differences in climate. In 1947, Rensch published Neuere Probleme der
Abstammungslehre: die Transspezifische Evolution (English translation 1959: Evolution above the Species
level). This looked at how the same evolutionary mechanisms involved in speciation might be extended to
explain the origins of the differences between the higher level taxa. His writings contributed to the rapid
acceptance of the synthesis in Germany.[25][26]
George Gaylord Simpson was responsible for showing that the modern synthesis was compatible
with paleontology in his book Tempo and Mode in Evolution published in 1944. Simpson's work was crucial
because so many paleontologists had disagreed, in some cases vigorously, with the idea that natural
selection was the main mechanism of evolution. It showed that the trends of linear progression (in for
example the evolution of the horse) that earlier paleontologists had used as support for neo-Lamarckism and
orthogenesis did not hold up under careful examination. Instead the fossil record was consistent with the
irregular, branching, and non-directional pattern predicted by the modern synthesis.[18][19]
 The botanist G. Ledyard Stebbins was another major contributor to the synthesis. His major work,
Variation and Evolution in Plants, was published in 1950. It extended the synthesis to encompass botany
including the important effects of hybridization and polyploidy in some kinds of plants.[18]

[edit] Further advances

The modern evolutionary synthesis continued to be developed and refined after the initial
establishment in the 1930s and 1940s. The work of W. D. Hamilton, George C. Williams, John Maynard
Smith and others led to the development of a gene-centric view of evolution in the 1960s. The synthesis as it
exists now has extended the scope of the Darwinian idea of natural selection to include subsequent scientific
discoveries and concepts unknown to Darwin, such as DNA and genetics, which allow rigorous, in many
cases mathematical, analyses of phenomena such as kin selection, altruism, and speciation.
A particular interpretation most commonly associated with Richard Dawkins, author of The Selfish
Gene, asserts that the gene is the only true unit of selection.[27] Dawkins further extended the Darwinian
idea to include non-biological systems exhibiting the same type of selective behavior of the 'fittest' such as
memes in culture. The synthesis continues to undergo regular review.[28] (See also Current research in
evolutionary biology).
[edit] After the synthesis

The structure of evolutionary biology.

The history and causes of evolution (center) are subject to various subdisciplines of evolutionary biology. The
areas of segments give an impression of the contributions of subdisciplines to the literature of evolutionary
biology.
 There are a number of discoveries in earth sciences and biology which have arisen since the
synthesis. Listed here are some of those topics which are relevant to the evolutionary synthesis, and which
seem soundly based.

[edit] Understanding of Earth history

The Earth is the stage on which the evolutionary play is performed. Darwin studied evolution in the
context of Charles Lyell's geology, but our present understanding of Earth history includes some critical
advances made during the last half-century.
• The age of the Earth has been revised upwards. It is now estimated at 4.56 billion years,
about one-third of the age of the universe. It is worth noting that the Phanerozoic only occupies the
last 1/9th of this period of time.[29]
• The triumph of Alfred Wegener's idea of continental drift came around 1960. The key
principle of plate tectonics is that the lithosphere exists as separate and distinct tectonic plates, which
ride on the fluid-like (visco-elastic solid) asthenosphere. This discovery provides a unifying theory for
geology, linking phenomena such as volcanos, earthquakes, orogeny, and providing data for many
paleogeographical questions.[30] One major question is still unclear: when did plate tectonics begin?
[31]
• Our understanding of the evolution of the Earth's atmosphere has progressed. The
substitution of oxygen for carbon dioxide in the atmosphere, which occurred in the Proterozoic,
caused probably by cyanobacteria in the form of stromatolites, caused changes leading to the
evolution of aerobic organisms.[32][33]
 • The identification of the first generally accepted fossils of microbial life was made by
geologists. These rocks have been dated as about 3.465 billion years ago.[34] Walcott was the first
geologist to identify pre-Cambrian fossil bacteria from microscopic examination of thin rock slices. He
also thought stromatolites were organic in origin. His ideas were not accepted at the time, but may
now be appreciated as great discoveries.[35]
• Information about paleoclimates is increasingly available, and being used in paleontology.
One example: the discovery of massive ice ages in the Proterozoic, following the great reduction of
CO2 in the atmosphere. These ice ages were immensely long, and led to a crash in microflora.[36]
See also Cryogenian period and Snowball Earth.
• Catastrophism and mass extinctions. A partial reintegration of catastrophism has occurred,
[37] and the importance of mass extinctions in large-scale evolution is now apparent. Extinction
events disturb relationships between many forms of life and may remove dominant forms and release
a flow of adaptive radiation amongst groups that remain. Causes include meteorite strikes (K–T
junction; Upper Devonian); flood basalt provinces (Deccan traps at K/T junction; Siberian traps at P–T
junction); and other less dramatic processes.[38][39]
Conclusion: Our present knowledge of earth history strongly suggests that large-scale geophysical
events influenced macroevolution and megaevolution. These terms refer to evolution above the species
level, including such events as mass extinctions, adaptive radiation, and the major transitions in evolution.
[40][41]
 [edit] Symbiotic origin of eukaryotic cell structures
Further information: Endosymbiont and Endosymbiotic theory
Once symbiosis was discovered in lichen and in plant roots (rhizobia in root nodules) in the 19th
century, the idea arose that the process might have occurred more widely, and might be important in
evolution. Anton de Bary invented the concept of symbiosis;[42] several Russian biologists promoted the
idea;[43] Edwin Wilson mentioned it in his epic text The Cell;[44] a book was published in the U.S.A. in 1927
with the title Symbionticism and the origin of species;[45] and there was a brief mention by Julian Huxley in
1930;[46] all in vain because sufficient evidence was lacking. Symbiosis as a major evolutionary force was
not discussed at all in the evolutionary synthesis.[47]
The role of symbiosis in cell evolution was revived partly by Joshua Lederberg,[48] and finally
brought to light by Lynn Margulis in a series of papers and books.[49][50] It turns out that some cell
organelles are of microbial origin: mitochondria and chloroplasts definitely, cilia, flagella and centrioles
possibly, and perhaps the nuclear membrane and much of the chromosome structure as well. What is now
clear is that the evolution of eukaryote cells is either caused by, or at least profoundly influenced by,
symbiosis with bacterial and archaean cells in the Proterozoic.
The origin of the eukaryote cell by symbiosis in several stages was not part of the evolutionary
synthesis. It is, at least on first sight, an example of megaevolution by big jumps. However, what symbiosis
provided was a copious supply of heritable variation from microorganisms, which was fine-tuned over a long
period to produce the cell structure we see today. This part of the process is consistent with evolution by
natural selection.[51]
 [edit] Trees of life
Further information: Last universal ancestor and Phylogenetic tree
The ability to analyse sequence in macromolecules (protein, DNA, RNA) provides evidence of
descent, and permits us to work out genealogical trees covering the whole of life, since now there are data on
every major group of living organisms. This project, begun in a tentative way in the 1960s, has become a
search for the universal tree or the universal ancestor, a phrase of Carl Woese.[52][53] The tree that results
has some unusual features, especially in its roots. There are two kingdoms of prokaryotes: bacteria and
archaea, both of which contributed genetic material to the eukaryotes, mainly by means of symbiosis. Also,
since bacteria can pass genetic material to other bacteria, their relationships look more like a web than a
tree. Once eukaryotes were established, their sexual reproduction produced the traditional branching tree-
like pattern, the only diagram Darwin put in the Origin. The last universal ancestor (LUA) would be a
prokaryotic cell before the split between the bacteria and archaea. LUA is defined as most recent organism
from which all organisms now living on Earth descend (some 3.5 to 3.8 billion years ago, in the Archean era).
[54][55]
This technique may be used to clarify relationships within any group of related organisms. It is now a
standard procedure, and examples are published regularly. April 2009 sees the publication of a tree covering
all the animal phyla, derived from sequences from 150 genes in 77 taxa.[56]
Early attempts to identify relationships between major groups were made in the 19th century by Ernst
Haeckel, and by comparative anatomists such as Thomas Henry Huxley and E. Ray Lankester. Enthusiasm
waned: it was often difficult to find evidence to adjudicate between different opinions. Perhaps for that
reason, the evolutionary synthesis paid surprisingly little attention to this activity. It is certainly a lively field of
research today.
 [edit] Evo-devo
Further information: Evolutionary developmental biology
What once was called embryology played a modest role in the evolutionary synthesis,[57] mostly
about evolution by changes in developmental timing (allometry and heterochrony).[58] Man himself was,
according to Bolk, a typical case of evolution by retention of juvenile characteristics (neoteny). He listed many
characters where "Man, in his bodily development, is a primate foetus that has become sexually mature".[59]
Unfortunately, his interpretation of these ideas was non-Darwinian, but his list of characters is both
interesting and convincing.[60]
Modern interest in Evo-devo springs from clear proof that development is closely controlled by
special genetic systems, and the hope that comparison of these systems will tell us much about the
evolutionary history of different groups.[61][62] In a series of experiments with the fruit-fly Drosophila, Edward
B. Lewis was able to identify a complex of genes whose proteins bind to the cis-regulatory regions of target
genes. The latter then activate or repress systems of cellular processes that accomplish the final
development of the organism.[63][64] Furthermore, the sequence of these control genes show co-linearity:
the order of the loci in the chromosome parallels the order in which the loci are expressed along the anterior-
posterior axis of the body. Not only that, but this cluster of master control genes programs the development of
all higher organisms.[65][66] Each of the genes contains a homeobox, a remarkably conserved DNA
sequence. This suggests the complex itself arose by gene duplication.[67][68][69] In his Nobel lecture, Lewis
said "Ultimately, comparisons of the [control complexes] throughout the animal kingdom should provide a
picture of how the organisms, as well as the [control genes] have evolved".
genetic regulatory system is inherited from a common ancestor, as it is in the evolution of vertebrate and
invertebrate eyes. The phenomenon is implicated in many cases of parallel evolution.[71]
A great deal of evolution may take place by changes in the control of development. This may be
relevant to punctuated equilibrium theory, for in development a few changes to the control system could
make a significant difference to the adult organism. An example is the giant panda, whose place in the
mammalia was long uncertain.[72] Apparently, the giant panda's evolution from Ursus to Ailuropoda required
the change of only a few genetic messages (5 or 6 perhaps), yet the phenotypic and lifestyle change from a
standard bear is considerable.[73][74] The transition could therefore be effected relatively swiftly.

[edit] Fossil discoveries

In the past thirty or so years there have been excavations in parts of the world which had scarcely
been investigated before. Also, there is fresh appreciation of fossils discovered in the 19th century, but then
denied or deprecated: the classic example is the Ediacaran biota from the immediate pre-Cambrian, after the
Cryogenian period. These soft-bodied fossils are the first record of multicellular life. The interpretation of this
fauna is still in flux.
Many outstanding discoveries have been made, and some of these have implications for evolutionary
theory. The discovery of feathered dinosaurs and early birds from the Lower Cretaceous of Liaoning, N.E.
China have convinced most students that birds did evolve from coelurosaurian theropod dinosaurs. Less well
known, but perhaps of equal evolutionary significance, are the studies on early insect flight, on stem
tetrapods from the Upper Devonian,[75][76] and the early stages of whale evolution.[77]
with external asymmetry. Their young are perfectly symmetrical, but the head is remodelled during a
metamorphosis, which entails the migration of one eye to the other side, close to the other eye. Some
species have both eyes on the left (turbot), some on the right (halibut, sole); all living and fossil flatfish to date
show an 'eyed' side and a 'blind' side.[78] The lack of an intermediate condition in living and fossil flatfish
species had led to debate about the origin of such a striking adaptation. The case was considered by
Lamark,[79] who thought flatfish precursors would have lived in shallow water for a long period, and by
Darwin, who predicted a gradual migration of the eye, mirroring the metamorphosis of the living forms.
Darwin's long-time critic St. George Mivart thought that the intermediate stages could have no selective
value,[80] and in the 6th edition of the Origin, Darwin made a concession to the possibility of acquired traits.
[81] Many years later the geneticist Richard Goldschmidt put the case forward as an example of evolution by
saltation, bypassing intermediate forms.[82][83]
A recent examination of two fossil species from the Eocene has provided the first clear picture of
flatfish evolution. The discovery of stem flatfish with incomplete orbital migration refutes Goldschmidt's ideas,
and demonstrates that "the assembly of the flatfish bodyplan occurred in a gradual, stepwise fashion".[84]
There are no grounds for thinking that incomplete orbital migration was maladaptive, because stem forms
with this condition ranged over two geological stages, and are found in localities which also yield flatfish with
the full cranial asymmetry. The evolution of flatfish falls squarely within the evolutionary synthesis.[78]

[edit] See also

Evolutionary biology portal
 Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Evolution
• The Origin of Species
• History of evolutionary thought
• Gene-centered view of evolution
• Particulate inheritance theory
• Population genetics
• Developmental systems theory
• Polymorphism (biology)

[edit] Footnotes
1. ^ "The scientific consensus around evolution is overwhelming". "Appendix: Frequently Asked
Questions" (php). Science and Creationism: A View from the National Academy of Sciences (Second
ed.). Washington, DC: The National Academy of Sciences. 1999. p. 28. ISBN ISBN-0-309-06406-6.
http://www.nap.edu/openbook.php?record_id=6024&page=27#p200064869970027001. Retrieved
September 24, 2009.
2. ^ Mayr, Ernst 2002. What evolution is. Weidenfeld & Nicolson, London. p270
3. ^ Huxley J.S. 1942. Evolution: the modern synthesis. Allen & Unwin, London. 2nd ed 1963;
3rd ed 1974.
 4. ^ Mayr & Provine 1998
5. ^ Mayr E. 1982. The growth of biological thought: diversity, evolution & inheritance . Harvard,
Cambs. p567 et seq.
6. ^ Smocovitis, V. Betty. 1996. Unifying Biology: the evolutionary synthesis and evolutionary
biology. Princeton University Press. p192
7. ^ Gould S.J. Ontogeny and phylogeny. Harvard 1977. p221-2
8. ^ Bowler P.J. 2003. Evolution: the history of an idea. pp236–256
9. ^ a b Gould The Structure of Evolutionary Theory p. 216
10.^ Kutschera U. 2003. A comparative analysis of the Darwin-Wallace papers and the
development of the concept of natural selection. Theory in Biosciences 122, 343-359
11.^ Bowler pp. 253–256
12.^ Mike Ambrose. "Mendel's Peas". Genetic Resources Unit, John Innes Centre, Norwich, UK.
http://www.jic.ac.uk/germplas/pisum/zgs4f.htm. Retrieved 2007-09-22.
13.^ Bateson, William 1894. Materials for the study of variation, treated with special regard to
discontinuity in the origin of species. The division of thought was between gradualists of the
Darwinian school, and saltationists such as Bateson. Mutations (as 'sports') and polymorphisms were
well known long before the Mendelian recovery.
14.^ Larson pp. 157–166
15.^ Grafen, Alan; Ridley, Mark (2006). Richard Dawkins: How A Scientist Changed the Way
We Think. New York, New York: Oxford University Press. p. 69. ISBN 0199291160.
16.^ Bowler pp. 271–272
17.^ Transactions of the Royal Society of Edinburgh, 52:399–433
18.^ a b c d e f g h Larson Evolution: The Remarkable History of a Scientific Theory pp. 221–243
19.^ a b c d e f Bowler Evolution: The history of an Idea pp. 325–339
 20.^ Gould The Structure of Evolutionary Theory pp. 503–518
21.^ Mayr & Provine 1998 p. 231
22.^ Dobzhansky T. 1951. Genetics and the Origin of Species. 3rd ed, Columbia University
Press N.Y.
23.^ Ford E.B. 1964, 4th edn 1975. Ecological genetics. Chapman and Hall, London.
24.^ Mayr and Provine 1998 pp. 33–34
25.^ Smith, Charles H.. "Rensch, Bernhard (Carl Emmanuel) (Germany 1900–1990)". Western
Kentucky University. http://www.wku.edu/~smithch/chronob/RENS1900.htm. Retrieved 2007-09-22.
26.^ Mayr and Provine 1998 pp. 298–299, 416
27.^ Bowler p.361
28.^ Pigliucci, Massimo 2007. Do we need an extended evolutionary synthesis? Evolution 61
12, 2743–2749.
29.^ Dalrymple, G. Brent 2001. The age of the Earth in the twentieth century: a problem (mostly)
solved. Special Publications, Geological Society of London 190, 205–221.
30.^ Van Andel, Tjeerd 1994. New views on an old planet: a history of global change . 2nd ed.
Cambridge.
31.^ Witz A. 2006. The start of the world as we know it. Nature 442, p128.
32.^ Schopf J.W. and Klein (eds) 1992. The Proterozoic biosphere: a multi-disciplinary study .
Cambridge University Press.
33.^ Lane, Nick 2002. Oxygen: the molecule that made the world. Oxford.
34.^ Schopf J.W. 1999. Cradle of life: the discovery of Earth's earliest fossils . Princeton.
35.^ Yochelson, Ellis L. 1998. Charles Doolittle Walcott: paleontologist. Kent State, Ohio.
36.^ Knoll A.H. and Holland H.D. 1995. Oxygen and Proterozoic evolution: an update. In
National Research Council, Effects of past climates upon life. National Academy, Washington D.C.
 37.^ Huggett, Richard J. 1997. Catastrophism. new ed. Verso.
38.^ Hallam A. and Wignall P.B. 1997. Mass extinctions and their aftermath. Columbia, N.Y.
39.^ Elewa A.M.T. (ed) 2008. Mass extinctions. Springer, Berlin.
40.^ The terms (or their equivalents) were used as part of the synthesis by Simpson G.G. 1944.
Tempo and mode in evolution, and Rensch B. 1947. Evolution above the species level. Columbia,
N.Y. They were also used by some non-Darwinian evolutionists such as Yuri Filipchenko and
Richard Goldschmidt. Here we use the terms as part of the evolutionary synthesis: they do not imply
any change in mechanism.
41.^ Maynard Smith J. and Szathmáry E. 1997. The major transitions in evolution. Oxford.
42.^ de Bary, H.A. 1879. Die Erscheinung der Symbiose. Strassburg.
43.^ Khakhina, Liya Nikolaevna 1992. Concepts of symbiogenesis: a historical and critical study
of the research of Russian scientists.
44.^ Wilson E.B. 1925. The cell in development and heredity . Macmillan, N.Y.
45.^ Walin I.E. 1927. Symbionticism and the origin of species. Williams & Wilkins, Baltimore.
46.^ Wells H.G., Huxley J. and Wells G.P. 1930. The science of life. London vol 2, p505. This
section (The ABC of genetics) was written by Huxley.
47.^ Sapp, January 1994. Evolution by association: a history of symbiosis. Oxford.
48.^ Lederberg J. 1952. Cell genetics and hereditary symbiosis. Physiological Reviews 32, 403–
430.
49.^ Margulis L and Fester R (eds) 1991. Symbiosis as a source of evolutionary innovation . MIT.
50.^ Margulis L. 1993. Symbiosis in cell evolution: microbial communities in the Archaean and
Proterozoic eras. Freeman, N.Y.
51.^ Maynard Smith J. and Szathmáry E. 1997. The major transitions in evolution. Oxford. The
origin of the eukaryote cell is one of the seven major transitions, according to these authors.
 52.^ Woese, Carl 1998. The Universal Ancestor. PNAS 95, 6854–6859.
53.^ Doolittle, W. Ford 1999. Phylogenetic classification and the Universal Tree. Science 284,
2124–2128.
54.^ Doolittle, W. Ford 2000. Uprooting the tree of life. Scientific American 282 (6): 90–95.
55.^ Villarreal LP, Witzany G. 2010. Viruses are essential agents within the roots and stem of
the tree of life. Journal of Theoretical Biology 262(4): 698-710.
56.^ Dunn, Casey W. et al 2009. Broad phylogenetic sampling improves resolution of the animal
tree of life. Nature 452, 745–749.
57.^ Laubichler M. and Maienschein J. 2007. From Embryology to Evo-Devo: a history of
developmental evolution. MIT.
58.^ de Beer, Gavin 1930. Embryology and evolution. Oxford; 2nd ed 1940 as Embryos and
ancestors; 3rd ed 1958, same title.
59.^ Bolk, L. 1926. Der Problem der Menschwerdung. Fischer, Jena.
60.^ short-list of 25 characters reprinted in Gould, Stephen Jay 1977. Ontogeny and phylogeny.
Harvard. p357
61.^ Raff R.A. and Kaufman C. 1983. Embryos, genes and evolution: the developmental-genetic
basis of evolutionary changes. Macmillan, N.Y.
62.^ Carroll, Sean B. 2005. Endless forms most beautiful: the new science of Evo-Devo and the
making of the animal kingdom. Norton, N.Y.
63.^ Lewis E.B. 1995. The bithorax complex: the first fifty years. Nobel Prize lecture. Repr. in
Ringertz N. (ed) 1997. Nobel lectures, Physiology or Medicine. World Scientific, Singapore.
64.^ Lawrence P. 1992. The making of a fly. Blackwell, Oxford.
65.^ Duncan I. 1987. The bithorax complex. Ann. Rev. Genetics 21, 285–319.
 66.^ Lewis E.B. 1992. Clusters of master control genes regulate the development of higher
organisms. J. Am. Medical Assoc. 267, 1524–1531.
67.^ McGinnis W. et al 1984. A conserved DNA sequence in homeotic genes of the Drosophila
antennipedia and bithorax complexes. Nature 308, 428–433.
68.^ Scott M.P. and Weiner A.J. 1984. Structural relationships among genes that control
developmental sequence homology between the antennipedia, ultrabithorax and fushi tarazu loci of
Drosophila. PNAS USA 81, 4115.
69.^ Gehring W. 1999. Master control systems in development and evolution: the homeobox
story. Yale.
70.^ Shubin N, Tabin C and Carroll S. 1997. Fossils, genes and the evolution of animal limbs.
Nature 388, 639–648.
71.^ Shubin N, Tabin C and Carroll S. 2009. Deep homology and the origins of evolutionary
novelty. Nature 457, p818–823.
72.^ Sarich V. 1976. The panda is a bear. Nature 245, 218–220.
73.^ Davies D.D. 1964. The giant panda: a morphological study of evolutionary mechanisms.
Fieldiana Memoires (Zoology) 3, 1–339.
74.^ Stanley Steven M. 1979. Macroevolution: pattern & process. Freeman, San Francisco.
p157
75.^ Clack, Jenny A. 2002. Gaining Ground: the origin and evolution of tetrapods . Bloomington,
Indiana. ISBN 0-253-34054-3
76.^ Home page - Jenny Clack
77.^ Both whale evolution and early insect flight are discussed in Raff R.A. 1996. The shape of
life. Chicago. These discussions provide a welcome synthesis of evo-devo and paleontology.
78.^ a b Janvier, Philip 2008. Squint of the fossil flatfish. Nature 454, 169
 79.^ Lamark J.B. 1809. Philosophie zoologique. Paris.
80.^ Mivart St G. 1871. The genesis of species. Macmillan, London.
81.^ Darwin, Charles 1872. The origin of species. 6th ed, Murray, London. p186–188. The whole
of Chapter 7 in this edition is taken up with answering critics of natural selection.
82.^ Goldschmidt R. Some aspects of evolution. Science 78, 539–547.
83.^ Goldschmidt R. 1940. The material basis of evolution. Yale.
84.^ Friedman, Matt 2008. The evolutionary origin of flatfish asymmetry. Nature 454, 209–212.

[edit] References
• Allen, Garland. Thomas Hunt Morgan: The Man and His Science, Princeton University Press,
1978 ISBN 0-691-08200-6
• Bowler, Peter J. (2003). Evolution:The History of an Idea. University of California Press.
ISBN 0-52023693-9.
• Dawkins, Richard. The Blind Watchmaker, W.W. Norton and Company, Reissue Edition
1996 ISBN 0-393-31570-3
• Dobzhansky, T. Genetics and the Origin of Species, Columbia University Press, 1937 ISBN
0-231-05475-0
• Fisher, R. A. The Genetical Theory of Natural Selection , Clarendon Press, 1930 ISBN 0-19-
850440-3
• Futuyma, D.J. Evolutionary Biology, Sinauer Associates, 1986, p. 12 0-87-893189-9
• Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Belknap Press of Harvard
University Press. ISBN 0-674-00613-5.
 • Haldane, J. B. S. The Causes of Evolution, Longman, Green and Co., 1932; Princeton
University Press reprint, ISBN 0-691-02442-1
• Huxley, J. S., ed. The New Systematics, Oxford University Press, 1940 ISBN 0-403-01786-6
• Huxley, J. S. Evolution: The Modern Synthesis, Allen and Unwin, 1942 ISBN 0-02-846800-7
• Larson, Edward J. (2004). Evolution:The Remarkable History of a Scientific Theory. Modern
Library. ISBN 0-679-64288-9.
• Margulis, Lynn and Dorion Sagan. "Acquiring Genomes: A Theory of the Origins of Species",
Perseus Books Group, 2002 ISBN 0-465-04391-7
• Mayr, E. Systematics and the Origin of Species, Columbia University Press, 1942; Harvard
University Press reprint ISBN 0-674-86250-3
• Mayr, E. and W. B. Provine, eds. The Evolutionary Synthesis: Perspectives on the
Unification of Biology, Harvard University Press, 1998 ISBN 0-674-27225-0
• Simpson, G. G. Tempo and Mode in Evolution, Columbia University Press, 1944 ISBN 0-
231-05847-0
• Smocovitis, V. Betty. Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology ,
Princeton University Press, 1996 ISBN 0-691-27226-9
• Wright, S. 1931. "Evolution in Mendelian populations". Genetics 16: 97–159.

[edit] External links

• Rose MR, Oakley TH, The new biology: beyond the Modern Synthesis. Biology Direct 2007,
2:30. A review of biology in light of recent innovations since the initiation of modern synthesis.
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 W000

Théorie synthétique de l'évolution

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Aller à : Navigation, rechercher

Pour les articles homonymes, voir TSE.

La théorie synthétique de l'évolution constitue le cadre conceptuel le plus largement utilisé dans
l'étude scientifique des processus d'évolution en biologie. Cette théorie est basée sur l'intégration de la
théorie de l'hérédité mendélienne et de la génétique des populations à la théorie darwinienne[1]. Cette
synthèse fut menée au cours des années 1930 et 1940 par R.A. Fisher, J.B.S Haldane, Sewall Wright, Julian
Huxley, Ernst Mayr, Bernhard Rensch, George Gaylord Simpson et George Ledyard Stebbins. Baptisée ainsi
par Julian Huxley en 1942, cette théorie est aussi appelée néodarwinisme ou synthèse néodarwinienne pour
souligner le fait qu'elle constitue une extension de la théorie originale de Charles Darwin qui ignorait les
mécanismes de l'hérédité génétique.
Sommaire
[masquer]
• 1
Histoire de la
théorie
synthétique de
l'évolution
• 2
Définition
• 3
L'évolution,
faits et théorie
• 4
Mécanismes
de l'évolution
• 4
.1
L'appa
rition
de
caract
ères
Histoire de la théorie synthétique de l'évolution [modifier]
Article détaillé : Histoire de la pensée évolutionniste.

Définition [modifier]
Aujourd'hui, l'évolution n'est plus envisagée comme la transformation d'individus isolés mais comme
celle de groupements d'individus de même espèce, c'est-à-dire des populations. Mais le principe de base est
le même : elle explique l'évolution par l'action de la sélection naturelle sur des populations.
Une population évolue quand la fréquence d'une version d'un gène appelée allèle (ou de plusieurs
allèles) s'y modifie. On voit ainsi se répandre dans certaines espèces des caractères ayant acquis, en raison
de changements du milieu, une valeur adaptative qu'ils ne possédaient pas auparavant ; les individus
porteurs de ces caractères sont particulièrement favorisés dans le nouveau milieu, auquel ils se trouvaient
en quelques sortes "préadaptés" ; ils constituent alors rapidement une grande partie de la population ou
même toute la population de l'espèce. Les caractères qui se répandent alors correspondent à des allèles
existant auparavant "silencieusement" au sein de l'espèce.
Lorsque l'ensemble des individus qui constituent une espèce forme plusieurs populations isolées,
chacune de ces populations peut acquérir des caractères particuliers et donner naissance à des variétés
différentes au sein de la même espèce. Si ces variations sont, par la suite, dans l'impossibilité de se croiser,
elles divergent de plus en plus et finalement sont interstériles : elles constituent alors des espèces distinctes.
La "barrière" qui sépare les variétés d'une même espèce peut être de nature variée.
L'évolution, faits et théorie [modifier]

Le membre antérieur de tous les Vertébrés présente une structure homologue.

 Une observation des êtres vivants révèle l'existence de nombreux points communs dans
l'organisation anatomique, physiologique et biochimique des organismes. Depuis l'Antiquité, l'Homme a tenté
de trouver un ordre dans la nature[2] selon les ressemblances et les différences qu'il observait chez les
animaux et les végétaux donnant naissance à diverses classifications des espèces. Si certaines espèces se
ressemblent beaucoup et d'autres moins, l'unité du vivant n'a guère été remise en cause ; elle a même été
renforcée d'abord au cours du XIXe siècle par la découverte de la cellule, unité vivante fondamentale de tous
les organismes, puis au XXe siècle par la découverte de l'ADN comme support de l'hérédité commun à tout
le vivant.
Le phénomène d'évolution permet d'expliquer ces ressemblances par l'existence de liens
généalogiques entre toutes les formes de vie : les organismes se ressemblent parce qu'ils partagent des
caractères hérités d'un ancêtre commun[3]. Des faits observés dans plusieurs disciplines scientifiques
corroborent cette théorie.
Tous les êtres vivants fonctionnent sur les mêmes bases moléculaires (ADN, ARN, protéines…) et
utilisent le même code génétique. Les différences entre les espèces et les variations entre individus au sein
d'une même espèce sont dues essentiellement à des différences dans la séquence des gènes et la structure
des chromosomes, différences provoquées par des mutations et des réarrangements chromosomiques
occasionnels. Ces évènements aléatoires, qui produisent des individus dont les gènes sont légèrement
différents de ceux de leurs parents, sont certes rares et - comme une coquille qui se glisse dans l'impression
d'un texte - plus souvent nocifs qu'utiles, mais suffisent pour introduire la variabilité, source de l'évolution[4].
Les différentes espèces d'un embranchement, malgré des aspects extérieurs très différents,
partagent un plan d'organisation invariable[5]. L'homologie des organes entre tous les membres d'un groupe
biologique se retrouve à n'importe quel niveau de la classification. Par exemple, chez tous les mammifères,
les membres sont organisés de la même façon : ceux d'une baleine, d'un lion, d'une chauve-souris ou d'un
homme comportent les mêmes os, même si leur forme est très variable et qu'ils sont employés à des
fonctions complètement différentes.
Par ailleurs, il existe une certaine variabilité des caractères au sein de chaque espèce, c'est-à-dire
qu'il n'existe aucun individu complètement identique à un autre sauf cas exceptionnels. Cette variation peut
être de plus ou moins grande ampleur, mais elle montre en tout cas que les êtres vivants ne sont pas fixes.
L'éventail de variations que présente une population vivante est le matériau de base avec lequel l'évolution
pourrait construire des organismes de plus en plus différents. On assiste directement à des transformations
au sein de populations d'êtres vivants pouvant être interprétées comme des spéciations en cours[6].
L'acquisition de nouveaux caractères chez certains virus et bactéries a également été constatée pour être
combattue, dans le cas de la résistance aux antibiotiques, ou au contraire utilisée, pour la capacité à digérer
des composés synthétiques.
Les fossiles témoignent de la disparition de certaines espèces d'êtres vivants depuis l'apparition de
la vie sur Terre il y a au moins 3,5 milliards d'années[7]. Certaines espèces sont appelées des formes
« intermédiaires » entre différents groupes et permettent de mettre en évidence le phénomène d'évolution[8].
Par exemple, l'archéoptéryx, qui serait une forme de transition entre certains dinosaures et les oiseaux, ou
Rodhocetus (ou Ambulocetus) dont l'anatomie semble se situer entre celle des mammifères terrestres et
celle des baleines.
Les détracteurs de l'évolution soulignent qu'elle n'est « qu'une théorie ». En fait il ne s'agit pas d'une
théorie au sens commun mais dans le sens scientifique du terme[9]. Cela signifie qu'il s'agit d'un ensemble
de connaissances basé sur quelques suppositions de départ, et développé par des démonstrations
rigoureuses à partir de ces principes. Le développement de la théorie est donc toujours susceptible d'être
complété, ou même corrigé sur certains points. En revanche il n'est pas correct de considérer l'ensemble de
l'édifice comme une simple hypothèse, sauf à disposer d'arguments forts contre les mécanismes de base de
la théorie qui ont été observés en conditions expérimentales et naturelles.
Mécanismes de l'évolution [modifier]
 Illustrations de cinq types de mutations chromosomiques.
L'évolution dépend de divers processus qui tendent à modifier la fréquence des allèles au sein de
populations : mutation : dérive génétique, étranglement génétique ; flux de gènes, brassage génétique ;
sélection naturelle : sélection stabilisante, directionnelle ou diversifiante ; sélection sexuelle : accouplement
non aléatoire, ... ; pédomorphose ; pléiotropie : pléiotropie antagoniste ; ...
L'évolution des espèces est permise par les mutations que subissent les gènes portés par les
chromosomes (constitués d'ADN). Tout être vivant possède de l'ADN (cela corrobore l'idée d'une origine
commune des espèces) : cette molécule est constituée de deux hélices complémentaires au niveau de leurs
bases azotées. Des mutations peuvent affecter cet ADN ; elles sont provoquées par des agents mutagènes
tels que rayons X, alpha, UV, ou tout simplement par la défaillance des organites responsables de la
réparation de l'ADN mal transcrit ou traduit. La théorie des monstres prometteurs met le stress provoqué par
des facteurs externes au premier plan de ces facteurs de mutation. Ces mutations affectent la séquence d'un
gène concerné (ordre des bases nucléotides d'un gène : adénine, thymine, guanine et cytosine).
Ces mutations sont à l'origine du polymorphisme des gènes, c'est-à-dire le fait que deux versions d'un même
gène (deux allèles) par exemple, sont présentes chacune dans au moins un pour cent de la population de
l'espèce considérée. Ces mutations créent donc de nouveaux allèles. Mais il ne faut pas oublier le
phénomène des familles multigéniques, qui a un impact très important dans l'évolution : un gène peut être
dupliqué et transposé sur un autre chromosome, ainsi tous les gènes issus de ce gène ancestral font partie
d'une famille multigénique.
L'environnement « encadre » ces mutations par le biais d'un phénomène appelé sélection naturelle :
un gène présentant un avantage pour une espèce dans un environnement donné, permettant à ses
représentants d'atteindre le mieux possible la maturité sexuelle, se répand chez les individus d'une même
espèce, a contrario s'il est néfaste, il disparait. Quant aux gènes neutres, ils se répandent de façon aléatoire
mais peuvent permettre de suivre l'évolution (mutation de gènes homéotiques). C'est donc l'environnement
qui décide de l'évolution des espèces, celles-ci évoluant pour être toujours plus adaptées à celui-ci. Donc le
caractère aléatoire des mutations de l'ADN est compensé par la sélection environnementale.
Il est donc très important de ne pas confondre évolution et innovation, ce qu'il faut retenir, c'est que
l'espèce qui survit est l'espèce la mieux adaptée, pendant une certaine durée, à son environnement.
Par ailleurs, il ne faut surtout pas voir l'évolution d'un point de vue généalogique, mais
phylogénétique, en effet les espèces ne descendent pas les unes des autres. Des phénomènes comme la
dérive génétique font que deux populations d'une même espèce isolées pendant une très longue période de
temps divergent et forment deux nouvelles espèces. Par exemple pour la lignée humaine, l'arbre
phylogénétique est buissonnant : plusieurs espèces Homo et Australopithèque ont vécu simultanément. Il est
aussi à noter que l'homme - contrairement aux idées reçues - ne descend pas "du singe", il a un ancêtre en
commun avec lui. Et cet ancêtre n'était pas un singe. Notre patrimoine génétique est très proche de celui des
chimpanzés, en effet les différences entre nous et celui-ci ne tiendraient qu'à la différence de quelques
gènes déterminants : ils nous permettent de garder des caractères juvéniles toute notre vie, nous permettant
de conserver la bipédie (le petit chimpanzé est quasi-bipède) et ils allongent la phase embryonnaire,
permettant la mitose de beaucoup plus de neurones.

L'évolution résulte de l'action de plusieurs mécanismes évolutifs susceptibles de modifier la fréquence des
génotypes dans une population, et par conséquent de modifier des caractères morphologiques,
physiologiques ou comportementaux.
Le processus est dans tous les cas identique, dans le même ordre :
 1. Apparition d'un nouveau caractère héréditaire chez un ou quelques individus.
2. Ce nouveau caractère se répand dans les populations, soit parce qu'il est plus adapté à
l'environnement, soit à cause d'un effet de hasard.

L'apparition de caractères nouveaux et héréditaires [modifier]

Les mécanismes de
l'évolution biologique

Mécanismes non aléatoires :

• sélection naturelle
• sélection
utilitaire
• sélection
sexuelle
• sélection de
parentèle
• sélection de
groupe
• sélection
stratégique
 • sélection artificielle

Mécanismes aléatoires :
• mutation génétique
• recombinaison
• dérive génétique

Conséquences de l'évolution :
• spéciation
• adaptation des
espèces
• radiation évolutive
A chaque génération, de nouveaux variants génétiques apparaissent qui peuvent eux-mêmes
transmettre leur patrimoine génétique au moment de la reproduction. Ces sources de diversité génétique
sont :
• les mutations ponctuelles dans la séquence d'ADN des gènes ;
• les réarrangements chromosomiques ;
• les recombinaisons ou brassages génétiques qui se produisent lors de la reproduction
sexuée et, chez les bactéries, lors de transferts d'ADN entre cellules.
 On peut ajouter à cette liste une autre source de diversité génétique que sont les migrations par
lesquelles le stock génétique dans une population donnée se voit renouvelé par l'arrivée d'autres membres
de l'espèce porteurs d'un pool génétique différent.

La mutation [modifier]
Les mutations résultent d'erreurs lors de la réplication des gènes. Ces modifications accidentelles
peuvent être transmises à la descendance si elles touchent les cellules germinales, donnant naissance à des
individus porteurs de gènes nouveaux, qu'aucun de leurs parents n'exprimaient.
Il existe plusieurs types de mutations (mutations ponctuelles, duplications de gènes, cassures des
chromosomes, insertions ou délétions de séquences). Ces différentes modifications suffisent à expliquer la
diversité génétique observée dans la nature.
Il peut s'y ajouter des mutations par échange de matériel génétique entre espèces, par différents
mécanismes naturels :
• Les bactéries sont capables d'intégrer et d'utiliser du matériel génétique (des plasmides)
échangé entre deux bactéries, voire simplement présent dans le milieu. Le plus souvent cela ne
conduit pas immédiatement à la définition d'une nouvelle espèce, mais le phénomène peut y
contribuer et il contribue de façon importante à la sélection naturelle (par exemple, de nombreuses
résistances aux antibiotiques se diffusent de cette façon).
• Les virus peuvent servir de vecteurs de transfert de matériel génétique d'une espèce vers
une autre, même très différente ; le plus souvent, l'infection virale conduit à la mort de la cellule
victime, mais ce n'est pas nécessairement le cas, et la cellule cible peut participer à la reproduction
 (notamment pour les plantes). Un des exemples les mieux connus (et les plus utilisés par la
recherche) est celui du tabac et des mosaïques.
• Les plantes peuvent s'hybrider, conduisant à de nouvelles espèces (le blé étant un exemple
typique).
Finalement, il faut mentionner les symbioses strictes, qui sont des créations de nouvelles espèces à
partir d'anciennes, mais sans échange ni fusion du matériel génétique (au moins dans un premier temps).
Les lichens et, selon l'hypothèse de l'endosymbiose, les organites dits « autonomes » à l'intérieur des
cellules (chloroplastes, mitochondries) relèvent de ce type d'association.
Les mutations augmentent la variabilité génétique au sein d'une population. Cependant, elles sont
rarement bénéfiques à l'individu qui les porte car elles peuvent détruire l'activité d'un gène essentiel.

La migration [modifier]
La migration est une force de l'évolution qui augmente la variabilité génétique d'une population. De
nouveaux arrivants en provenance d'une population éloignée amènent des allèles nouveaux, augmentant le
pool génétique de la population d'accueil.

Le nouveau caractère se répand dans les populations [modifier]

Les mécanismes de l'évolution supposent qu'un ensemble d'individus, regroupés en une espèce sur
la base de caractères partagés, évoluent « conjointement. » Se pose donc le problème de la spéciation,
c'est-à-dire de l'évolution des espèces au-delà des simples individus.
 La dérive aléatoire [modifier]
Article détaillé : dérive génétique.
Lorsqu'une petite population est isolée du reste d'une population-mère (par exemple, par isolement
sur une petite île), certains types d'allèles tendent à disparaître, du fait de faible nombre de copies
distribuées dans le pool génétique de la population au départ. Au fil des générations, la population dérive
alors, acquérant des caractéristiques qui lui sont propres (ex: petitesse (mammouth nain de Sicile),
gigantisme (varan de Komodo) parmi les espèces insulaires). La dérive est une force de l'évolution qui
diminue la variabilité génétique au sein de la population. La dérive influe sur la fréquence des allèles. Elle ne
crée en aucun cas de nouveaux allèles. Certains allèles vont être surreprésentés et d'autres sous-
représentés par rapport à la population d'origine. Cette modification de la fréquence s'explique par une erreur
d'échantillonnage. (hasard)
 La sélection naturelle [modifier]

Sélection naturelle et mutation.

Article détaillé : sélection naturelle.
Elle diminue la variabilité génétique d'une population en enlevant certaines combinaisons génétiques
défavorables dans le milieu où elles se produisent. Cependant elle peut aussi maintenir à une fréquence
stable, des nouveaux variants génétiques, sans pour autant éliminer les anciennes versions. C'est le cas
pour l'allèle de la drépanocytose. La sélection naturelle participe donc à l'augmentation de la diversité
génétique.
Une image donnée par Richard Dawkins [modifier]
Pour se trouver mille ancêtres différents, il faut remonter en arrière d'une dizaine de générations, ce
qui représente quelques siècles. Or, avant le XIXe siècle, la moitié au moins des enfants mouraient en bas
âge : on peut donc se demander combien de nos mille derniers ancêtres sont morts en bas âge, et la
réponse n'est pas « au moins la moitié », mais bien entendu zéro, par définition. Nous ne sommes donc
nullement représentatifs de l'humanité passée, car descendants d'une longue lignée de gens qui ont tous eu
la chance d'amener une progéniture à l'âge de la procréation (c'est-à-dire survivre et trouver un partenaire,
entre autres). Une génération peut avoir de la chance. Quand une dizaine en a coup sur coup, on peut
supposer que cette « chance » correspond en fait à un ensemble de facteurs favorables qui se retrouvent de
l'une à l'autre (facteurs qui ne sont pas tous génétiques : ils peuvent être culturels, religieux, économiques,
etc.) Cette considération à elle seule montre que même à notre échelle, nous avons participé un tout petit
peu, sur les quelques derniers siècles, à l'évolution. D'ailleurs notre simple choix d'un conjoint se révèle,
comme le montre le biologiste Geoffrey Miller (The Mating Mind) obéir à des choix pas toujours conscients
qu'on peut rapprocher de l'eugénisme.
Richard Dawkins pousse plus avant ses spéculations en considérant l'être vivant comme un
conteneur à gènes (ce serait en tout cas le point de vue des gènes, s'ils en avaient un !). Pour lui les produits
des gènes - cellules, tissus, organes, organismes, sociétés - servent aux gènes à se répliquer et à survivre.
En quelque sorte, à la question : « De l'œuf ou la poule, qui est le premier ? », il répond : « La poule est le
moyen trouvé par l'œuf pour faire d'autres œufs. ». Ce renversement de perspective considère l'être vivant
comme la marionnette (Dawkins utilise le terme de véhicule-robot) de ses gènes.
Tout en la reconnaissant ingénieuse, Stephen Jay Gould a mis en garde contre une prise trop à la
lettre de cette vision. Les gènes ne possèdent ni intention, ni projet au sens que nous donnons à ces termes.
Dawkins n'a utilisé le terme qu'en tant que métaphore parlante, mais ses lecteurs n'ont pas toujours saisi la
nuance.
En outre, il convient de se rappeler que les facteurs de survie ne sont pas seulement génétiques, ce
qui est bien évident chez l'homme, mais ce qui est aussi le cas chez de nombreux animaux (par exemple, le
chant des oiseaux, essentiel dans leur reproduction, dépend d'un apprentissage).

Devenir des mutations et caractères hérités [modifier]

• Les mutations se font toujours au hasard, et sont le plus souvent létales. La plupart du
temps, elles perturbent gravement le fonctionnement de l'organisme mutant. Bien sûr, il arrive aussi
qu'elles soient neutres (en modifiant un caractère sans pour autant le rendre moins fonctionnel) ou
avantageuses; le cas est beaucoup plus rare, mais les mutations portent sur des millions d'individus
pendant des dizaines de milliers de générations ou davantage, ce qui assure un facteur multiplicatif
de 1010 à chacun de ces évènements aléatoires.
• C'est la sélection naturelle, mise en évidence par Darwin, qui se charge de « faire le tri »
entre les différentes variations. Il s'agit d'un processus purement mécanique : les mutants
défavorisés auront tendance soit à mourir plus jeunes que les autres, soit à trouver plus difficilement
un compagnon de reproduction. Dans un cas comme dans l'autre, ils laisseront moins (voire pas du
tout) de descendants. Les mutants neutres mourront en moyenne au même âge que les autres,
auront globalement autant d'enfants et pourront donc répandre leurs nouveaux caractères dans une
population sous l'effet du hasard. Quant aux mutants avantagés, ils seront plus compétitifs et auront
donc en moyenne plus de descendants. Une mutation qui permet de mieux échapper à un prédateur,
de mieux s'orienter, de mieux séduire le sexe opposé, d'avoir plus de descendants, de mieux résister
 aux maladies, de mieux tirer profit d'une vie en société aura tendance à faire baisser le taux de
mortalité chez les mutants ou à améliorer leur succès reproductif.
• Génération après génération, la sélection naturelle favorise les mutations qui se transmettent
le plus, dans les populations (la dérive génétique, qui se fait au hasard, permet également à
quelques mutations neutres de se fixer dans le génome de l'espèce). L'adaptation des populations
tend donc progressivement à s'améliorer ; l'émergence d'organes aussi complexes et aussi
fonctionnels que l'œil humain ou le radar de la chauve-souris s'explique par un tel processus de
sélection cumulative. En revanche, l'existence de caractères sans utilité adaptative apparente peut
s'expliquer par la fixation au hasard de mutations neutres. L'hypothèse dite de « l'auto-stop », ou
« hitch hiking » en anglais, explique la fixation des mutations neutres par la mutation sur le même
chromosome de deux gênes, l'une étant neutre et l'autre étant positive qui se transmettront
simultanément.

Conséquences de l'évolution [modifier]

Apparition d'espèces nouvelles [modifier]

Article détaillé : spéciation.
La spéciation désigne l'apparition de nouvelles espèces à partir d'une espèce ancestrale. Une
espèce est un ensemble d'individus interféconds (interfertiles), isolés du point de vue reproductif des autres
êtres vivants. L'apparition d'une barrière reproductrice (arrêt des échanges génétiques entre elles) au sein de
cet ensemble homogène entraine l'apparition de deux groupes isolés (isolement reproductif). Les deux
groupes vont ensuite évoluer indépendamment l'un de l'autre, et vont progressivement accumuler des
différences génétiques, jusqu'à ce que l'isolement devienne irréversible.
 • La spéciation dite allopatrique concerne deux ou plusieurs populations isolées
géographiquement, séparées les unes des autres par des barrières géographiques (océan,
montagne…). Un petit isolat peut aussi se former en périphérie de l'aire globale de distribution de
l'espèce. Chaque population va alors pouvoir évoluer indépendamment, d'autant plus rapidement
qu'elle est petite (les mutations se fixent plus facilement, pour des raisons mathématiques, dans les
petites populations) et ainsi accumuler des remaniements chromosomiques pour finir par former une
nouvelle espèce.
• La spéciation sympatrique : il arrive aussi qu'un isolement reproducteur apparaisse au sein
d'une population qui ne sera jamais séparée géographiquement. Il suffit pour cela que quelques
individus changent de période de reproduction ou de signaux pour la parade nuptiale pour que les
autres représentants de l'espèce cessent rapidement de s'accoupler avec eux ; il y a alors formation
de deux communautés se partageant le même espace, mais s'isolant sur le plan de la reproduction.
Les deux populations, en évoluant chacune de leur côté, pourront finir par se transformer en deux
espèces complètement différentes.
Comme les spéciations sont des phénomènes relativement courts, qui se produisent dans de petites
populations isolées, on observe souvent une certaine discontinuité entre les différentes espèces dans les
archives fossiles. Une espèce peut ne pas changer pendant très longtemps, puis être rapidement (à l'échelle
géologique) remplacée par une autre.
 Les mutations à l'origine des grandes modifications évolutives [modifier]
Certaines mutations minimes permettent à l'évolution d'agir très rapidement. On connait des gènes
régulateurs, dits gènes homéotiques, qui déterminent les grandes lignes du développement et du plan
d'organisation de chaque partie du corps. Lorsque ces gènes (qui régulent eux-mêmes quelques milliers
d'autres gènes) mutent, l'individu obtenu est souvent porteur de caractères nettement différents de ceux de
ses parents. Certains « sauts évolutifs » s'expliquent par ce type de « macro-mutations » (que Goldschmidt
avait découvert en proposant sa théorie du « monstre prometteur »). Par exemple, l'apparition de doigts et la
disparition des rayons au bout de la nageoire des poissons crossoptérygiens (c'est-à-dire la formation des
premières ébauches de pattes) s'expliquent par un simple décalage de l'activité de deux gènes
homéotiques : Hox-a et Hox-b. Cette étape évolutive-là n'a pas nécessité des milliers, ni même des dizaines
de mutations différentes : les gènes homéotiques ont permis l'émergence rapide de caractères radicalement
nouveaux. Même si les « macro-mutations » donnent le plus souvent des « monstres » incapables de
survivre et même si le processus, lent et sûr, de sélection cumulative de milliers de mutations minuscules a
apparemment joué un rôle plus important dans l'évolution, il est de fait que certains « monstres
prometteurs » ont permis à l'évolution de faire de grands bonds en avant.
D'autres mutations modifient la chronologie du développement : on parle d'hétérochronies. Elles
peuvent être à l'origine de l'apparition d'une espèce plus juvénile (paedomorphique) ou, au contraire, plus
adulte (hypermorphique) que son ancêtre. On connait plusieurs exemples d'histoires évolutives qui ont fait
intervenir les hétérochronies. L'axolotl, un urodèle mexicain qui passe sa vie entière à l'état larvaire sans
jamais se métamorphoser, est paedomorphique par rapport à son ancêtre l'ambystome. Une simple
hétérochronie a permis l'apparition d'une espèce complètement différente. De même, il semblerait que
l'homme soit paedomorphique par rapport à ses ancêtres simiens, car il garde toute sa vie des caractères
juvéniles que le chimpanzé et le gorille perdent. De plus, les phases générales de son développement sont
ralenties par rapport à celles des grands singes.
L'apparition de nouveaux gènes s'explique principalement par la duplication de gènes préexistants.
On connaît plusieurs « familles de gènes » dont les différents membres sont apparemment issus de plusieurs
duplications d'un même gène ancestral. Par la suite, chacun des exemplaires « dupliqués » a pu muter et
évoluer indépendamment des autres. Plusieurs familles multigéniques sont de toute évidence apparues
grâce à ce processus.
De même, des duplications de l'ensemble du génome (polyploïdisations) ont également pu jouer. Par
exemple, on estime que deux épisodes de polyploïdisations se sont produits depuis la séparation des
deutérostomiens (soient essentiellement les échinodermes et les chordés dont les vertébrés) et des
protostomiens (les invertébrés) sur la branche des vertébrés. On retrouve ainsi que beaucoup de gènes,
comme HedgeHog impliqué dans le développement des polarités chez la drosophile, existent en trois ou
quatre exemplaires chez les vertébrés (Desert HedgeHog, Indian HedgeHog et Sonic HedgeHog) pour des
fonctions similaires.

Approche évo-dévo : évolution et développement [modifier]

Depuis les années 1980, les travaux visant à relier la génétique et l'embryologie apportent un nouvel
éclairage sur les mécanismes de l'évolution biologique. Ces nouvelles approches réactualisent certains
aspects de théories plus anciennes comme la théorie du monstre prometteur de Goldschmidt.
des gènes dans la construction de l'individu (l'ontogenèse) et dans les réorganisations anatomiques qui se
produisent au cours de l'histoire évolutive (la phylogenèse). En plus de confirmer l'ascendance commune
entre les espèces vivantes, ces découvertes montrent que de petites modifications dans la séquence
génétique au niveau de ces gènes voire une simple différence dans la période pendant laquelle de tels
gènes sont exprimés au cours de l'embryogenèse, peuvent avoir des effets très importants sur la
morphologie de l'organisme. Cela montre que l'évolution ne consiste pas seulement à créer de nouveaux
gènes par mutation mais aussi à changer la façon dont les gènes sont exprimés.
Parmi les autres découvertes de la biologie moléculaire, les mécanismes épigénétiques constituent
une autre voie par laquelle de l'information peut être transmise "à côté" du seul code génétique. Ce type de
transmission peut impliquer l'ADN, l'ARN ou les protéines ou les organites du cytoplasme cellulaire et peut
être réversible ou non. L'un des processus épigénétiques les plus étudiés est la méthylation de l'ADN et son
lien avec les modifications de la chromatine : ce mécanisme consiste en l'inactivation d'un gène par
changement de la conformation de la chromatine du chromosome qui le porte. Ce changement rend le gène
silencieux sans en affecter la séquence nucléotidique. Par conséquent, ce gène sera transmis "intact" mais
silencieux aux générations suivantes. On pourra donc observer la résurgence d'un caractère ancestral plus
tard dans la lignée simplement par réactivation du gène (via une reconformation de la chromatine,
déclenchée par un évènement stressant dans l'environnement). L'importance de ces mécanismes
épigénétiques est encore difficile à mesurer mais ils peuvent avoir des conséquences importantes dans
l'explication de phénomènes qui ne respectent pas les lois de l'hérédité mendélienne.
Controverses actuelles [modifier]
Adaptationnisme et Neutralisme [modifier]
Au sein de la communauté scientifique, le débat concernant les théories modernes de l'évolution se
porte donc sur le rôle que joue le "hasard" (la dérive génétique (?!), produisant la diversité des caractères)
par rapport à la sélection naturelle. Les théories neutralistes ont ainsi été critiquées par des biologistes
tenant d'une vision adaptationniste de l'évolution pour lesquels les pressions de sélection (naturelle et
sexuelle) sont les principales forces dans l'évolution des espèces, le "hasard" ne jouant finalement que peu
de rôle.
• Argumentation : Les adaptationnistes s'appuient notamment sur les phénomènes de
convergence évolutive par lesquels différentes espèces aboutissent à la même solution évolutive
lorsqu'elles sont soumises au même problème adaptatif[10].
La forme fuselée de l'ichthyosaure, de la baleine ou du manchot qui ont la forme des poissons
témoignent de la force de la pression de la sélection naturelle qui plaide en faveur des adaptationnistes.
Les ichthyosaures étaient des Les manchots à jugulaire
Le hareng est un poisson osseux.
reptiles. des oiseaux aquatiques.
 Article détaillé : théorie neutraliste de l'évolution.
La Théorie Synthétique de l'Evolution s'impose dans le monde scientifique comme théorie de
référence. Dans les années 1970, elle connait d'importants développements théoriques, parfois iconoclastes.
Par exemple, la théorie neutraliste de l'évolution moléculaire proposée par le généticien japonais Motoo
Kimura en 1970, repose sur l'idée que l'évolution, au niveau des gènes, doit au moins autant sinon plus au
hasard (c'est-à-dire à une dérive) qu'à la sélection naturelle.
• Argumentation : La plupart des mutations ne sont pas adaptatives. Une énorme partie des
mutations est neutre ne procurant ni avantage, ni handicap, et une autre partie importante est
handicapante. Les mutations procurant un avantage sont quasiment inexistantes. Cela est
l'argument fort de la théorie neutraliste qui veut que le hasard ou la dérive génétique soit presque
l'unique clé de l'évolution. La sélection naturelle ne jouerait qu'un rôle marginal selon la théorie
neutraliste.
 • Il est tout aussi dangereux d'adopter le seul point de vue « adaptationniste », selon lequel
chaque caractère d'un être vivant pris isolément aurait été « façonné » par la sélection naturelle.
Dans cette optique, tout devrait avoir une utilité adaptative, y compris la couleur des yeux, le
chiasme optique et l'appendice.
L'expérience montre en fait qu'il existe beaucoup de caractères neutres en ce qui concerne la
descendance. Certains gènes sont par ailleurs pléiotropes - c'est-à-dire qu'ils commandent plusieurs
caractères à la fois. Une mutation avantageuse entraîne donc avec elle d'autres caractères ; quand
la mutation se répandra dans la population, ses « effets secondaires » se répandront en même
temps, sans avoir d'utilité en eux-mêmes... en tout cas à ce moment-là. Un exemple flagrant est celui
de la drépanocytose, une maladie héréditaire, qui perdure parce qu'elle confère une certaine
résistance au paludisme. La mutation ayant entrainé son apparition n'est pas neutre, mais la grave
maladie qu'elle entraine est en quelque sorte compensée par un avantage sélectif, vis-à-vis d'une
autre maladie.

Equilibres ponctués [modifier]

Article détaillé : équilibre ponctué.
 Podarcis sicula. Des lézards de l'espèce p. Sicula déposés sur l'ile de Prod Mrcaru en 1971 ont
évolué en 36 ans de sorte à disposer d'un nouvel organe de digestion absent chez l'espèce d'origine : les
valves sécales.
En 1972, S. J. Gould et Niles Eldredge publient leur théorie des équilibres ponctués destinée à
combler ce qui est pour eux une faille de la TSE : les paléontologues (comme Simpson) affirmaient en effet
que l'on devrait voir les espèces se transformer toujours très graduellement, et que l'on devrait donc trouver
des fossiles correspondant à toutes les étapes de la spéciation[11].
Cette thèse baptisée gradualisme phylétique peut se résumer sous l'expression répétée à plusieurs
reprises par Darwin dans son Origine des espèces : "Natura non facit saltum" (La Nature ne fait pas de saut).
A l'inverse, certains biologistes (comme Ernst Mayr) pensaient que les apparitions de nouvelles espèces
correspondaient à des évènements rares et ponctuels, que l'on peut représenter sous forme d'un escalier,
les paliers étant pour les périodes où une population ne subit aucun changement et les sauts correspondant
à une modification rapide des caractéristiques qui aboutissent sur quelques générations à une nouvelle
espèce.
Pour Gould et Eldredge, ce sont les biologistes qui ont raison ; les paléontologues auraient en fait
mal interprété les fossiles. Le débat sera néanmoins ravivé avec la critique des paléontologues Simon
Conway-Morris, Derek Briggs et Richard Fortey qui contredisent les interprétations de Gould sur les schistes
de Burgess censés illustrer la thèse des équilibres ponctués. Leur analyse plus précise de ces fossiles
montre certes une grande diversité anatomiques mais pas de quoi conclure qu'il y aurait là les traces d'un
"saut" au moment de l'explosion cambrienne avec de nombreuses lignées qui auraient brutalement disparu
du monde vivant tel que le prétendait Gould.
• Argumentation : Récemment, un cas d'évolution ultra-rapide a été enregistré[12].

Néo-Lamarckisme [modifier]
• Lamarck et Charles Darwin décrivaient l'hérédité des caractères acquis (bien qu'ils
n'emploient pas cette expression, la distinction entre caractères acquis et innés date de la fin du
XIXe siècle). Darwin mentionne même dans la première édition de L'Origine des espèces les « effets
cumulatifs du dressage » de génération en génération chez des chiens d'arrêt (pointers). Darwin est
en réalité le premier à proposer une théorie des caractères acquis, sous le nom de "théorie de la
pangenèse", dans son ouvrage De la variation des animaux et des plantes sous l'action de la
domestication. C'est August Weismann qui fera la démonstration, non de l'impossibilité de la
transmission des caractères acquis, mais de l'impossibilité de la transmission des mutilations,
 soutenue à la fin du XIXe siècle par les néo-lamarckiens. La théorie de Weismann exclut la
transmission des caractères acquis de manière purement théorique. La théorie synthétique postule
aujourd'hui que les gènes ne peuvent être modifiés naturellement que par des mutations aléatoires,
mais il s'agit, de l'aveu même de Francis Crick, d'un "dogme" de la biologie moléculaire.
[réf. nécessaire]
• On croyait en conséquence, jusqu'à la fin du XXe siècle, que l'adaptation individuelle ne se
transmettait pas. Toutefois, au début du XXIe siècle, de nouvelles expériences et observations ont
rouvert la porte à l'hypothèse d'une transmission de l'adaptation individuelle dans certains cas
(notamment la taille, par rapport aux conditions d'alimentation), non par la modification des gènes,
mais par la modification de leurs conditions d'expression, et, par là, de leur niveau d'activité, avec
toutes les conséquences. Ces phénomènes sont qualifiés d'épigénétique. En outre, on a découvert
que si les mutations sont bien aléatoires, les probabilités qu'elles soient réparées sont dépendantes
des conditions extérieures : les mécanismes d'auto-réparation et d'entretien de l'intégrité du génome
sont sous contrôle de l'état des cellules, ce qui fait que les cellules laissent passer plus ou moins de
mutations selon leur adaptation à leur milieu. Au final, l'adaptation acquise apparait bien comme un
élément de l'hérédité.
Une autre approche de l'évolution est le néo-lamarckisme. L’action en faveur de Lamarck se traduit
par l’organisation d’un congrès international à Paris en 1947 sous l’égide du CNRS sous le thème de
« paléontologie et transformisme » dont les actes paraissent en 1950 chez Albin Michel. Il réunit nombre des
plus grandes autorités françaises ; outre Grassé, y participèrent Lucien Cuénot (1866-1951), Pierre Teilhard
de Chardin (1881-1955), Maurice Caullery..[13]. Ceux-ci s’étaient alors opposés à des tenants du néo-
darwinisme, brillants biologistes également, comme John Burdon Sanderson Haldane (1892-1964) et
George Gaylord Simpson (1902-1984)[14]. Grassé fait l’éloge de Lamarck d’autres façons comme dans un
article de l’Encyclopædia Universalis en affirmant que Lamarck a été injustement calomnié et qu’il doit être
réhabilité[15].
• Argumentation : La transmission de caractères acquis à la descendance, pourrait passer par
un support épigénétique selon de récentes études[16].

Accroissement de la complexité au cours de l'évolution [modifier]

Article connexe : Modèle évolutif r/K.
Jean-Baptiste Lamarck est le premier a voir proposé une théorie de l'évolution où une tendance à la
complexification des êtres vivants sous l'effet de leur dynamique interne joue un rôle important. Charles
Darwin semble avoir rejeté cette idée parce qu'il ne voulait pas de l' ordre naturel qu'elle impliquait et qu'il
assimilait abusivement à une intervention divine[17]. Cette confusion semble avoir perduré jusque chez les
évolutionnistes actuel, comme en témoignent les difficultés autour de cette question.
Il reste difficile de mesurer la complexité dans le monde vivant. Elle est souvent estimée par le
nombre de types cellulaires identifiables chez un organisme, mais cette estimation n'est pas, ou peu,
corrélée à la taille du génome ou au nombre de gènes (complexité du génome) par exemple et ne saurait en
aucun cas être une mesure indiscutable.[réf. nécessaire] Ainsi, le génome humain comprend 3 milliards de
bases, le blé 12 milliards. Richard Dawkins signale également que les animaux à sang froid ont un génome
plus long que des animaux de taille comparable à sang chaud.
Rapporté à la quantité d'êtres vivants, l'accroissement de la complexité semble être un phénomène
marginal dans l'évolution. En effet, la plupart des espèces sont encore au stade unicellulaire (plus de 99 %
des êtres vivants sont des bactéries[18]), seules quelques lignées ont évolué vers une complexification.
 La sélection naturelle a tout de même favorisé, dans de nombreux cas, des organismes complexes.
Chez les mammifères, on observe une tendance à l'accroissement de la taille du cerveau, et des
comportements. On l'explique généralement par une « course aux armements » entre proies et prédateurs,
ou à une co-évolution antagoniste, à cause d'une compétition perpétuelle dans les populations (voir la
théorie de la reine rouge). Un système nerveux plus élaboré peut sauver la vie de l'animal. De même, des
« fédérations de cellules », ébauches d'organismes pluricellulaires, ont pu dans certaines circonstances être
plus efficaces que les bactéries isolées pour leur permettre de survivre. Cependant, il est plausible que
l'accroissement de la complexité ait pu, dans certains cas, correspondre à un caractère neutre et être fixé par
la dérive génétique[réf. nécessaire].
Si on parcourt l'histoire évolutive de l'ensemble du monde vivant on assiste à un accroissement de la
complexité dans de nombreuses lignées: insecte, mammifères. Cette complexité semble liée à une meilleure
adaptation des populations à un environnement fluctuant. En effet dans les espèces simples (bactéries) les
effectifs des populations varient au gré des variations des conditions, ce qui est moins le cas pour les
espèces plus complexes (humains, insectes sociaux). Par exemple, un organisme volumineux résiste mieux
aux changements de température. Un comportement élaboré permet d'effectuer des réserves de nourriture,
de prévoir les saisons.
Cependant, comme le rappelle Stephen Jay Gould là encore ( op.cit.), le phénomène est biaisé en
raison de ce qu'il nomme le mur de complexité minimale en dessous duquel on ne peut plus concevoir
d'organisme viable et pouvant se reproduire. Le génome peut donc se complexifier autant que le lui
permettent le temps et les ressources de l'environnement, mais n'a pas la latitude de se simplifier en de ça
de ce "mur". On est dans le phénomène bien connu en probabilités de marche au hasard avec une barrière
absorbante pour les individus, et de ce fait réfléchissante pour l'espèce : ne sera préservé que ce qui ne la
franchit pas.
Remises en question de la théorie [modifier]
Article détaillé : Historique des critiques des théories de l'évolution.
Les paléoanthropologues mettent en doute certains des aspects de cette théorie. Yves Coppens et
Pascal Picq vont, entre autres, tenter de démontrer que l'évolution biologique a précédé l'évolution culturelle,
dans le cas de notre espèce, et qu'en ce sens " Le développement technique et culturel dépasse le
développement biologique"; c'est-à-dire que les explications biologiques des comportements culturels
humains sont moins significatives que les explications sociales. Ils parleront aussi de "sélection culturelle",
en opposition à la sélection naturelle supposément s'opérant. Une des thèses les plus importantes sur le
sujet de l'explication biologique des comportements sociaux humains provient de E.O. Wilson, fondateur de
la sociobiologie.

Notes et références [modifier]

1. ↑ Source David, P. & Samadi, S. (2000) La Théorie de l'évolution, Flammarion.
2. ↑ Par exemple Aristote dans son Histoire des animaux [archive]
3. ↑ Guillaume Lecointre, Hervé Le Guyader, Classification phylogénétique du vivant, éd. Belin,
2001, p. 43.
4. ↑ Génomes et évolution - Innovations génétiques [archive], CNRS, (page consultée le 2
juillet 2008).
5. ↑ Parenté et diversité des organismes [archive], Université Pierre-et-Marie-Curie, Paris,
(page consultée le 2 juillet 2008).
6. ↑ Neil Campbell & al., Biologie, éd. De Boeck, 2006, p. 516.
 7. ↑ Apparition de la vie [archive], École Supérieure de Physique et de Chimie Industrielle,
(page consultée le 2 juillet 2008).
8. ↑ Cyril Langlois, L'Évolution montrée par les fossiles [archive], École normale supérieure de
Lyon, (page consultée le 2 juillet 2008).
9. ↑ Charles Sullivan, Cameron Mcpherson Smith, Darwin : ce n'est pas une histoire de
singe [archive], Association française pour l'information scientifique, (page consultée le 26 avril
2008).
10.↑ Par exemple, les yeux des vertébrés et des céphalopodes sont très similaires alors même
que l'ancêtre commun de ces taxons ne possédait pas de tels yeux. Les yeux de ces deux taxons
sont donc des adaptations analogues au problème de la vision montrant, pour les adaptationnistes,
que, malgré la diversité des espèces, les contraintes adaptatives sont suffisamment fortes pour
contraindre la place laissée au "hasard". De même le degré de ressemblance entre le Thylacine qui
est un marsupial, et le chien qui est un placentaire est tel qu'il faut être un spécialiste pour
différencier leurs squelettes respectifs.
11.↑ Plusieurs scientifiques considèrent la théorie de la mutation rapide des espèces d'Hugo
De Vries comme la base de cette théorie saltationniste.
12.↑ En 2004, une équipe scientifique dirigée par Duncan Irschick et Anthony Herrel est allée
sur l'ile de Prod Mrcaru en mer Adriatique afin d'observer un lézard que l'on connait bien puisque très
présent en France sous le nom de lézard des ruines. Ce lézard avait été introduit en effet sur l'ile en
1971 par l'équipe du professeur Eviatar Nevo, un spécialiste de l'évolution. Celui-ci n'ayant pas pu
suivre son projet personnellement, les conflits yougoslaves ayant entrainé une interdiction de facto
de poursuite de l'expérience, c'est 36 années plus tard que, Duncan Irschick et Anthony Herrel
découvrirent comme Podarcis sicula avait évolué de façon fulgurante, confirmant de façon frappante
l'hypothèse des équilibres ponctués. Le lézard avait grandi, sa mâchoire devenue plus puissante, et
 il avait changé de régime alimentaire, d'insectivore il était devenu herbivore, et des valves sécales
étaient apparues au niveau des intestins pour l'aider à digérer les herbes... Voir site : [pdf]
(en)PNAS, vol. 105, n°12, pages 4792-4795 [archive] (25 mars 2008).
13.↑ cf. Blanc (1990) : 238.
14.↑ cf. Blanc (1990) : 11.
15.↑ cf. Lestienne et Lapidus (2000) : 45.
16.↑ Par exemple, L’obésité serait non pas uniquement un effet direct touchant les individus
atteints eux-mêmes mais également un effet transgénérationnel. Des données chez l'homme et chez
l'animal semblent montrer que les effets d'une sous-alimentation subies par des individus pourraient
en effet être transmises aux descendants. Des modifications épigénétiques (n’altérant donc pas
l'ADN) n’étant pas effacées lors du passage par la lignée germinale sont désormais décrites chez
plusieurs espèces. En fait, chez les plantes il existe une corrélation entre le niveau d'expression d'un
gène et sa méthylation. Pareillement, chez les mammifères nous témoignons de la méthylation d'une
séquence transposable qui est insérée à proximité d'un gène particulier. Le degré de méthylation
d'un transposon pouvant enfin moduler l'expression du gène dans lequel il s'est inséré. (Revue :
Cahiers de nutrition et de diététique ISSN 0007-9960 CODEN CNDQA8 ; Source : 2002, vol. 37,
no4, pp. 261-272 ; Editeur : Masson, Paris, FRANCE (1966) ; Site : CAT.INIST du CNRS [archive]).
17.↑ Cf. André Pichot, Histoire de la notion de vie, éd. Gallimard, coll. TEL, 1993, pp. 820-838.
18.↑ Stephen, Jay Gould rappelle d'ailleurs, dans La vie est belle, que la biomasse bactérienne
sur Terre est supérieure à celle des mammifères

Voir aussi [modifier]

 [Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

 Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Th%C3%A9orie_synth%C3%A9tique_de_l
%27%C3%A9volution ».
Catégorie : Théorie sur l'évolution | [+]
 W000

Evolution as theory and fact

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Biology portal · v • d • e
 The statement "evolution is both a theory and a fact" is often seen in biological literature.[1][2][3][4][5]
[6][7] Evolution is a "theory" in the scientific sense of the term "theory"; it is an established scientific model
that explains observations and makes predictions through mechanisms such as natural selection.
When scientists say "evolution is a fact" they are using one of two meanings of the word "fact". One
meaning is empirical, and when this is what scientists mean, then "evolution" is used to mean observed
changes in allele frequencies or traits of a population over successive generations.
Another way "fact" is used is to refer to a certain kind of theory, one that has been so powerful and
productive for such a long time that it is universally accepted by scientists. When scientists say evolution is a
fact in this sense, they mean it is a fact that all living organisms have descended from a common ancestor (or
ancestral gene pool) [8] even though this cannot be directly observed. This implies more tangibly that it is a
fact that humans share a common ancestor with other primates.
Contents
[hide]
• 1 Evolution, fact and theory
• 1.1 Evolution
• 1.2 Fact
• 1.3 Theory
• 2 Evolution compared with gravity
• 3 Evolution as theory and fact in the literature
• 3.1 Evolution as fact and theory
• 3.2 Evolution as fact not theory
• 4 Predictive power
• 5 Related concepts and terminology
• 6 See also
• 7 Notes
• 8 References
• 9 External links
[edit] Evolution, fact and theory
Evolution has been described as "fact and theory", "fact not theory", and "only a theory, not a fact".
This illustrates a confusion in terminology that hampers discussion.[9][10] The meanings of the terms
"evolution", "fact", and "theory" are described below.

[edit] Evolution
Main articles: Introduction to evolution and Evolution
Evolution is usually defined simply as changes in trait or gene frequency in a population of organisms
from one generation to the next. However, "evolution" is often used to include the following additional claims:
1. Differences in trait composition between isolated populations over many generations may
result in the origin of new species.
2. All living organisms alive today have descended from a common ancestor (or ancestral gene
pool).
According to Douglas Futuyma:
Biological evolution may be slight or substantial; it embraces everything from slight changes in
the proportion of different alleles within a population (such as those determining blood types) to
the successive alterations that led from the earliest proto-organism to snails, bees, giraffes, and
dandelions.[11]

The term "evolution", especially when referred to as a "theory", is also used more broadly to
incorporate processes such as natural selection and genetic drift.
[edit] Fact
Main article: Scientific fact
Fact is often used by scientists to refer to experimental or empirical data or objective verifiable
observations.[12][13][14][15] "Fact" is also used in a wider sense to mean any theory for which there is
overwhelming evidence.
A fact is a hypothesis that is so firmly supported by evidence that we assume it is true, and act
as if it were true. —Douglas Futyuma[16]
Evolution is a fact in the sense that it is overwhelmingly validated by the evidence. Frequently,
evolution is said to be a fact in the same way as the Earth revolving around the Sun is a fact.[16][17] The
following quotation from H. J. Muller, "One Hundred Years Without Darwin Are Enough" explains the point.
There is no sharp line between speculation, hypothesis, theory, principle, and fact, but only a
difference along a sliding scale, in the degree of probability of the idea. When we say a thing is a fact,
then, we only mean that its probability is an extremely high one: so high that we are not bothered by
doubt about it and are ready to act accordingly. Now in this use of the term fact, the only proper one,
evolution is a fact.[3]
The National Academy of Science (U.S.) makes a similar point:
Scientists most often use the word "fact" to describe an observation. But scientists can also
use fact to mean something that has been tested or observed so many times that there is no longer a
compelling reason to keep testing or looking for examples. The occurrence of evolution in this sense is
fact. Scientists no longer question whether descent with modification occurred because the evidence
is so strong.[18]
 Philosophers of science argue that we do not know anything with absolute certainty: even direct
observations may be "theory laden" and depend on assumptions about our senses and the measuring
instruments used. In this sense all facts are provisional.[1][19]

[edit] Theory
Main article: Scientific theory
A scientific theory is a well-supported body of interconnected statements that explains observations
and can be used to make testable predictions. Scientific theories describe the coherent framework into which
observable data fit. The "theory of evolution" is the framework that best explains observed changes of
species over time and best predicts the new observations that continue to be made in evolutionary biology
and related sciences.
The scientific definition of the word "theory" is different from the colloquial sense of the word.
Colloquially, "theory" can mean a conjecture, an opinion, or a speculation that does not have to be based on
facts or make testable predictions. In science, the meaning of theory is more rigorous: a theory must be
based on observed facts and make testable predictions.
In science, a current theory is a theory that has no equally acceptable or more acceptable alternative
theory, and has survived attempts at falsification. That is, there have been no observations made that
contradict it to this point and, indeed, every observation ever made either supports the current theory or at
least does not falsify it by contradicting it completely. A revision of the current theory, or the generation of a
new theory is necessary if new observations contradict the current theory, as the current findings are in need
of a new explanation (see scientific revolution or paradigm shift). However, the falsification of a theory does
not falsify the facts on which the theory is based.
[edit] Evolution compared with gravity
See also: Gravity and History of gravitational theory
The application of the terms "fact" and "theory" to evolution is similar to their use in describing
gravity.[1][20]
The most obvious fact of gravity is that objects in our everyday experience tend to fall downwards
when not otherwise prevented from doing so. People throughout history have wondered what causes this
effect. Many explanations have been proposed over the centuries. Aristotle, Galileo, Newton, and Einstein
have all developed useful models of gravity, each of which constitutes a theory of gravity. (Newton, for
example, realized that the fact of gravity can be extended to the tendency of any two masses to attract one
another.) The word "gravity", therefore, can be used to refer to the observed facts (i.e., that masses attract
one another) and the theory used to explain the facts (the reason why masses attract one another). In this
way, gravity is both a theory and a fact.
In the study of biological species, the facts include the existence of many different species in
existence today, some very similar to each other and some very dissimilar, the remains of extinct species in
the fossil record, and so forth. In species that rapidly reproduce, for example fruit flies, the process of change
from generation to generation — that is, evolutionary change — has been observed in the laboratory.[21] The
observation of fruit fly populations changing over time is also an example of a fact. So evolution is a fact just
as the observations of gravity are a fact.
There have been many attempts to explain these biological observations over the years.
Lamarckism, transmutationism and orthogenesis were all non-Darwinian theories that attempted to explain
the observations of species and fossils, as well as other evidence. However, the modern theory of evolution
is the explanation for all relevant observations regarding the development of life, based on a model that
explains all the available data and observations (and provides testable predictions). Thus, evolution is not
only a fact but also a theory, just as gravity is both a fact and a theory.

[edit] Evolution as theory and fact in the literature

The confusion over the word evolution and the distinction between "fact" and "theory" is largely due
to authors using evolution to refer to three related yet distinct ideas: first, the changes that occur within
species over generations; second, the mechanism thought to drive change; and third, the concept of
common descent. However, among biologists there is a consensus that evolution is a fact:
• American zoologist and paleontologist George Simpson stated that "Darwin... finally and
definitely established evolution as a fact." [22]
• H. J. Muller wrote, "If you like, then, I will grant you that in an absolute sense evolution is not
a fact, or rather, that it is no more a fact than that you are hearing or reading these words." [3]
• Kenneth R. Miller writes, "evolution is as much a fact as anything we know in science." [23]
• Ernst Mayr observed, "The basic theory of evolution has been confirmed so completely that
most modern biologists consider evolution simply a fact. How else except by the word evolution can
we designate the sequence of faunas and floras in precisely dated geological strata? And
evolutionary change is also simply a fact owing to the changes in the content of gene pools from
generation to generation."[7]

[edit] Evolution as fact and theory

Commonly "fact" is used to refer to the observable changes in organisms' traits over generations
while the word "theory" is reserved for the mechanisms that cause these changes:
 • Paleontologist Stephen Jay Gould writes, "Evolution is a theory. It is also a fact. And facts
and theories are different things, not rungs in a hierarchy of increasing certainty. Facts are the
world's data. Theories are structures of ideas that explain and interpret facts. Facts do not go away
when scientists debate rival theories to explain them. Einstein's theory of gravitation replaced
Newton's, but apples did not suspend themselves in mid-air, pending the outcome. And humans
evolved from ape-like ancestors whether they did so by Darwin's proposed mechanism or by some
other yet to be discovered."[2]
• Similarly, biologist Richard Lenski says, "Scientific understanding requires both facts and
theories that can explain those facts in a coherent manner. Evolution, in this context, is both a fact
and a theory. It is an incontrovertible fact that organisms have changed, or evolved, during the
history of life on Earth. And biologists have identified and investigated mechanisms that can explain
the major patterns of change."[6]
• Biologist T. Ryan Gregory says, "biologists rarely make reference to 'the theory of evolution,'
referring instead simply to 'evolution' (i.e., the fact of descent with modification) or 'evolutionary
theory' (i.e., the increasingly sophisticated body of explanations for the fact of evolution). That
evolution is a theory in the proper scientific sense means that there is both a fact of evolution to be
explained and a well-supported mechanistic framework to account for it." [20]

[edit] Evolution as fact not theory

Other commentators, focusing on the changes in species over generations and in some cases
common ancestry have stressed that evolution is a fact to emphasize the weight of supporting evidence while
denying it is helpful to use the term "theory":
 • R. C. Lewontin wrote, "It is time for students of the evolutionary process, especially those
who have been misquoted and used by the creationists, to state clearly that evolution is a fact, not
theory."[24]
• Douglas Futuyma writes in his Evolutionary Biology book, "The statement that organisms
have descended with modifications from common ancestors—the historical reality of evolution—is not
a theory. It is a fact, as fully as the fact of the earth's revolution about the sun." [11]
• Richard Dawkins says, "One thing all real scientists agree upon is the fact of evolution itself.
It is a fact that we are cousins of gorillas, kangaroos, starfish, and bacteria. Evolution is as much a
fact as the heat of the sun. It is not a theory, and for pity's sake, let's stop confusing the
philosophically naive by calling it so. Evolution is a fact." [25]
• Neil Campbell wrote in his 1990 biology textbook, "Today, nearly all biologists acknowledge
that evolution is a fact. The term theory is no longer appropriate except when referring to the various
models that attempt to explain how life evolves... it is important to understand that the current
questions about how life evolves in no way implies any disagreement over the fact of evolution." [4]

[edit] Predictive power

A central tenet in science is that a scientific theory is supposed to have predictive power, and
verification of predictions are seen as an important and necessary support for the theory. The theory of
evolution has provided such predictions. Four examples are:
• Genetic information must be transmitted in a molecular way that will be almost exact but
permit slight changes. Since this prediction was made, biologists have discovered the existence of
DNA, which has a mutation rate of roughly 10−9 per nucleotide per cell division; this provides just
such a mechanism.[26]
• Some DNA sequences are shared by very different organisms. It has been predicted by the
theory of evolution that the differences in such DNA sequences between two organisms should
roughly resemble both the biological difference between them according to their anatomy and the
time that had passed since these two organisms have separated in the course of evolution, as seen
in fossil evidence. The rate of accumulating such changes should be low for some sequences,
namely those that code for critical RNA or proteins, and high for others that code for less critical RNA
or proteins; but for every specific sequence, the rate of change should be roughly constant over time.
These results have been experimentally confirmed. Two examples are DNA sequences coding for
rRNA, which is highly conserved, and DNA sequences coding for fibrinopeptides (amino acid chains
that are discarded during the formation of fibrin), which are highly non-conserved.[26]
• Prior to 2004, paleontologists had found fossils of amphibians with necks, ears, and four
legs, in rock no older than 365 million years old. In rocks more than 385 million years old they could
only find fish, without these amphibian characteristics. Evolutionary theory predicted that since
amphibians evolved from fish, an intermediate form should be found in rock dated between 365 and
385 million years ago. Such an intermediate form should have many fish-like characteristics,
conserved from 385 million years ago or more, but also have many amphibian characteristics as well.
In 2004, an expedition to islands in the Canadian arctic searching specifically for this fossil form in
rocks that were 375 million years old discovered fossils of Tiktaalik.[27]
• Evolutionary theory predicts that novel inventions can arise, while creationists predict that
new "information" cannot arise, and that the Second Law of Thermodynamics only allows for
"information" to be lost.[28] In an ongoing experiment, Richard Lenski observed that E. coli evolved
 the ability to metabolize citrate, which constitutes a novel invention, and an increase in the
information of the DNA of the E. coli.[29]

[edit] Related concepts and terminology

• Speculative or conjectural explanations are called hypotheses. Well-tested explanations are
called theories.
• "Fact" does not mean "absolute certainty". In the words of Stephen J. Gould: In science,
"fact" can only mean "confirmed to such a degree that it would be perverse to withhold provisional
assent."[2]
• "Proof" of a theory does not exist in natural sciences. Proof only exists in formal sciences,
such as mathematics. Experimental observation of the predictions made by a hypothesis or theory is
called validation.
• A scientific law is a concept related to a scientific theory. Very well-established "theories" that
rely on a simple principle are often called scientific "laws". For example, it is common to encounter
reference to "the law of gravity", "the law of natural selection", or the "laws of evolution."

[edit] See also

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Epistemology
 • Evidence of common descent
• Evolution is just a theory, not a fact (in objections to evolution)
• Misconceptions about evolution (in List of misconceptions)
• Theory vs. Fact (in Creation-evolution controversy)

[edit] Notes
1. ^ a b c Moran, Laurence (1993-01-22). "Evolution is a Fact and a Theory". Talk.origins.
http://www.talkorigins.org/faqs/evolution-fact.html. Retrieved 2007-10-18.
2. ^ a b c Gould, Stephen Jay (1981-05-01). "Evolution as Fact and Theory". Discover 2 (5): 34–
37. http://www.stephenjaygould.org/library/gould_fact-and-theory.html. Reprinted in:
• Vetter, Herbert F. (ed.) (1982). Speak Out Against The New Right. Beacon Press.
ISBN 0807004863.
• Gould, Stephen Jay (1994-04-01). Hen's Teeth and Horse's Toes. New York: Norton.
ISBN 0393017168.
3. ^ a b c Muller, H. J. (1959). "One hundred years without Darwin are enough". School Science
and Mathematics 59: 304–305. doi:10.1111/j.1949-8594.1959.tb08235.x.
http://www.skepticfiles.org/evolut/100pcnts.htm. Reprinted in:
• Zetterberg, Peter (ed.) (1983-05-01). Evolution Versus Creationism: The Public
Education Controversy. Phoenix AZ: ORYX Press. ISBN 0897740610.
4. ^ a b Campbell, Neil A.; Reece, Jane B. (2002-02-05). Biology 6th ed.. Benjamin Cummings.
p. 1175. ISBN 0805366245.
 5. ^ Dobzhansky, Theodosius (1973-03-01). "Nothing in biology makes sense except in the light
of evolution". American Biology Teacher 35. http://people.delphiforums.com/lordorman/light.htm.
Reprinted in:
• Zetterberg, Peter (ed.) (1983-05-01). Evolution Versus Creationism: The Public
Education Controversy. Phoenix AZ: ORYX Press. ISBN 0897740610.
6. ^ a b Lenski, Richard E. (2000). "Evolution: Fact and Theory". American Institute of Biological
Sciences. http://www.actionbioscience.org/evolution/lenski.html. Retrieved 2007-10-18.
7. ^ a b Mayr, Ernst (1988). Toward a New Philosophy of Biology: Observations of an
Evolutionist. Cambridge: Harvard University Press. ISBN 0-674-89666-1.
8. ^ Cavalier-Smith T (2006). "Cell evolution and Earth history: stasis and revolution". Philos
Trans R Soc Lond B Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842. PMID 16754610.
9. ^ Is "Evolution" a "Theory" or "Fact" or Is This Just a Trivial Game of Semantics? by Casey
Luskin
10.^ Committee for Skeptical Inquiry — Evolution & Creationism: Terminology in Conflict by
Richard Joltes
11.^ a b Futuyma, Douglas J. (1997). , Evolutionary Biology, 3rd ed.. Sinauer Associates. p. 751.
ISBN 0878931899.
12.^ Wordnet entry for phrase "scientific fact"
13.^ United States National Park Service Glossary
14.^ Webster's New Millennium Dictionary of English, Preview Edition (v 0.9.6) , Copyright ©
2003–2006 Lexico Publishing Group, LLC
15.^ Webster's Encyclopedic Unabridged Dictionary of the English Language (1996) gives a
third meaning of the word "fact" as (3) A truth known by actual experience or observation; something
known to be true: 'Scientists gather facts about plant growth.'
 16.^ a b Hypotheses, Facts, and the Nature of Science, —Douglas Futyuma
17.^ Guardian article by Richard Dawkins, Jerry Coyne
18.^ Science and Creationism: A View from the National Academy of Sciences, Second Edition
(1999), National Academy of Sciences (NAS), National Academy Press, Washington DC, 2006.
19.^ Wilkins, JS (1997). "Evolution and Philosophy:Is Evolution Science, and What Does
'Science' Mean?". The TalkOrigins Archive. http://www.talkorigins.org/faqs/evolphil/falsify.html.
Retrieved 2009-08-17.
20.^ a b Gregory, T. Ryan (2007). "Evolution as Fact, Theory, and Path". Evolution: Education
and Outreach 1 (1): 46–52. doi:10.1007/s12052-007-0001-z.
21.^ Dobzhansky T, Pavlovsky O (1971). "Experimentally created incipient species of
Drosophila". Nature 230 (5292): 289–92. doi:10.1038/230289a0. PMID 5549403.
22.^ Robinson, B.A. (2005-08-30). "Is the theory of evolution merely a "theory"?".
http://www.religioustolerance.org/ev_stat.htm. Retrieved 2007-10-18.
23.^ "Miller, Kenneth S. (2007). Finding Darwin's God: A Scientist's Search for Common Ground
Between God and Evolution (P.S.). New York, N.Y: Harper Perennial. ISBN 0061233501.
24.^ Lewontin, R. C. (1981). "Evolution/Creation Debate: a time for truth". Bioscience 31: 559.
Reprinted in:
• Zetterberg, Peter, ed. (1983-05-01). Evolution Versus Creationism: the public
education controversy. Phoenix AZ: Oryx Press. ISBN 0897740610.
25.^ Natural History article : The Illusion of Design by Richard Dawkins
26.^ a b Bruce Alberts; Alexander Johnson; Julian Lewis; Martin Raff; Keith Roberts; Peter
Walter (March, 2002). Molecular Biology of the Cell (4th ed.). Routledge. ISBN 0-8153-3218-1.
27.^ "Shubin, Neil. (2008). Your Inner Fish. Pantheon. ISBN 9780375424472.
28.^ TalkOrigins.org
 29.^ NS:bacteria make major evolutionary shift in the lab

[edit] References
• J.P. Franck, et al., "Evolution of a satellite DNA family in tilapia." Annual Meeting Canadian
Federation of Biological Societies. Halifax, (1990).
• M. Losseau-Hoebeke, "The biology of four haplochromine species of Lake Kivu (Zaire) with
evolutionary implications." Thesis, Dept. Ichthyology, Rhodes University, Grahamstown, (1992).

[edit] External links

• Not Just a Theory Discredits the assertion that evolution is "just a theory", with an
explanation of the meaning of the word 'theory' in a scientific context.
• Talk Origins Response to the claim that no examples of speciation have been observed.
• Glenn Branch; Louise S. Mead (2008-06-06). "“Theory” in Theory and Practice" (pdf). Evo
Edu Outreach (2008) 1:287–289. Springer Science + Business Media.
http://www.springerlink.com/content/fr258627q2x3t378/fulltext.pdf. Retrieved 2008-07-21.
Retrieved from "http://en.wikipedia.org/wiki/Evolution_as_theory_and_fact"

Categories: Creationist objections to evolution | Biology theories | Evolutionary biology

• This page was last modified on 18 August 2010 at 16:24.

 w000

Timeline of evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

For the history of evolutionary biology, see History of evolutionary thought.
This article is about the timeline of evolution in a general aspect. See also the timeline of human
evolution.
This timeline of the evolution of life outlines the major events in the development of life on the planet
Earth (See Organism). For a thorough explanatory context, see the history of Earth, and geologic time scale.
The dates given in this article are estimates based on scientific evidence.
In biology, evolution is the process by which populations of organisms acquire and pass on novel
traits from generation to generation. Its occurrence over large stretches of time explains the origin of new
species and ultimately the vast diversity of the biological world. Contemporary species are related to each
other through common descent, products of evolution and speciation over billions of years.

Contents
[hide]
• 1 Basic timeline
• 2 Detailed timeline
• 2.1 Hadean Eon
• 2.2 Archean Eon
• 2.3 Proterozoic Eon
• 2.4 Phanerozoic Eon
• 2.4.1 Paleozoic Era
• 2.4.2 Mesozoic Era
• 2.4.3 Cenozoic Era
• 3 See also
• 4 Further reading
• 5 References
• 6 External links

[edit] Basic timeline

 Life on Earth
view • discuss • edit
-4500 —
–
-4000 —
–
-3500 —
–
-3000 —
–
-2500 —
–
-2000 —
–
-1500 —
–
-1000 —
–
-500 —
–
0—
Life?
 Photosynthesis?
Eukaryotes
Complex multicellular life
Animals
Land plants
"Dinosaurs"
Mammals
Flowers

←
Formation
of Earth

←
Meteorite bombardment

←
Atmospheric oxygen

←
Ediacara biota
 ←
Cambrian explosion

←
Modern-looking
humans

Axis scale: millions of years ago.

The basic timeline is a 4.5 billion year old Earth, with (very approximate) dates:
• 3.8 billion years of simple cells (prokaryotes),
• 3 billion years of photosynthesis,
• 2 billion years of complex cells (eukaryotes),
• 1 billion years of multicellular life,
• 600 million years of simple animals,
• 570 million years of arthropods (ancestors of insects, arachnids and crustaceans),
• 550 million years of complex animals,
• 500 million years of fish and proto-amphibians,
• 475 million years of land plants,
• 400 million years of insects and seeds,
• 360 million years of amphibians,
• 300 million years of reptiles,
• 200 million years of mammals,
 • 150 million years of birds,
• 130 million years of flowers,
• 65 million years since the non-avian dinosaurs died out,
• 2.5 million years since the appearance of the genus Homo,
• 200,000 years since humans started looking like they do today,
• 25,000 years since Neanderthals died out.

[edit] Detailed timeline

Note that Ma, megaannum, means "million years ago".
Also note that Ka means "thousand years ago" and Ya means "years ago"

[edit] Hadean Eon

3800 Ma and earlier.
Date Event

4600 Ma The planet Earth forms from the accretion disc

revolving around the young Sun.

4500 Ma According to one plausible theory, the planet Earth

and the planet Theia collide, sending countless moonlets into
orbit around the young Earth. These moonlets eventually
 coalesce to form the Moon.[1] The gravitational pull of the
new Moon stabilises the Earth's fluctuating axis of rotation
and sets up the conditions in which life formed.[2]

4100 Ma The surface of the Earth cools enough for the crust to
solidify. The atmosphere and the oceans form.[3] PAH infall,
[4] and iron sulfide synthesis along deep ocean platelet
boundaries, may have led to the RNA world of competing
organic compounds.

Between 4500 and 3500 Ma The earliest life appears, possibly derived from self-
reproducing RNA molecules.[5][6] The replication of these
organisms requires resources like energy, space, and smaller
building blocks, which soon become limited, resulting in
competition, with natural selection favouring those molecules
which are more efficient at replication. DNA molecules then
take over as the main replicators and these archaic genomes
soon develop inside enclosing membranes which provide a
stable physical and chemical environment conducive to their
replication: proto-cells.[7][8][9]

3900 Ma Late Heavy Bombardment: peak rate of impact

events upon the inner planets by meteors. This constant
disturbance may have obliterated any life that had evolved to
that point, or possibly not, as some early microbes could have
 survived in hydrothermal vents below the Earth's surface;[10]
or life might have been transported to Earth by a meteor.[11]

Somewhere between 3900 - 2500 Ma Cells resembling prokaryotes appear.[12] These first
organisms are chemoautotrophs: they use carbon dioxide as
a carbon source and oxidize inorganic materials to extract
energy. Later, prokaryotes evolve glycolysis, a set of
chemical reactions that free the energy of organic molecules
such as glucose and store it in the chemical bonds of ATP.
Glycolysis (and ATP) continue to be used in almost all
organisms, unchanged, to this day.[13][14]

[edit] Archean Eon

3800 Ma – 2500 Ma
Date Event

3500 Ma Lifetime of the last universal ancestor;[15][16] the split between bacteria and
archaea occurs.[17]
Bacteria develop primitive forms of photosynthesis which at first do not produce
oxygen.[18] These organisms generate ATP by exploiting a proton gradient, a mechanism
still used in virtually all organisms.
 3000 Ma Photosynthesizing cyanobacteria evolve; they use water as a reducing agent,
thereby producing oxygen as waste product.[19] The oxygen initially oxidizes dissolved
iron in the oceans, creating iron ore. The oxygen concentration in the atmosphere
subsequently rises, acting as a poison for many bacteria. The moon is still very close to
the earth and causes tides 1,000 feet (305 m) high. The earth is continually wracked by
hurricane force winds. These extreme mixing influences are thought to stimulate
evolutionary processes. (See Oxygen catastrophe)

[edit] Proterozoic Eon

2500 Ma – 542 Ma
Date Event

By 1850 Ma Eukaryotic cells appear.[20][21] Eukaryotes contain membrane-bound

organelles with diverse functions, probably derived from prokaryotes engulfing each
other via phagocytosis. (See Endosymbiosis)

By 1200 Ma Sexual reproduction first appears, increasing the rate of evolution.[22]

1200 Ma Simple multicellular organisms evolve, mostly consisting of cell colonies of

limited complexity.

850–630 Ma A global glaciation may have occurred.[23][24] Opinion is divided on whether it

 increased or decreased biodiversity or the rate of evolution.[25][26][27]

580–542 Ma The Ediacaran biota represent the first large, complex multicellular organisms -
although their affinities remain a subject of debate.[28]

580–500 Ma Most modern phyla of animals begin to appear in the fossil record during the
Cambrian explosion.[29][30]

580–540 Ma The accumulation of atmospheric oxygen allows the formation of an ozone

layer.[31] This blocks ultraviolet radiation, permitting the colonisation of the land.[31]

560 Ma Earliest fungi

[edit] Phanerozoic Eon

542 Ma – present
The Phanerozoic Eon, literally the "period of well-displayed life", marks the appearance in the fossil
record of abundant, shell-forming and/or trace-making organisms. It is subdivided into three eras, the
Paleozoic, Mesozoic and Cenozoic, which are divided by major mass extinctions.

[edit] Paleozoic Era

542 Ma – 251.0 Ma
Date Event
535 Ma Major diversification of living things in the oceans: chordates, arthropods (e.g.
trilobites, crustaceans), echinoderms, mollusks, brachiopods, foraminifers and
radiolarians, etc.

530 Ma The first known footprints on land date to 530 Ma, indicating that early animal
explorations may have predated the development of terrestrial plants.[32]

525 Ma Earliest graptolites.

510 Ma First cephalopods (Nautiloids) and chitons.

505 Ma Fossilization of the Burgess Shale.

485 Ma First vertebrates with true bones (jawless fishes).

450 Ma Land arthropod burrows (millipedes) appear, along with the first complete
conodonts and echinoids.

440 Ma First agnathan fishes: Heterostraci, Galeaspida, and Pituriaspida.

434 Ma The first primitive plants move onto land,[33][citation needed] having evolved from
green algae living along the edges of lakes.[34] They are accompanied by fungi, which
may have aided the colonisation of land through symbiosis.
420 Ma Earliest ray-finned fishes, trigonotarbid arachnids, and land scorpions.

410 Ma First signs of teeth in fish. Earliest nautiid nautiloids, lycophytes, and
trimerophytes.

395 Ma First lichens, stoneworts. Earliest harvestman, mites, hexapods (springtails), and
ammonoids.

363 Ma By the start of the Carboniferous Period, the Earth begins to be recognisable.
Insects roamed the land and would soon take to the skies; sharks swam the oceans as
top predators,[35] and vegetation covered the land, with seed-bearing plants and forests
soon to flourish.
Four-limbed tetrapods gradually gain adaptations which will help them occupy a
terrestrial life-habit.

360 Ma First crabs and ferns. Land flora dominated by seed ferns.

350 Ma First large sharks, ratfishes, and hagfish.

340 Ma Diversification of amphibians.

330 Ma First amniote vertebrates (Paleothyris).

305 Ma Earliest diapsid reptiles (e.g. Petrolacosaurus).

280 Ma Earliest beetles, seed plants and conifers diversify while lepidodendrids and
 sphenopsids decrease. Terrestrial temnospondyl amphibians and pelycosaurs (e.g.
Dimetrodon) diversify in species.
251.4 Ma The Permian-Triassic extinction event eliminates over 90-95% of marine species.
Terrestrial organisms were not as seriously affected as the marine biota. This "clearing of
the slate" may have led to an ensuing diversification, but life on land took 30M years to
completely recover.[36]

[edit] Mesozoic Era

Date Event

From 251.4 Ma The Mesozoic Marine Revolution begins: increasingly well-adapted and
diverse predators pressurise sessile marine groups; the "balance of power" in the
oceans shifts dramatically as some groups of prey adapt more rapidly and
effectively than others.

245 Ma Earliest ichthyosaurs.

240 Ma Increase in diversity of gomphodont cynodonts and rhynchosaurs.

225 Ma Earliest dinosaurs (prosauropods), first cardiid bivalves, diversity in cycads,

bennettitaleans, and conifers. First teleost fishes.

215 Ma First mammals (e.g. Eozostrodon), minor vertebrate extinctions occur

220 Ma

Eoraptor, among the earliest dinosaurs, appeared in the fossil record 230
million years ago.
Gymnosperm forests dominate the land; herbivores grow to huge sizes in
order to accommodate the large guts necessary to digest the nutrient-poor plants.
[citation needed], first flies and turtles (Odontochelys). First Coelophysoid dinosaurs

200 Ma The first accepted evidence for viruses (at least, the group Geminiviridae)
exists.[37] Viruses are still poorly understood and may have arisen before "life"
itself, or may be a more recent phenomenon.
Major extinctions in terrestrial vertebrates and large amphibians. Earliest
examples of Ankylosaurian dinosaurs

195 Ma First pterosaurs with specialized feeding (Dorygnathus). First sauropod

dinosaurs. Diversification in small, ornithischian dinosaurs: heterodontosaurids,
fabrosaurids, and scelidosaurids.
190 Ma Pliosaurs appear in the fossil record. First lepidopteran insects
(Archaeolepis), hermit crabs, modern starfish, irregular echinoids, corbulid bivalves,
and tubulipore bryozoans. Extensive development of sponge reefs.

176 Ma First members of the Stegosauria group of dinosaurs

170 Ma Earliest salamanders, newts, cryptoclidid & elasmosaurid plesiosaurs, and

cladotherian mammals. Cynodonts become extinct while sauropod dinosaurs
diversify.

165 Ma First rays and glycymeridid bivalves.

161 Ma Ceratopsian dinosaurs appear in the fossil record (Yinlong)

155 Ma First blood-sucking insects (ceratopogonids), rudist bivalves, and

cheilosome bryozoans. Archaeopteryx, a possible ancestor to the birds, appears in
the fossil record, along with triconodontid and symmetrodont mammals. Diversity in
stegosaurian and theropod dinosaurs.

130 Ma The rise of the Angiosperms: These flowering plants boast structures that
attract insects and other animals to spread pollen. This innovation causes a major
burst of animal evolution through co-evolution. First freshwater pelomedusid turtles.

115 Ma First monotreme mammals.

 110 Ma First hesperornithes, toothed diving birds. Earliest limopsid, verticordiid, and
thyasirid bivalves.

106 Ma Spinosaurus, the largest theropod dinosaur, appears in the fossil record.
100 Ma Earliest bees.

90 Ma Extinction of ichthyosaurs. Earliest snakes and nuculanid bivalves. Large

diversification in angiosperms: magnoliids, rosids, hamamelidids, monocots, and
ginger. Earliest examples of ticks.

80 Ma First ants and termites.

70 Ma Multituberculate mammals increase in diversity. First yoldiid bivalves.

68 Ma Tyrannosaurus, the largest terrestrial predator of North America appears in

the fossil record. First species of Triceratops.

[edit] Cenozoic Era

65.5 Ma – present
Date Event
 65.5 Ma The Cretaceous–Tertiary extinction event eradicates about half of all animal
species, including mosasaurs, pterosaurs, plesiosaurs, ammonites, belemnites, rudist
and inoceramid bivalves, most planktic foraminifers, and all of the dinosaurs excluding
their descendants the birds [38]

From 65 Ma Rapid dominance of conifers and ginkgos in high latitudes, along with
mammals becoming the dominant species. First psammobiid bivalves. Rapid
diversification in ants.

63 Ma Evolution of the creodonts, an important group of carnivorous mammals.

60 Ma Diversification of large, flightless birds. Earliest true primates, along with the
first semelid bivalves, edentates, carnivorous and lipotyphlan mammals, and owls. The
ancestors of the carnivorous mammals (miacids) were alive.

56 Ma Gastornis, a large, flightless bird appears in the fossil record, becoming an

apex predator at the time.

55 Ma Modern bird groups diversify (first song birds, parrots, loons, swifts,
woodpeckers), first whale (Himalayacetus), earliest rodents, lagomorphs, armadillos,
appearance of sirenians, proboscideans, perissodactyl and artiodactyl mammals in the
fossil record. Angiosperms diversify. The ancestor (according to theory) of the species
in Carcharodon, the early mako shark Isurus hastalis, is alive.

52 Ma First bats appear (Onychonycteris).

50 Ma Peak diversity of dinoflagellates and nanofossils, increase in diversity of
anomalodesmatan and heteroconch bivalves, brontotheres, tapirs, rhinoceroses, and
camels appear in the fossil record, diversification of primates.

40 Ma Modern type butterflies and moths appear. Extinction of Gastornis.

Basilosaurus, one of the first of the giant whales, appeared in the fossil record.
37 Ma First Nimravid carnivores ("False Saber-toothed Cats") - these species are
unrelated to modern-type felines

35 Ma Grasses evolve from among the angiosperms; grasslands begin to expand.

Slight increase in diversity of cold-tolerant ostracods and foraminifers, along with major
extinctions of gastropods, reptiles, and amphibians. Many modern mammal groups
begin to appear: first glyptodonts, ground sloths, dogs, peccaries, and the first eagles
and hawks. Diversity in toothed and baleen whales.

33 Ma Evolution of the thylacinid marsupials (Badjcinus).

30 Ma First balanids and eucalypts, extinction of embrithopod and brontothere

mammals, earliest pigs and cats.

28 Ma Paraceratherium appears in the fossil record, the largest terrestrial mammal

that ever lived.

25 Ma First deer.
20 Ma First giraffes and giant anteaters, increase in bird diversity.

15 Ma Mammut appears in the fossil record, first bovids and kangaroos, diversity in
Australian megafauna.

10 Ma Grasslands and savannas are established, diversity in insects, especially ants

and termites, horses increase in body size and develop high-crowned teeth, major
diversification in grassland mammals and snakes.

6.5 Ma First hominin (Sahelanthropus).

6 Ma Australopithecines diversify (Orrorin, Ardipithecus)

5 Ma First tree sloths and hippopotami, diversification of grazing herbivores, large

carnivorous mammals, burrowing rodents, kangaroos, birds, and small carnivores,
vultures increase in size, decrease in the number of perissodactyl mammals. Extinction
of Nimravid carnivores

4.8 Ma Mammoths appear in the fossil record.

4 Ma Evolution of Australopithecus, Stupendemys appears in the fossil record as the

largest freshwater turtle.
 3 Ma The Great American Interchange, where various land and freshwater faunas
migrated between North and South America. Armadillos, opossums, hummingbirds,
and vampire bats traveled to North America while horses, tapirs, saber-toothed cats,
and deer entered South America. The first short-faced bears ( Arctodus) appear.

2.7 Ma Evolution of Paranthropus

2.5 Ma The earliest species of Smilodon evolve

2 Ma First members of the genus Homo appear in the fossil record. Diversification of
conifers in high latitudes. The eventual ancestor of cattle, Bos primigenius evolves in
India

1.7 Ma Extinction of australopithecines.

1.2 Ma Evolution of Homo antecessor. The last members of Paranthropus die out.

600 ka Evolution of Homo heidelbergensis

350 ka Evolution of Neanderthals

300 ka Gigantopithecus, a giant relative of the orangutan dies out from Asia
200 ka Anatomically modern humans appear in Africa.[39][40][41] Around 50,000
years before present they start colonising the other continents, replacing the
Neanderthals in Europe and other hominins in Asia.
 40 ka The last of the giant monitor lizards (Megalania) die out

30 ka Extinction of Neanderthals

15 ka The last Woolly rhinoceros (Coelodonta) are believed to have gone extinct

11 ka The giant short-faced bears (Arctodus) vanish from North America, with the
last Giant Ground Sloths dying out. All Equidae become extinct in North America

10 ka The Holocene Epoch starts 10,000[42] years ago after the Late Glacial
Maximum. The last mainland species of Woolly mammoth (Mammuthus primigenius)
die out, as does the last Smilodon species

6 ka Small populations of American Mastodon die off in places like Utah and
Michigan

4500 ya The last members of a dwarf race of Woolly Mammoths vanish from Wrangel
Island near Alaska

383 ya The last wild Aurochs die out

37 ya The Thylacine goes extinct in a Tasmanian zoo, the last member of the family
Thylacinidae

Present day The impact of Homo sapiens sapiens is felt in all corners of the globe and
contributing to a dramatically rising extinction rate, estimated in 1995 to be 100 times
 the background rate.[43][44]

[edit] See also

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Evolutionary history of life
• Evolutionary history of plants
• Extinction events
• Geologic time scale
• History of Earth
• Natural history
• Sociocultural evolution
• Timeline of human evolution
• Timeline of plant evolution

[edit] Further reading

• The Ancestor's Tale by Richard Dawkins, for a list of ancestors common to humans and
other living species
[edit] References
1. ^ Planetary Science Institute page on the Giant Impact Hypothesis. Hartmann and Davis
belonged to the PSI. This page also contains several paintings of the impact by Hartmann himself.
2. ^ "Because the Moon helps stabilize the tilt of the Earth's rotation, it prevents the Earth from
wobbling between climatic extremes. Without the Moon, seasonal shifts would likely outpace even
the most adaptable forms of life." Making the Moon Astrobiology Magazine. (URL accessed on
August 7, 2010)
3. ^ "However, once the Earth cooled sufficiently, sometime in the first 700 million years of its
existence, clouds began to form in the atmosphere, and the Earth entered a new phase of
development." How the Oceans Formed (URL accessed on January 9, 2005)
4. ^ *The 'PAH World'
5. ^ Gilbert, Walter (February 1986). "The RNA World". Nature 319: 618.
doi:10.1038/319618a0.
6. ^ Joyce, G.F. (2002). "The antiquity of RNA-based evolution". Nature 418 (6894): 214–21.
doi:10.1038/418214a. PMID 12110897.
7. ^ Hoenigsberg, H. (December 2003). "Evolution without speciation but with selection: LUCA,
the Last Universal Common Ancestor in Gilbert’s RNA world". Genetic and Molecular Research 2 (4):
366–375. PMID 15011140. http://www.funpecrp.com.br/gmr/year2003/vol4-2/gmr0070_full_text.htm.
Retrieved 2008-08-30. (also available as PDF)
8. ^ Trevors, J. T. and Abel, D. L. (2004). "Chance and necessity do not explain the origin of
life". Cell Biol. Int. 28 (11): 729–39. doi:10.1016/j.cellbi.2004.06.006. PMID 15563395.
9. ^ Forterre, P., Benachenhou-Lahfa, N., Confalonieri, F., Duguet, M., Elie, C. and Labedan, B.
(1992). "The nature of the last universal ancestor and the root of the tree of life, still open questions".
BioSystems 28 (1-3): 15–32. doi:10.1016/0303-2647(92)90004-I. PMID 1337989.
 10.^ Steenhuysen, Julie (May 21, 2009). "Study turns back clock on origins of life on Earth".
Reuters.com. Reuters. http://www.reuters.com/article/scienceNews/idUSTRE54J5PX20090520?
pageNumber=1&virtualBrandChannel=0. Retrieved May 21, 2009.
11.^ " Between about 3.8 billion and 4.5 billion years ago, no place in the solar system was safe
from the huge arsenal of asteroids and comets left over from the formation of the planets. Sleep and
Zahnle calculate that Earth was probably hit repeatedly by objects up to 500 kilometers across "
Geophysicist Sleep: Martian underground may have harbored early life (URL accessed on January 9,
2005)
12.^ Carl Woese, J Peter Gogarten, "When did eukaryotic cells (cells with nuclei and other
internal organelles) first evolve? What do we know about how they evolved from earlier life-forms?"
Scientific American, October 21, 1999.
13.^ Romano AH, Conway T. (1996) Evolution of carbohydrate metabolic pathways. Res
Microbiol. 147(6-7):448-55 PMID 9084754
14.^ Knowles JR (1980). "Enzyme-catalyzed phosphoryl transfer reactions". Annu. Rev.
Biochem. 49: 877–919. doi:10.1146/annurev.bi.49.070180.004305. PMID 6250450.
15.^ Doolittle, W. Ford (February, 2000). Uprooting the tree of life. Scientific American 282 (6):
90–95.
16.^ Nicolas Glansdorff, Ying Xu & Bernard Labedan: The Last Universal Common Ancestor :
emergence, constitution and genetic legacy of an elusive forerunner. Biology Direct 2008, 3:29.
17.^ Hahn, Jürgen; Pat Haug (1986). "Traces of Archaebacteria in ancient sediments". System
Applied Microbiology 7 (Archaebacteria '85 Proceedings): 178–83.
18.^ Olson JM (May 2006). "Photosynthesis in the Archean era". Photosyn. Res. 88 (2): 109–17.
doi:10.1007/s11120-006-9040-5. PMID 16453059.
 19.^ Buick R (August 2008). "When did oxygenic photosynthesis evolve?". Philos. Trans. R.
Soc. Lond., B, Biol. Sci. 363 (1504): 2731–43. doi:10.1098/rstb.2008.0041. PMID 18468984.
20.^ Knoll, Andrew H.; Javaux, E.J, Hewitt, D. and Cohen, P. (2006). "Eukaryotic organisms in
Proterozoic oceans". Philosophical Transactions of the Royal Society of London, Part B 361 (1470):
1023–38. doi:10.1098/rstb.2006.1843. PMID 16754612.
21.^ Fedonkin, M. A. (March 2003). "The origin of the Metazoa in the light of the Proterozoic
fossil record" (PDF). Paleontological Research 7 (1): 9–41. doi:10.2517/prpsj.7.9.
http://www.vend.paleo.ru/pub/Fedonkin_2003.pdf. Retrieved 2008-09-02.
22.^ Nicholas J. Butterfield, "Bangiomorpha pubescens n. gen., n. sp.: implications for the
evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic radiation of eukaryotes"
23.^ Hoffman, P.F.; Kaufman, A.J., Halverson, G.P., Schrag, D.P. (1998-08-28). "A
Neoproterozoic Snowball Earth". Science 281 (5381): 1342. doi:10.1126/science.281.5381.1342.
PMID 9721097. http://www.sciencemag.org/cgi/content/full/281/5381/1342?
ijkey=48d78da67bab492803c333f50c0dd84fbbef109c. Retrieved 2007-05-04. Full online article (pdf
260 Kb)
24.^ Kirschvink, J.L. (1992). "Late Proterozoic low-latitude global glaciation: The snowball
Earth". in Schopf, JW, and Klein, C. (PDF). The Proterozoic Biosphere: A Multidisciplinary Study .
Cambridge University Press, Cambridge. pp. 51–52.
http://www.gps.caltech.edu/~jkirschvink/pdfs/firstsnowball.pdf.
25.^ http://researchpages.net/media/resources/2007/06/21/richtimhywelfinal.pdf
26.^ Corsetti, F.A.; Awramik, S.M.; Pierce, D. (2003-04-15). "A complex microbiota from
snowball Earth times: Microfossils from the Neoproterozoic Kingston Peak Formation, Death Valley,
USA". Proceedings of the National Academy of Sciences 100 (8): 4399–4404.
doi:10.1073/pnas.0730560100. PMID 12682298. PMC 153566.
http://www.pnas.org/cgi/content/abstract/100/8/4399. Retrieved 2007-06-28.
27.^ Corsetti, F.A.; Olcott, A.N.; Bakermans, C. (2006). "The biotic response to Neoproterozoic
Snowball Earth". Palaeogeography, Palaeoclimatology, Palaeoecology 232 (232): 114–130.
doi:10.1016/j.palaeo.2005.10.030.
28.^ Narbonne, Guy (June 2006). "The Origin and Early Evolution of Animals". Department of
Geological Sciences and Geological Engineering, Queen's University.
http://geol.queensu.ca/people/narbonne/recent_pubs1.html. Retrieved 2007-03-10.
29.^ The Cambrian Period
30.^ The Cambrian Explosion – Timing
31.^ a b Formation of the Ozone Layer
32.^ "The oldest fossils of footprints ever found on land hint that animals may have beaten
plants out of the primordial seas. Lobster-sized, centipede-like or slug like animals such as
Protichnites and Climactichnites made the prints wading out of the ocean and scuttling over sand
dunes about 530 million years ago. Previous fossils indicated that animals didn't take this step until
40 million years later." Oldest fossil footprints on land
33.^ "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician
Period (~450–440 Ma) on the basis of fossil spores" Transition of plants to land
34.^ "The land plants evolved from the algae, more specifically green algae, as suggested by
certain common biochemical traits" The first land plants
35.^ "The ancestry of sharks dates back more than 200 million years before the earliest known
dinosaur. Introduction to shark evolution, geologic time and age determination
36.^ Sahney, S. and Benton, M.J. (2008). "Recovery from the most profound mass extinction of
all time" (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759.
doi:10.1098/rspb.2007.1370. PMID 18198148. PMC 2596898.
http://journals.royalsociety.org/content/qq5un1810k7605h5/fulltext.pdf.
37.^ "Viruses of nearly all the major classes of organisms—animals, plants, fungi and
bacteria/archaea—probably evolved with their hosts in the seas, given that most of the evolution of
life on this planet has occurred there. This means that viruses also probably emerged from the
waters with their different hosts, during the successive waves of colonisation of the terrestrial
environment." Origins of Viruses (URL accessed on January 9, 2005)
38.^ Chiappe, Luis M., & Dyke, Gareth J. (2002). "The Mesozoic Radiation of Birds". Annual
Review of Ecology & Systematics 33: 91–124. doi:10.1146/annurev.ecolsys.33.010802.150517.
39.^ The Oldest Homo Sapiens: - URL retrieved May 15, 2009
40.^ Alemseged, Z., Coppens, Y., Geraads, D. (2002). "Hominid cranium from Homo:
Description and taxonomy of Homo-323-1976-896". Am J Phys Anthropol 117 (2): 103–12.
doi:10.1002/ajpa.10032. PMID 11815945.
41.^ Stoneking, Mark; Soodyall, Himla (1996). "Human evolution and the mitochondrial
genome". Current Opinion in Genetics & Development 6 (6): 731–6. doi:10.1016/S0959-
437X(96)80028-1.
42.^ "International Stratigraphic Chart". International Commission on Stratigraphy.
http://www.stratigraphy.org/cheu.pdf. Retrieved 2009-02-03.
43.^ The American Museum of Natural History National Survey Reveals Biodiversity Crisis (URL
accessed on February 23, 2006)
44.^ J.H.Lawton and R.M.May, Extinction Rates, Oxford University Press, Oxford, UK
 [edit] External links
• Berkeley Evolution
• Tree of Life Web Project - explore complete phylogenetic tree interactively
• A more compact timeline at the TalkOrigins Archive
• Palaeos - The Trace of Life on Earth
• University of Waikato - Sequence of Plant Evolution
• University of Waikato - Sequence of Animal Evolution
• Graphical Timeline of evolution
• History of Life on Earth
• Exploring Time from Planck Time to the lifespan of the universe
• Interactive Plant Evolution Timeline - from the University of Cambridge Ensemble Project

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Evolutionary history of life

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The evolutionary history of life on Earth traces the processes by which living and fossil organisms
evolved. It stretches from the origin of life on Earth, thought to be over 3,500 million years ago, to the present
day. The similarities between all present day organisms indicate the presence of a common ancestor from
which all known species have diverged through the process of evolution.[1]
 Microbial mats of coexisting bacteria and archaea were the dominant form of life in the early Archean
and many of the major steps in early evolution are thought to have taken place within them.[2] The evolution
of oxygenic photosynthesis, around 3,500 million years ago, eventually led to the oxygenation of the
atmosphere, beginning around 2,400 million years ago.[3] The earliest evidence of eukaryotes (complex cells
with organelles), dates from 1,850 million years ago[4][5], and while they may have been present earlier, their
diversification accelerated when they started using oxygen in their metabolism. Later, around 1,700 million
years ago, multicellular organisms began to appear, with differentiated cells performing specialised functions.
[6]
The earliest land plants date back to around 450 million years ago,[7] though evidence suggests that
algal scum formed on the land as early as 1,200 million years ago. Land plants were so successful that they
are thought to have contributed to the late Devonian extinction event.[8] Invertebrate animals appear during
the Vendian period,[9] while vertebrates originated about 525 million years ago during the Cambrian
explosion.[10]
During the Permian period, synapsids, including the ancestors of mammals, dominated the land,[11]
but the Permian–Triassic extinction event 251 million years ago came close to wiping out all complex life.[12]
During the recovery from this catastrophe, archosaurs became the most abundant land vertebrates,
displacing therapsids in the mid-Triassic.[13] One archosaur group, the dinosaurs, dominated the Jurassic
and Cretaceous periods,[14] with the ancestors of mammals surviving only as small insectivores.[15] After
the Cretaceous–Tertiary extinction event 65 million years ago killed off the non-avian dinosaurs[16] mammals
increased rapidly in size and diversity.[17] Such mass extinctions may have accelerated evolution by
providing opportunities for new groups of organisms to diversify.[18]
Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early
Cretaceous, between 130 million years ago and 90 million years ago, probably helped by coevolution with
pollinating insects. Flowering plants and marine phytoplankton are still the dominant producers of organic
matter. Social insects appeared around the same time as flowering plants. Although they occupy only small
parts of the insect "family tree", they now form over half the total mass of insects. Humans evolved from a
lineage of upright-walking apes whose earliest fossils date from over 6 million years ago. Although early
members of this lineage had chimp-sized brains, there are signs of a steady increase in brain size after about
3 million years ago.
Contents
[hide]
• 1 Earliest history of Earth
• 2 Earliest evidence for life on Earth
• 3 Origins of life on Earth
• 3.1 Life "seeded" from elsewhere
• 3.2 Independent emergence on Earth
• 3.2.1 Replication first: RNA world
• 3.2.2 Metabolism first: Iron-sulfur world
• 3.2.3 Membranes first: Lipid world
• 3.2.4 The clay theory
• 4 Environmental and evolutionary impact of microbial mats
• 5 Diversification of eukaryotes
• 6 Multicellular organisms and sexual reproduction
• 6.1 Multicellularity
• 6.2 Evolution of sexual reproduction
• 6.3 Fossil evidence for multicellularity and sexual
reproduction
• 7 Emergence of animals
• 8 Colonization of land
• 8.1 Evolution of soil
-4500 —
–
-4000 —
–
-3500 —
–
-3000 —
–
-2500 —
–
-2000 —
–
-1500 —
–
-1000 —
–
-500 —
–
0—
Hadean
Archean
Protero
-zoic
 Phanero
-zoic
Eo
Paleo
Meso
Neo
Paleo
Meso
Neo
Paleo
Meso
Ceno

←
Solar system formed

←
Impact formed Moon

←
? Cool surface, oceans, atmosphere

←
Late Heavy Bombardment

←
? Earliest evidence of life

←
Oxygenation of atmosphere

←
Earliest multicellular organism[19]
 ←
Earliest known fungi

←
Earliest known cnidarians

←
? Cambrian explosion

←
Earliest land invertebrates and plants

←
Earliest land vertebrates
 ←
Earliest known dinosaur

←
Extinction of non-avian dinosaurs

Scale:
Millions of years
Main article: History of the Earth
The oldest meteorite fragments found on Earth are about 4,540 million years old, this, coupled
primarily with the dating of ancient lead deposits, has put the estimated age of Earth at around that time.[20]
About 40 million years later a planetoid struck the Earth, throwing into orbit the material that formed the
Moon.[21]
Until recently the oldest rocks found on Earth were about 3,800 million years old,[20] leading
scientists to believe for decades that Earth's surface had been molten until then. Accordingly, they named
this part of Earth's history the Hadean eon, whose name means "hellish".[22] However analysis of zircons
formed 4,400 to 4,000 million years ago indicates that Earth's crust solidified about 100 million years after the
planet's formation and that the planet quickly acquired oceans and an atmosphere, which may have been
capable of supporting life.[23]
 Evidence from the Moon indicates that from 4,000 to 3,800 million years ago it suffered a Late Heavy
Bombardment by debris that was left over from the formation of the Solar system, and the Earth should have
experienced an even heavier bombardment due to its stronger gravity.[22][24] While there is no direct
evidence of conditions on Earth 4,000 to 3,800 million years ago, there is no reason to think that the Earth
was not also affected by this late heavy bombardment.[25] This event may well have stripped away any
previous atmosphere and oceans; in this case gases and water from comet impacts may have contributed to
their replacement, although volcanic outgassing on Earth would have contributed at least half.[26]

[edit] Earliest evidence for life on Earth

The earliest identified organisms were minute and relatively featureless, their fossils look like small
rods, which are very difficult to tell apart from structures that arise through abiotic physical processes. The
oldest undisputed evidence of life on Earth, interpreted as fossilized bacteria, dates to 3,000 million years
ago.[27] Other finds in rocks dated to about 3,500 million years ago have been interpreted as bacteria,[28]
with geochemical evidence also seeming to show the presence of life 3,800 million years ago.[29] However
these analyses were closely scrutinized, and non-biological processes were found which could produce all of
the "signatures of life" that had been reported.[30][31] While this does not prove that the structures found had
a non-biological origin, they cannot be taken as clear evidence for the presence of life. Currently, the oldest
unchallenged evidence for life is geochemical signatures from rocks deposited 3,400 million years ago,[27]
[32] although these statements have not been thoroughly examined by critics.
[edit] Origins of life on Earth

Evolutionary tree showing the divergence of modern species from their common ancestor in the
center.[33] The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.
Further information: Evidence of common descent, Common descent, and Homology (biology)
 Biologists reason that all living organisms on Earth must share a single last universal ancestor,
because it would be virtually impossible that two or more separate lineages could have independently
developed the many complex biochemical mechanisms common to all living organisms.[34][35] As previously
mentioned the earliest organisms for which fossil evidence is available are bacteria, cells far too complex to
have arisen directly from non-living materials.[36] The lack of fossil or geochemical evidence for earlier
organisms has left plenty of scope for hypotheses, which fall into two main groups: 1) that life arose
spontaneously on Earth or 2) that it was "seeded" from elsewhere in the universe.[37]

[edit] Life "seeded" from elsewhere

Main articles: Panspermia, Life on Mars, Fermi paradox, and Rare Earth hypothesis
The idea that life on Earth was "seeded" from elsewhere in the universe dates back at least to the
fifth century BC.[38] In the twentieth century it was proposed by the physical chemist Svante Arrhenius,[39]
by the astronomers Fred Hoyle and Chandra Wickramasinghe,[40] and by molecular biologist Francis Crick
and chemist Leslie Orgel.[41] There are three main versions of the "seeded from elsewhere" hypothesis:
from elsewhere in our Solar system via fragments knocked into space by a large meteor impact, in which
case the only credible source is Mars;[42] by alien visitors, possibly as a result of accidental contamination by
micro-organisms that they brought with them;[41] and from outside the Solar system but by natural means.
[39][42] Experiments suggest that some micro-organisms can survive the shock of being catapulted into
space and some can survive exposure to radiation for several days, but there is no proof that they can
survive in space for much longer periods.[42] Scientists are divided over the likelihood of life arising
independently on Mars,[43] or on other planets in our galaxy.[42]
[edit] Independent emergence on Earth
Main article: Abiogenesis
Life on earth is based on carbon and water. Carbon provides stable frameworks for complex
chemicals and can be easily extracted from the environment, especially from carbon dioxide. The only other
element with similar chemical properties, silicon, forms much less stable structures and, because most of its
compounds are solids, would be more difficult for organisms to extract. Water is an excellent solvent and has
two other useful properties: the fact that ice floats enables aquatic organisms to survive beneath it in winter;
and its molecules have electrically negative and positive ends, which enables it to form a wider range of
compounds than other solvents can. Other good solvents, such as ammonia, are liquid only at such low
temperatures that chemical reactions may be too slow to sustain life, and lack water's other advantages.[44]
Organisms based on alternative biochemistry may however be possible on other planets.[45]
Research on how life might have emerged unaided from non-living chemicals focuses on three
possible starting points: self-replication, an organism's ability to produce offspring that are very similar to
itself; metabolism, its ability to feed and repair itself; and external cell membranes, which allow food to enter
and waste products to leave, but exclude unwanted substances.[46] Research on abiogenesis still has a long
way to go, since theoretical and empirical approaches are only beginning to make contact with each other.
[47][48]

[edit] Replication first: RNA world

Main articles: Last universal ancestor and RNA world
 Even the simplest members of the three modern domains of life use DNA
to record their "recipes" and a complex array of RNA and protein molecules to
"read" these instructions and use them for growth, maintenance and self-
replication. This system is far too complex to have emerged directly from non-
living materials.[36] The discovery that some RNA molecules can catalyze both
their own replication and the construction of proteins led to the hypothesis of
earlier life-forms based entirely on RNA.[49] These ribozymes could have formed
an RNA world in which there were individuals but no species, as mutations and
horizontal gene transfers would have meant that the offspring in each generation
were quite likely to have different genomes from those that their parents started
with.[50] RNA would later have been replaced by DNA, which is more stable and
therefore can build longer genomes, expanding the range of capabilities a single
organism can have.[50][51][52] Ribozymes remain as the main components of
ribosomes, modern cells' "protein factories".[53]
Although short self-replicating RNA molecules have been artificially
produced in laboratories,[54] doubts have been raised about where natural non-
biological synthesis of RNA is possible.[55] The earliest "ribozymes" may have
been formed of simpler nucleic acids such as PNA, TNA or GNA, which would
have been replaced later by RNA.[56][57]
In 2003 it was proposed that porous metal sulfide precipitates would
assist RNA synthesis at about 100 °C (212 °F) and ocean-bottom pressures near
The replicator in
virtually all known life is
deoxyribonucleic acid.
DNA's structure and
replication systems are
far more complex than
those of the original
replicator.[36]
hydrothermal vents. In this hypothesis lipid membranes would be the last major cell components to appear
and until then the proto-cells would be confined to the pores.[58]

[edit] Metabolism first: Iron-sulfur world

Main article: Iron-sulfur world theory
A series of experiments starting in 1997 showed that early stages in the formation of proteins from
inorganic materials including carbon monoxide and hydrogen sulfide could be achieved by using iron sulfide
and nickel sulfide as catalysts. Most of the steps required temperatures of about 100 °C (212 °F) and
moderate pressures, although one stage required 250 °C (482 °F) and a pressure equivalent to that found
under 7 kilometres (4.3 mi) of rock. Hence it was suggested that self-sustaining synthesis of proteins could
have occurred near hydrothermal vents.[59]

[edit] Membranes first: Lipid world

It has been suggested that double-walled "bubbles" of
lipids like those that form the external membranes of cells may = water-attracting heads of lipid
have been an essential first step.[60] Experiments that
simulated the conditions of the early Earth have reported the = water-repellent tails
formation of lipids, and these can spontaneously form
liposomes, double-walled "bubbles", and then reproduce
themselves. Although they are not intrinsically information-
carriers as nucleic acids are, they would be subject to natural

Cross-section through a liposome.

selection for longevity and reproduction. Nucleic acids such as RNA might then have formed more easily
within the liposomes than they would have outside.[61]

[edit] The clay theory

Main articles: Graham Cairns-Smith#Clay Theory and RNA world
RNA is complex and there are doubts about whether it can be produced non-biologically in the wild.
[55] Some clays, notably montmorillonite, have properties that make them plausible accelerators for the
emergence of an RNA world: they grow by self-replication of their crystalline pattern; they are subject to an
analog of natural selection, as the clay "species" that grows fastest in a particular environment rapidly
becomes dominant; and they can catalyze the formation of RNA molecules.[62] Although this idea has not
become the scientific consensus, it still has active supporters.[63]
Research in 2003 reported that montmorillonite could also accelerate the conversion of fatty acids
into "bubbles", and that the "bubbles" could encapsulate RNA attached to the clay. These "bubbles" can then
grow by absorbing additional lipids and then divide. The formation of the earliest cells may have been aided
by similar processes.[64]
A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides, lipids
and amino acids.[65]

[edit] Environmental and evolutionary impact of microbial mats

Main articles: Microbial mat and Oxygen catastrophe
 Modern stromatolites in Shark Bay, Western Australia.
Microbial mats are multi-layered, multi-species colonies of bacteria and other organisms that are
generally only a few millimeters thick, but still contain a wide range of chemical environments, each of which
favors a different set of micro-organisms.[66] To some extent each mat forms its own food chain, as the by-
products of each group of micro-organisms generally serve as "food" for adjacent groups.[67]
Stromatolites are stubby pillars built as microbes in mats slowly migrate upwards to avoid being
smothered by sediment deposited on them by water.[66] There has been vigorous debate about the validity
of alleged fossils from before 3,000 million years ago,[68] with critics arguing that so-called stromatolites
could have been formed by non-biological processes.[30] In 2006 another find of stromatolites was reported
from the same part of Australia as previous ones, in rocks dated to 3,500 million years ago.[69]
In modern underwater mats the top layer often consists of photosynthesizing cyanobacteria which
create an oxygen-rich environment, while the bottom layer is oxygen-free and often dominated by hydrogen
sulfide emitted by the organisms living there.[67] It is estimated that the appearance of oxygenic
photosynthesis by bacteria in mats increased biological productivity by a factor of between 100 and 1,000.
The reducing agent used by oxygenic photosynthesis is water, which is much more plentiful than the
geologically-produced reducing agents required by the earlier non-oxygenic photosynthesis.[70] From this
point onwards life itself produced significantly more of the resources it needed than did geochemical
processes.[71] Oxygen is toxic to organisms that are not adapted to it, but greatly increases the metabolic
efficiency of oxygen-adapted organisms.[72][73]
Oxygen became a significant component of Earth's atmosphere about 2,400 million years ago.[74]
Although eukaryotes may have been present much earlier,[75][76] the oxygenation of the atmosphere was a
prerequisite for the evolution of the most complex eukaryotic cells, from which all multicellular organisms are
built.[77] The boundary between oxygen-rich and oxygen-free layers in microbial mats would have moved
upwards when photosynthesis shut down overnight, and then downwards as it resumed on the next day. This
would have created selection pressure for organisms in this intermediate zone to acquire the ability to
tolerate and then to use oxygen, possibly via endosymbiosis, where one organism lives inside another and
both of them benefit from their association.[2]
Cyanobacteria have the most complete biochemical "toolkits" of all the mat-forming organisms.
Hence they are the most self-sufficient of the mat organisms and were well-adapted to strike out on their own
both as floating mats and as the first of the phytoplankton, providing the basis of most marine food chains.[2]
 Eukaryotes
Bikonta

Apusozoa

[edit] Diversification of eukaryotes Archaeplastida (Land plants

Main article: Eukaryote green algae, red algae, and
glaucophytes)
Eukaryotes may have been present long before the
oxygenation of the atmosphere,[75] but most modern eukaryotes
Chromalveolata
require oxygen, which their mitochondria use to fuel the
production of ATP, the internal energy supply of all known cells.
[77] In the 1970s it was proposed and, after much debate, widely
accepted that eukaryotes emerged as a result of a sequence of Rhizaria
endosymbioses between "procaryotes". For example: a
predatory micro-organism invaded a large procaryote, probably
an archaean, but the attack was neutralized, and the attacker Excavata
took up residence and evolved into the first of the mitochondria;
one of these chimeras later tried to swallow a photosynthesizing
cyanobacterium, but the victim survived inside the attacker and
the new combination became the ancestor of plants; and so on.
After each endosymbiosis began, the partners would have Unikonta
eliminated unproductive duplication of genetic functions by re-
arranging their genomes, a process which sometimes involved Amoebozoa
transfer of genes between them.[80][81][82] Another hypothesis
proposes that mitochondria were originally sulfur- or hydrogen- Opisthokonta
metabolising endosymbionts, and became oxygen-consumers
later.[83] On the other hand mitochondria might have been part Metazoa
of eukaryotes' original equipment.[84] (Animals)

Choanozoa

Eumycota
(Fungi)
 There is a debate about when eukaryotes first appeared: the presence of steranes in Australian
shales may indicate that eukaryotes were present 2,700 million years ago;[76] however an analysis in 2008
concluded that these chemicals infiltrated the rocks less than 2,200 million years ago and prove nothing
about the origins of eukaryotes.[85] Fossils of the alga Grypania have been reported in 1,850 million-year-old
rocks (originally dated to 2,100 million years ago but later revised[5]), and indicates that eukaryotes with
organelles had already evolved.[86] A diverse collection of fossil algae were found in rocks dated between
1,500 million years ago and 1,400 million years ago.[87] The earliest known fossils of fungi date from 1,430
million years ago.[88]

[edit] Multicellular organisms and sexual reproduction

[edit] Multicellularity
Main articles: Multicellular organism , Evolution of multicellularity , and Sexual reproduction

A slime mold solves a maze. The mold (yellow) explored and filled the maze (left). When the
researchers placed sugar (red) at two separate points, the mold concentrated most of its mass there and left
only the most efficient connection between the two points (right).[89]
 The simplest definitions of "multicellular", for example "having multiple cells", could include colonial
cyanobacteria like Nostoc. Even a professional biologist's definition such as "having the same genome but
different types of cell" would still include some genera of the green alga Volvox, which have cells that
specialize in reproduction.[90] Multicellularity evolved independently in organisms as diverse as sponges and
other animals, fungi, plants, brown algae, cyanobacteria, slime moulds and myxobacteria.[5][91] For the sake
of brevity this article focuses on the organisms that show the greatest specialization of cells and variety of
cell types, although this approach to the evolution of complexity could be regarded as "rather
anthropocentric".[92]
The initial advantages of multicellularity may have included: increased resistance to predators, many
of which attacked by engulfing; the ability to resist currents by attaching to a firm surface; the ability to reach
upwards to filter-feed or to obtain sunlight for photosynthesis;[93] the ability to create an internal environment
that gives protection against the external one;[92] and even the opportunity for a group of cells to behave
"intelligently" by sharing information.[89] These features would also have provided opportunities for other
organisms to diversify, by creating more varied environments than flat microbial mats could.[93]
Multicellularity with differentiated cells is beneficial to the organism as a whole but disadvantageous
from the point of view of individual cells, most of which lose the opportunity to reproduce themselves. In an
asexual multicellular organism, rogue cells which retain the ability to reproduce may take over and reduce the
organism to a mass of undifferentiated cells. Sexual reproduction eliminates such rogue cells from the next
generation and therefore appears to be a prerequisite for complex multicellularity.[93]
The available evidence indicates that eukaryotes evolved much earlier but remained inconspicuous
until a rapid diversification around 1,000 million years ago. The only respect in which eukaryotes clearly
surpass bacteria and archaea is their capacity for variety of forms, and sexual reproduction enabled
eukaryotes to exploit that advantage by producing organisms with multiple cells that differed in form and
function.[93]

[edit] Evolution of sexual reproduction

Main article: Evolution of sexual reproduction
The defining characteristic of sexual reproduction is recombination, in which each of the offspring
receives 50% of its genetic inheritance from each of the parents.[94] Bacteria also exchange DNA by
bacterial conjugation, the benefits of which include resistance to antibiotics and other toxins, and the ability to
utilize new metabolites.[95] However conjugation is not a means of reproduction, and is not limited to
members of the same species – there are cases where bacteria transfer DNA to plants and animals.[96]
The disadvantages of sexual reproduction are well-known: the genetic reshuffle of recombination
may break up favorable combinations of genes; and since males do not directly increase the number of
offspring in the next generation, an asexual population can out-breed and displace in as little as 50
generations a sexual population that is equal in every other respect.[94] Nevertheless the great majority of
animals, plants, fungi and protists reproduce sexually. There is strong evidence that sexual reproduction
arose early in the history of eukaryotes and that the genes controlling it have changed very little since then.
[97] How sexual reproduction evolved and survived is an unsolved puzzle.[98]
The Red Queen Hypothesis suggests that sexual reproduction provides protection against parasites,
because it is easier for parasites to evolve means of overcoming the defenses of genetically identical clones
than those of sexual species that present moving targets, and there is some experimental evidence for this.
However there is still doubt about whether it would explain the survival of sexual species if multiple similar
clone species were present, as one of the clones may survive the attacks of parasites for long enough to out-
breed the sexual species.[94]
The Mutation Deterministic Hypothesis assumes that each organism has more than one harmful
mutation and the combined effects of these mutations are more harmful than the sum of the harm done by
each individual mutation. If so, sexual recombination of genes will reduce the harm that bad mutations do to
offspring and at the same time eliminate some bad mutations from the gene pool by isolating them in
individuals that perish quickly because they have an above-average number of bad mutations. However the
evidence suggests that the MDH's assumptions are shaky, because many species have on average less
than one harmful mutation per individual and no species that has been investigated shows evidence of
synergy between harmful mutations.[94]

Horodyskia apparently re-arranged itself into fewer but larger main masses as the sediment grew
deeper round its base.[5]
 The random nature of recombination causes the relative abundance of alternative traits to vary from
one generation to another. This genetic drift is insufficient on its own to make sexual reproduction
advantageous, but a combination of genetic drift and natural selection may be sufficient. When chance
produces combinations of good traits, natural selection gives a large advantage to lineages in which these
traits become genetically linked. On the other hand the benefits of good traits are neutralized if they appear
along with bad traits. Sexual recombination gives good traits the opportunities to become linked with other
good traits, and mathematical models suggest this may be more than enough to offset the disadvantages of
sexual reproduction.[98] Other combinations of hypotheses that are inadequate on their own are also being
examined.[94]
The following hypotheses attempt to explain how and why sex evolved:
• It may have enabled organisms to repair genetic damage.[99] The most primitive form of sex
may have been one organism repairing damaged DNA by replicating an undamaged strand from a
similar organism.[100]
• Sexual reproduction may have originated from selfish parasitic genetic elements propagating
themselves by transfer to new hosts.[101]
• It may have evolved from cannibalism, where some of the victim's DNA was incorporated into
the cannibal organism.[100]
• Sexual reproduction may have evolved from ancient haloarchaea through a combination of
jumping genes, and swapping plasmids.[102]
• Or it may have evolved as a form of vaccination in which infected hosts exchanged
weakened symbiotic copies of parasitic DNA as protection against more virulent versions. The
meiosis stage of sexual reproduction may then have evolved as a way of removing the symbiotes.
[103]
 Bacteria also exchange DNA by bacterial conjugation, the benefits of which include resistance to
antibiotics and other toxins, and the ability to utilize new metabolites.[95] However conjugation is not a
means of reproduction and is not limited to members of the same species, and there are cases where
bacteria transfer DNA to plants and animals.[104] Nevertheless it may be an example of the "selfish genetic
element" hypothesis, as it transfers DNA by means of such a "selfish gene", the F-plasmid.[100]

[edit] Fossil evidence for multicellularity and sexual reproduction

The Francevillian Group Fossil, dated to 2,100 million years ago, is the
earliest known fossil organism that is clearly multicellular.[19] They may have
had differentiated cells.[106] Another early multicellular fossil, Qingshania,
[note 1] dated to 1,700 million years ago, appears to consist of virtually
identical cells. The red alga called Bangiomorpha, dated at 1,200 million years
ago, is the earliest known organism which certainly has differentiated,
specialized cells, and is also the oldest known sexually-reproducing organism.
[93] The 1,430 million-year-old fossils interpreted as fungi appear to have been
multicellular with differentiated cells.[88] The "string of beads" organism
Horodyskia, found in rocks dated from 1,500 million years ago to 900 million
years ago, may have been an early metazoan;[5] however it has also been
interpreted as a colonial foraminiferan.[105]
Horodyskia may
[edit] Emergence of animals have been an early
Main articles: Animal, Ediacara biota, Cambrian Explosion, Burgess metazoan,[5] or a colonial
shale type fauna, and Stem group foraminiferan[105]
 Animals are multicellular eukaryotes,[note 2] and are Bilaterians
distinguished from plants, algae, and fungi by lacking cell walls.
[108] All animals are motile,[109] if only at certain life stages. All Deuterostomes
animals except sponges have bodies differentiated into separate (chordates
tissues, including muscles, which move parts of the animal by hemichordates
contracting, and nerve tissue, which transmits and processes echinoderms
signals.[110] Protostomes
The earliest widely-accepted animal fossils are rather Ecdysozoa
modern-looking cnidarians (the group that includes jellyfish, sea (arthropods,
anemones and hydras), possibly from around 580 million years nematodes, tardigrades,
ago, although fossils from the Doushantuo Formation can only etc.)
be dated approximately. Their presence implies that the
cnidarian and bilaterian lineages had already diverged.[111] Lophotrochozoa
(molluscs, annelids,
The Ediacara biota, which flourished for the last brachiopods, etc.)
40 million years before the start of the Cambrian,[112] were the
first animals more than a very few centimeters long. Many were
flat and had a "quilted" appearance, and seemed so strange that
there was a proposal to classify them as a separate kingdom,
Vendozoa.[113] Others, however, been interpreted as early
molluscs (Kimberella[114][115]), echinoderms (Arkarua[116]),
and arthropods (Spriggina,[117] Parvancorina[118]). There is
still debate about the classification of these specimens, mainly
because the diagnostic features which allow taxonomists to
classify more recent organisms, such as similarities to living organisms, are generally absent in the
Ediacarans. However there seems little doubt that Kimberella was at least a triploblastic bilaterian animal, in
other words significantly more complex than cnidarians.[119]
The small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and
Mid Cambrian periods. The earliest, Cloudina, shows signs of successful defense against predation and may
indicate the start of an evolutionary arms race. Some tiny Early Cambrian shells almost certainly belonged to
molluscs, while the owners of some "armor plates", Halkieria and Microdictyon, were eventually identified
when more complete specimens were found in Cambrian lagerstätten that preserved soft-bodied animals.
[120]

Opabinia made the largest single contribution to modern interest in the Cambrian explosion.[121]
In the 1970s there was already a debate about whether the emergence of the modern phyla was
"explosive" or gradual but hidden by the shortage of Pre-Cambrian animal fossils.[120] A re-analysis of
fossils from the Burgess Shale lagerstätte increased interest in the issue when it revealed animals, such as
Opabinia, which did not fit into any known phylum. At the time these were interpreted as evidence that the
modern phyla had evolved very rapidly in the "Cambrian explosion" and that the Burgess Shale's "weird
wonders" showed that the Early Cambrian was a uniquely experimental period of animal evolution.[122] Later
discoveries of similar animals and the development of new theoretical approaches led to the conclusion that
many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups[123] – for example that
Opabinia was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it
may have been closely related to the modern tardigrades.[124] Nevertheless there is still much debate about
whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it
appears unique in the history of animals.[125]

Acanthodians were among the earliest vertebrates with jaws[126]

Most of the animals at the heart of the Cambrian explosion debate are protostomes, one of the two
main groups of complex animals. One deuterostome group, the echinoderms, many of which have hard
calcite "shells", are fairly common from the Early Cambrian small shelly fauna onwards.[120] Other
deuterostome groups are soft-bodied, and most of the significant Cambrian deuterostome fossils come from
the Chengjiang fauna, a lagerstätte in China.[127] The Chengjiang fossils Haikouichthys and Myllokunmingia
appear to be true vertebrates,[128] and Haikouichthys had distinct vertebrae, which may have been slightly
mineralized.[129] Vertebrates with jaws, such as the Acanthodians, first appeared in the Late Ordovician.
[130]
[edit] Colonization of land
Adaptation to life on land is a major challenge: all land organisms need to avoid drying-out and all
those above microscopic size have to resist gravity; respiration and gas exchange systems have to change;
reproductive systems cannot depend on water to carry eggs and sperm towards each other.[131][132]
Although the earliest good evidence of land plants and animals dates back to the Ordovician period
(488 to 444 million years ago), modern land ecosystems only appeared in the late Devonian, about
385 to 359 million years ago.[133]

[edit] Evolution of soil

Before the colonization of land, soil, a combination of mineral particles and decomposed organic
matter, did not exist. Land surfaces would have been either bare rock or unstable sand produced by
weathering. Water and any nutrients in it would have drained away very quickly.[133]

Lichens growing on concrete

 Films of cyanobacteria, which are not plants but use the same photosynthesis mechanisms, have
been found in modern deserts, and only in areas that are unsuitable for vascular plants. This suggests that
microbial mats may have been the first organisms to colonize dry land, possibly in the Precambrian. Mat-
forming cyanobacteria could have gradually evolved resistance to desiccation as they spread from the seas
to tidal zones and then to land.[133] Lichens, which are symbiotic combinations of a fungus (almost always
an ascomycete) and one or more photosynthesizers (green algae or cyanobacteria),[134] are also important
colonizers of lifeless environments,[133] and their ability to break down rocks contributes to soil formation in
situations where plants cannot survive.[134] The earliest known ascomycete fossils date from 423 to 419
million years ago in the Silurian.[133]
Soil formation would have been very slow until the appearance of burrowing animals, which mix the
mineral and organic components of soil and whose feces are a major source of the organic components.
[133] Burrows have been found in Ordovician sediments, and are attributed to annelids ("worms") or
arthropods.[133][135]

[edit] Plants and the Late Devonian wood crisis

Main article: Evolutionary history of plants
Reconstruction of Cooksonia, a vascular plant from the Silurian.
 Fossilized trees from the Mid-Devonian Gilboa fossil forest.
In aquatic algae, almost all cells are capable of photosynthesies and are nearly independent. Life on
land required plants to become internally more complex and specialized: photosynthesis was most efficient
at the top; roots were required in order to extract water from the ground; the parts in between became
supports and transport systems for water and nutrients.[131][136]
Spores of land plants, possibly rather like liverworts, have been found in Mid Ordovician rocks dated
to about 476 million years ago. In Mid Silurian rocks 430 million years ago there are fossils of actual plants
including clubmosses such as Baragwanathia; most were under 10 centimetres (3.9 in) high, and some
appear closely related to vascular plants, the group that includes trees.[136]
By the Late Devonian 370 million years ago, trees such as Archaeopteris were so abundant that they
changed river systems from mostly braided to mostly meandering, because their roots bound the soil firmly.
[137] In fact they caused a "Late Devonian wood crisis",[138] because:
• They removed more carbon dioxide from the atmosphere, reducing the greenhouse effect
and thus causing an ice age in the Carboniferous period.[139] In later ecosystems the carbon dioxide
"locked up" in wood is returned to the atmosphere by decomposition of dead wood. However the
earliest fossil evidence of fungi that can decompose wood also comes from the Late Devonian.[140]
• The increasing depth of plants' roots led to more washing of nutrients into rivers and seas by
rain. This caused algal blooms whose high consumption of oxygen caused anoxic events in deeper
waters, increasing the extinction rate among deep-water animals.[139]
 [edit] Land invertebrates
Animals had to change their feeding and excretory systems, and most land animals developed
internal fertilization of their eggs. The difference in refractive index between water and air required changes
in their eyes. On the other hand in some ways movement and breathing became easier, and the better
transmission of high-frequency sounds in air encouraged the development of hearing.[132]
Some trace fossils from the Cambrian-Ordovician boundary about 490 million years ago are
interpreted as the tracks of large amphibious arthropods on coastal sand dunes, and may have been made
by euthycarcinoids,[141] which are thought to be evolutionary "aunts" of myriapods.[142] Other trace fossils
from the Late Ordovician a little over 445 million years ago probably represent land invertebrates, and there
is clear evidence of numerous arthropods on coasts and alluvial plains shortly before the Silurian-Devonian
boundary, about 415 million years ago, including signs that some arthropods ate plants.[143] Arthropods
were well pre-adapted to colonise land, because their existing jointed exoskeletons provided protection
against desiccation, support against gravity and a means of locomotion that was not dependent on water.
[144]
The fossil record of other major invertebrate groups on land is poor: none at all for non-parasitic
flatworms, nematodes or nemerteans; some parasitic nematodes have been fossilized in amber; annelid
worm fossils are known from the Carboniferous, but they may still have been aquatic animals; the earliest
fossils of gastropods on land date from the Late Carboniferous, and this group may have had to wait until leaf
litter became abundant enough to provide the moist conditions they need.[132]
The earliest confirmed fossils of flying insects date from the Late Carboniferous, but it is thought that
insects developed the ability to fly in the Early Carboniferous or even Late Devonian. This gave them a wider
range of ecological niches for feeding and breeding, and a means of escape from predators and from
unfavorable changes in the environment.[145] About 99% of modern insect species fly or are descendants of
flying species.[146]

[edit] Land vertebrates

Main article: Tetrapod

Acanthostega changed views about the early evolution of tetrapods[147]

"Fish"

Acanthostega
 Ichthyostega

Tulerpeton

Early
amphibians

Anthracosauria

Amniotes
 Family tree of tetrapods[148]
Tetrapods, vertebrates with four limbs, evolved from other rhipidistians over a relatively short
timespan during the Late Devonian, between 370 million years ago and 360 million years ago.[149] From the
1950s to the early 1980s it was thought that tetrapods evolved from fish that had already acquired the ability
to crawl on land, possibly in order to go from a pool that was drying out to one that was deeper. However in
1987 nearly-complete fossils of Acanthostega from about 363 million years ago showed that this Late
Devonian transitional animal had legs and both lungs and gills, but could never have survived on land: its
limbs and its wrist and ankle joints were too weak to bear its weight; its ribs were too short to prevent its
lungs from being squeezed flat by its weight; its fish-like tail fin would have been damaged by dragging on
the ground. The current hypothesis is that Acanthostega, which was about 1 metre (3.3 ft) long, was a wholly
aquatic predator that hunted in shallow water. Its skeleton differed from that of most fish, in ways that
enabled it to raise its head to breathe air while its body remained submerged, including: its jaws show
modifications that would have enabled it to gulp air; the bones at the back of its skull are locked together,
providing strong attachment points for muscles that raised its head; the head is not joined to the shoulder
girdle and it has a distinct neck.[147]
The Devonian proliferation of land plants may help to explain why air-breathing would have been an
advantage: leaves falling into streams and rivers would have encouraged the growth of aquatic vegetation;
this would have attracted grazing invertebrates and small fish that preyed on them; they would have been
attractive prey but the environment was unsuitable for the big marine predatory fish; air-breathing would have
been necessary because these waters would have been short of oxygen, since warm water holds less
dissolved oxygen than cooler marine water and since the decomposition of vegetation would have used
some of the oxygen.[147]
Later discoveries revealed earlier transitional forms between Acanthostega and completely fish-like
animals.[150] Unfortunately there is then a gap of about 30 million years between the fossils of ancestral
tetrapods and Mid Carboniferous fossils of vertebrates that look well-adapted for life on land. Some of these
look like early relatives of modern amphibians, most of which need to keep their skins moist and to lay their
eggs in water, while others are accepted as early relatives of the amniotes, whose water-proof skins and
eggs enable them to live and breed far from water.[148]
[edit] Dinosaurs, birds and mammals
Main articles: Dinosaur evolution, Origin of Birds, and Evolution of mammals
Amniotes
Synapsids
Early synapsids
(extinct)

Pelycosaurs

Extinct pelycosaurs

Therapsids

Extinct therapsids

Mammaliformes

Extinct mammaliformes

Mammals
 Sauropsids

Diapsids
 Squ
(lizards and
snakes)
Archosaurs
Extinct
archosaurs

Crocodilians

Pterosaurs
(extinct)

Dinosaurs
Theropods
Extinct
theropods
Birds

Sauropods
(extinct)

Ornithischians
(extinct)
 Possible family tree of dinosaurs, birds and mammals[152][153]
Amniotes, whose eggs can survive in dry environments, probably evolved in the Late Carboniferous
period, between 330 million years ago and 314 million years ago. The earliest fossils of the two surviving
amniote groups, synapsids and sauropsids, date from around 313 million years ago.[152][153] The synapsid
pelycosaurs and their descendants the therapsids are the most common land vertebrates in the best-known
Permian fossil beds, between 229 million years ago and 251 million years ago. However at the time these
were all in temperate zones at middle latitudes, and there is evidence that hotter, drier environments nearer
the Equator were dominated by sauropsids and amphibians.[154]
The Permian-Triassic extinction wiped out almost all land vertebrates,[155] as well as the great
majority of other life.[156] During the slow recovery from this catastrophe, estimated to be 30M years,[157] a
previously obscure sauropsid group became the most abundant and diverse terrestrial vertebrates: a few
fossils of archosauriformes ("shaped like archosaurs") have been found in Late Permian rocks,[158] but by
the Mid Triassic archosaurs were the dominant land vertebrates. Dinosaurs distinguished themselves from
other archosaurs in the Late Triassic, and became the dominant land vertebrates of the Jurassic and
Cretaceous periods, between 199 million years ago and 65 million years ago.[159]
During the Late Jurassic, birds evolved from small, predatory theropod dinosaurs.[160] The first birds
inherited teeth and long, bony tails from their dinosaur ancestors,[160] but some developed horny, toothless
beaks by the very Late Jurassic[161] and short pygostyle tails by the Early Cretaceous.[162]
While the archosaurs and dinosaurs were becoming more dominant in the Triassic, the mammaliform
successors of the therapsids could only survive as small, mainly nocturnal insectivores. This apparent set-
back may actually have promoted the evolution of mammals, for example nocturnal life may have
accelerated the development of endothermy ("warm-bloodedness") and hair or fur.[163] By 195 million years
ago in the Early Jurassic there were animals that were very nearly mammals.[164] Unfortunately there is a
gap in the fossil record throughout the Mid Jurassic.[165] However fossil teeth discovered in Madagascar
indicate that true mammals existed at least 167 million years ago.[166] After dominating land vertebrate
niches for about 150 million years, the dinosaurs perished 65 million years ago in the Cretaceous–Tertiary
extinction along with many other groups of organisms.[167] Mammals throughout the time of the dinosaurs
had been restricted to a narrow range of taxa, sizes and shapes, but increased rapidly in size and diversity
after the extinction,[168][169] with bats taking to the air within 13 million years,[170] and cetaceans to the sea
within 15 million years.[171]

[edit] Flowering plants

Main articles: Flowering plant and Gymnosperm
 Gymnosperms Gymnosp
Gnetales
(gymnosperm)

Welwitschia Cycads and

The 250,000 to 400,000 species of flowering plants outnumber all other ground plants combined,
(gymnosperm) (gymnosperm)

Bennettitales

Ephedra
(gymnosperm)

Gingk

Bennettitales

Angiosperms Gnetales
(flowering plants) (gymnosperm)

Conifers
(gymnosperm)

One possible family tree of flowering plants.[172]

Another possible family tree.[173]

[edit] Social insects
Main article: Social insects
The social insects are remarkable because the great majority of individuals in each colony are sterile.
This appears contrary to basic concepts of evolution such as natural selection and the selfish gene. In fact
there are very few eusocial insect species: only 15 out of approximately 2,600 living families of insects
contain eusocial species, and it seems that eusociality has evolved independently only 12 times among
arthropods, although some eusocial lineages have diversified into several families. Nevertheless social
insects have been spectacularly successful; for example although ants and termites account for only about
2% of known insect species, they form over 50% of the total mass of insects. Their ability to control a territory
appears to be the foundation of their success.[174]
 These termite mounds have survived a bush fire.
The sacrifice of breeding opportunities by most individuals has long been explained as a
consequence of these species' unusual haplodiploid method of sex determination, which has the paradoxical
consequence that two sterile worker daughters of the same queen share more genes with each other than
they would with their offspring if they could breed.[175] However Wilson and Hölldobler argue that this
explanation is faulty: for example, it is based on kin selection, but there is no evidence of nepotism in
colonies that have multiple queens. Instead, they write, eusociality evolves only in species that are under
strong pressure from predators and competitors, but in environments where it is possible to build "fortresses";
after colonies have established this security, they gain other advantages though co-operative foraging. In
support of this explanation they cite the appearance of eusociality in bathyergid mole rats,[174] which are not
haplodiploid.[176]
The earliest fossils of insects have been found in Early Devonian rocks from about 400 million years
ago, which preserve only a few varieties of flightless insect. The Mazon Creek lagerstätten from the Late
Carboniferous, about 300 million years ago, include about 200 species, some gigantic by modern standards,
and indicate that insects had occupied their main modern ecological niches as herbivores, detritivores and
insectivores. Social termites and ants first appear in the Early Cretaceous, and advanced social bees have
been found in Late Cretaceous rocks but did not become abundant until the Mid Cenozoic.[177]

[edit] Humans
Main article: Human evolution
Modern humans evolved from a lineage of upright-walking apes that has been traced back over 6
million years ago to Sahelanthropus.[178] The first known stone tools were made about 2.5 million years ago,
apparently by Australopithecus garhi, and were found near animal bones that bear scratches made by these
tools.[179] The earliest hominines had chimp-sized brains, but there has been a fourfold increase in the last
3 million years; a statistical analysis suggests that hominine brain sizes depend almost completely on the
date of the fossils, while the species to which they are assigned has only slight influence.[180] There is a
long-running debate about whether modern humans evolved all over the world simultaneously from existing
advanced hominines or are descendants of a single small population in Africa, which then migrated all over
the world less than 200,000 years ago and
replaced previous hominine species.[181] There is
also debate about whether anatomically-modern
humans had an intellectual, cultural and
technological "Great Leap Forward" under
100,000 years ago and, if so, whether this was
due to neurological changes that are not visible in
fossils.[182]

[edit] Mass extinctions

Main article: Mass extinction
Life on earth has suffered occasional
mass extinctions at least since 542 million years
ago. Although they are disasters at the time, mass
extinctions have sometimes accelerated the
evolution of life on earth. When dominance of
particular ecological niches passes from one
group of organisms to another, it is rarely because
the new dominant group is "superior" to the old

Apparent extinction intensity, i.e. the fraction of

genera going extinct at any given time, as reconstructed
from the fossil record. (Graph not meant to include
recent epoch of Holocene extinction event)
and usually because an extinction event eliminates the old dominant group and makes way for the new one.
[183][184]
The fossil record appears to show that the gaps between mass extinctions are becoming longer and
the average and background rates of extinction are decreasing. Both of these phenomena could be
explained in one or more ways:[185]
• The oceans may have become more hospitable to life over the last 500 million years and less
vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to
greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of
eutrophication and anoxic events; and marine ecosystems became more diversified so that food
chains were less likely to be disrupted.[186][187]
• Reasonably complete fossils are very rare, most extinct organisms are represented only by
partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly
assigned parts of the same organism to different genera which were often defined solely to
accommodate these finds – the story of Anomalocaris is an example of this. The risk of this mistake is
higher for older fossils because these are often unlike parts of any living organism. Many of the
"superfluous" genera are represented by fragments which are not found again and the "superfluous"
genera appear to become extinct very quickly.[185]
 Biodiversity in the fossil record, which is
"the number of distinct genera
alive at any given time; that is, those
whose first occurrence predates and
whose last occurrence postdates that
time"[188]
shows a different trend: a fairly swift rise from
542 to 400 million years ago; a slight decline from
400 to 200 million years ago, in which the devastating
Permian–Triassic extinction event is an important factor;
and a swift rise from 200 million years ago to the
Thousands of genera
present.[188]

[edit] The present

Oxygenic photosynthesis accounts for virtually
all of the production of organic matter from non-organic
ingredients. Production is split about evenly between
land and marine plants, and phytoplankton are the
dominant marine producers.[189]
The processes that drive evolution are still
operating. Well-known examples include the changes in
coloration of the peppered moth over the last 200 years

Phanerozoic biodiversity as shown by the

fossil record
and the more recent appearance of pathogens that are resistant to antibiotics.[190][191] There is even
evidence that humans are still evolving, and possibly at an accelerating rate over the last 40,000 years.[192]

[edit] See also

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[edit] Footnotes
1. ^ Name given as in Butterfield's paper "Bangiomorpha pubescens ..." (2000). A fossil fish,
also from China, has also been named Qingshania. The name of one of these will have to change.
 2. ^ Myxozoa were thought to be an exception, but are now thought to be heavily modified
members of the Cnidaria: Jímenez-Guri, E., Philippe, H., Okamura, B. and Holland, P. W. H. (July
2007). "Buddenbrockia is a cnidarian worm". Science 317 (116): 116–118.
doi:10.1126/science.1142024. PMID 17615357.
http://www.sciencemag.org/cgi/content/abstract/317/5834/116. Retrieved 2008-09-03.

[edit] References
1. ^ Futuyma, Douglas J. (2005). Evolution. Sunderland, Massachusetts: Sinuer Associates,
Inc. ISBN 0-87893-187-2.
2. ^ a b c Nisbet, E.G., and Fowler, C.M.R. (December 7, 1999). "Archaean metabolic evolution
of microbial mats". Proceedings of the Royal Society: Biology 266 (1436): 2375.
doi:10.1098/rspb.1999.0934. - abstract with link to free full content (PDF)
3. ^ Anbar, A.; Duan, Y.; Lyons, T.; Arnold, G.; Kendall, B.; Creaser, R.; Kaufman, A.; Gordon,
G. et al. (2007). "A whiff of oxygen before the great oxidation event?". Science (New York, N.Y.) 317
(5846): 1903–1906. doi:10.1126/science.1140325. PMID 17901330. edit
4. ^ Knoll, Andrew H.; Javaux, E.J, Hewitt, D. and Cohen, P. (2006). "Eukaryotic organisms in
Proterozoic oceans". Philosophical Transactions of the Royal Society of London, Part B 361 (1470):
1023–38. doi:10.1098/rstb.2006.1843. PMID 16754612.
5. ^ a b c d e f Fedonkin, M. A. (March 2003). "The origin of the Metazoa in the light of the
Proterozoic fossil record" (PDF). Paleontological Research 7 (1): 9–41. doi:10.2517/prpsj.7.9.
http://www.vend.paleo.ru/pub/Fedonkin_2003.pdf. Retrieved 2008-09-02.
6. ^ Bonner, J.T. (1998) The origins of multicellularity. Integr. Biol. 1, 27–36
 7. ^ "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician
Period (~450-440 m.y.a.) on the basis of fossil spores" Transition of plants to land
8. ^ Algeo, T.J.; Scheckler, S. E. (1998). "Terrestrial-marine teleconnections in the Devonian:
links between the evolution of land plants, weathering processes, and marine anoxic events".
Philosophical Transactions of the Royal Society B: Biological Sciences 353 (1365): 113–130.
doi:10.1098/rstb.1998.0195.
9. ^ "Metazoa: Fossil Record". http://www.ucmp.berkeley.edu/phyla/metazoafr.html.
10.^ Shu et al.; Luo, H-L.; Conway Morris, S.; Zhang, X-L.; Hu, S-X.; Chen, L.; Han, J.; Zhu, M.
et al. (November 4, 1999). "Lower Cambrian vertebrates from south China". Nature 402: 42–46.
doi:10.1038/46965.
11.^ Hoyt, Donald F. (1997). "Synapsid Reptiles".
http://www.csupomona.edu/~dfhoyt/classes/zoo138/SYNAPSID.HTML.
12.^ Barry, Patrick L. (January 28, 2002). "The Great Dying". Science@NASA. Science and
Technology Directorate, Marshall Space Flight Center, NASA.
http://science.nasa.gov/headlines/y2002/28jan_extinction.htm. Retrieved March 26, 2009.
13.^ Tanner LH, Lucas SG & Chapman MG (2004). "Assessing the record and causes of Late
Triassic extinctions" (PDF). Earth-Science Reviews 65 (1-2): 103–139. doi:10.1016/S0012-
8252(03)00082-5. http://nmnaturalhistory.org/pdf_files/TJB.pdf. Retrieved 2007-10-22.
14.^ Benton, M.J. (2004). Vertebrate Paleontology. Blackwell Publishers. xii-452. ISBN 0-632-
05614-2.
15.^ "Amniota - Palaeos". http://www.palaeos.org/Amniota.
16.^ Fastovsky DE, Sheehan PM (2005). "The extinction of the dinosaurs in North America".
GSA Today 15 (3): 4–10. doi:10.1130/1052-5173(2005)015<4:TEOTDI>2.0.CO;2.
http://www.gsajournals.org/perlserv/?request=get-document&doi=10.1130%2F1052-
5173%282005%29015%3C4%3ATEOTDI%3E2.0.CO%3B2. Retrieved 2007-05-18.
17.^ "Dinosaur Extinction Spurred Rise of Modern Mammals". News.nationalgeographic.com.
http://news.nationalgeographic.com/news/2007/06/070620-mammals-dinos.html. Retrieved 2009-03-
08.
18.^ Van Valkenburgh, B. (1999). "Major patterns in the history of carnivorous mammals".
Annual Review of Earth and Planetary Sciences 26: 463–493. doi:10.1146/annurev.earth.27.1.463.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.27.1.463.
19.^ a b El Albani, Abderrazak; Bengtson, Stefan; Canfield, Donald E.; Bekker, Andrey;
Macchiarelli, Reberto (July 2010). "Large colonial organisms with coordinated growth in oxygenated
environments 2.1 Gyr ago". Nature 466 (7302): 100–104. doi:10.1038/nature09166. PMID 11931142.
20.^ a b
• Dalrymple, G.B. (1991). The Age of the Earth. California: Stanford University Press.
ISBN 0-8047-1569-6.
• Newman, W.L. (July 2007). "Age of the Earth". Publications Services, USGS.
http://pubs.usgs.gov/gip/geotime/age.html. Retrieved 2008-08-29.
• Dalrymple, G.B. (2001). "The age of the Earth in the twentieth century: a problem
(mostly) solved". Geological Society, London, Special Publications 190: 205–221.
doi:10.1144/GSL.SP.2001.190.01.14.
http://sp.lyellcollection.org/cgi/content/abstract/190/1/205. Retrieved 2007-09-20.
21.^ Galimov, E.M. and Krivtsov, A.M. (December 2005). "Origin of the Earth-Moon System". J.
Earth Syst. Sci. 114 (6): 593–600. doi:10.1007/BF02715942. [1]
22.^ a b Cohen, B.A., Swindle, T.D. and Kring, D.A. (December 2000). "Support for the Lunar
Cataclysm Hypothesis from Lunar Meteorite Impact Melt Ages". Science 290 (5497): 1754–1756.
doi:10.1126/science.290.5497.1754. PMID 11099411.
http://www.sciencemag.org/cgi/content/abstract/290/5497/1754. Retrieved 2008-08-31.
23.^
• "Early Earth Likely Had Continents And Was Habitable" . University of Colorado.
2005-11-17. http://www.colorado.edu/news/releases/2005/438.html. Retrieved 2009-01-11.
• Cavosie, A.J., Valley, J.W., Wilde, S. A. and the Edinburgh Ion Microprobe Facility
(July 15, 2005). "Magmatic δ18O in 4400-3900 Ma detrital zircons: A record of the alteration
and recycling of crust in the Early Archean". Earth and Planetary Science Letters 235 (3-4):
663–681. doi:10.1016/j.epsl.2005.04.028. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6V61-4GDKB05-
3&_coverDate=07%2F15%2F2005&_alid=382434001&_rdoc=1&_fmt=&_orig=search&_qd=
1&_cdi=5801&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10
&md5=be47e49c535d059be188b66c6e596dd5.
24.^ Britt, R.R. (2002-07-24). "Evidence for Ancient Bombardment of Earth". Space.com.
http://www.space.com/scienceastronomy/planetearth/earth_bombarded_020724.html. Retrieved
2006-04-15.
25.^ Valley, J.W., Peck, W.H., King, E.M. and Wilde, S.A. (April 2002). "A cool early Earth"
(PDF). Geology 30 (4): 351–354. doi:10.1130/0091-7613(2002)030<0351:ACEE>2.0.CO;2.
http://www.geology.wisc.edu/zircon/Valley2002Cool_Early_Earth.pdf. Retrieved 2008-09-13.
26.^ Dauphas, N., Robert, F. and Marty, B. (December 2000). "The Late Asteroidal and
Cometary Bombardment of Earth as Recorded in Water Deuterium to Protium Ratio". Icarus 148 (2):
508–512. doi:10.1006/icar.2000.6489.
27.^ a b Brasier, M., McLoughlin, N., Green, O. and Wacey, D. (June 2006). "A fresh look at the
fossil evidence for early Archaean cellular life" (PDF). Philosophical Transactions of the Royal
Society: Biology 361 (1470): 887–902. doi:10.1098/rstb.2006.1835. PMID 16754605. PMC 1578727.
http://physwww.mcmaster.ca/~higgsp/3D03/BrasierArchaeanFossils.pdf. Retrieved 2008-08-30.
28.^
• Schopf, J. W. (April 1993). "Microfossils of the Early Archean Apex Chert: New
Evidence of the Antiquity of Life". Science 260 (5108): 640–646.
doi:10.1126/science.260.5108.640. PMID 11539831.
http://www.sciencemag.org/cgi/content/abstract/260/5108/640. Retrieved 2008-08-30.
• Altermann, W. and Kazmierczak, J. (2003). "Archean microfossils: a reappraisal of
early life on Earth". Res Microbiol 154 (9): 611–7. doi:10.1016/j.resmic.2003.08.006.
PMID 14596897.
29.^ Mojzsis, S.J., Arrhenius, G., McKeegan, K.D., Harrison, T.M., Nutman, A.P. and Friend,
C.R.L. (November 1996). "Evidence for life on Earth before 3,800 million years ago". Nature 384
(6604): 55–59. doi:10.1038/384055a0. PMID 8900275.
http://www.nature.com/nature/journal/v384/n6604/abs/384055a0.html. Retrieved 2008-08-30.
30.^ a b Grotzinger, J.P. and Rothman, D.H. (1996). "An abiotic model for stomatolite
morphogenesis". Nature 383: 423–425. doi:10.+1038/383423a0.
31.^
• Fedo, C.M. and Whitehouse, M.J. (May 2002). "Metasomatic Origin of Quartz-
Pyroxene Rock, Akilia, Greenland, and Implications for Earth's Earliest Life". Science 296
(5572): 1448–1452. doi:10.1126/science.1070336. PMID 12029129.
http://www.sciencemag.org/cgi/content/abstract/296/5572/1448. Retrieved 2008-08-30.
• Lepland, A., van Zuilen, M.A., Arrhenius, G., Whitehouse, M.J. and Fedo, C.M.
(January 2005). "Questioning the evidence for Earth's earliest life — Akilia revisited". Geology
 33 (1): 77–79. doi:10.1130/G20890.1.
http://geology.geoscienceworld.org/cgi/content/abstract/33/1/77. Retrieved 2008-08-30.
32.^ Schopf, J. (2006). "Fossil evidence of Archaean life". Philosophical Transactions of the
Royal Society of London: B Biological Sciences 361 (1470): 869–85. doi:10.1098/rstb.2006.1834.
PMID 16754604.
33.^ Ciccarelli, F.D., Doerks, T., von Mering, C., Creevey, C.J., et al (2006). "Toward automatic
reconstruction of a highly resolved tree of life". Science 311 (5765): 1283–7.
doi:10.1126/science.1123061. PMID 16513982.
34.^ Mason, S.F. (1984). "Origins of biomolecular handedness". Nature 311 (5981): 19–23.
doi:10.1038/311019a0. PMID 6472461.
35.^ Orgel, L.E. (October 1994). "The origin of life on the earth" (PDF). Scientific American 271
(4): 76–83. doi:10.1038/scientificamerican1094-76. PMID 7524147.
http://courses.washington.edu/biol354/The%20Origin%20of%20Life%20on%20Earth.pdf. Retrieved
2008-08-30. Also available as a web page
36.^ a b c Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. p. 6.
ISBN 0632044446.
37.^ Villarreal LP, Witzany G (October 2009). "Viruses are essential agents within the roots and
stem of the tree of life". J. Theor. Biol. 262 (4): 698–710. doi:10.1016/j.jtbi.2009.10.014.
PMID 19833132.
38.^ O'Leary, M.R. (2008). Anaxagoras and the Origin of Panspermia Theory. iUniverse, Inc..
ISBN 0595495966.
39.^ a b Arrhenius, S. (1903). "The Propagation of Life in Space". Die Umschau volume=7.
Reprinted in Goldsmith, D.,, ed. The Quest for Extraterrestrial Life. University Science Books.
ISBN 0198557043.
 40.^ Hoyle, F. and Wickramasinghe, C. (1979). "On the Nature of Interstellar Grains".
Astrophysics and Space Science 66: 77–90. doi:10.1007/BF00648361.
41.^ a b Crick, F (1973). "Directed Panspermia". Icarus 19: 341–348. doi:10.1016/0019-
1035(73)90110-3.
42.^ a b c d Warmflash, D. and Weiss, B. (November 2005). "Did Life Come From Another
World?". Scientific American: 64–71. http://www.sciam.com/article.cfm?articleID=00073A97-5745-
1359-94FF83414B7F0000&pageNumber=1&catID=2. Retrieved 2008-09-02.
43.^ Ker, Than (August 2007). "Claim of Martian Life Called 'Bogus'". space.com.
http://www.space.com/news/070823_mars_life.html. Retrieved 2008-09-02.
44.^ Bennett, J. O. (2008). "What is life?". Beyond UFOs: The Search for Extraterrestrial Life
and Its Astonishing Implications for Our Future . Princeton University Press. pp. 82–85.
ISBN 0691135495. http://books.google.com/?
id=lEQKnip7St4C&pg=PA84&dq=life+earth+carbon+water. Retrieved 2009-01-11.
45.^ Schulze-Makuch, D., Irwin, L. N. (April 2006). "The prospect of alien life in exotic forms on
other worlds". Naturwissenschaften 93 (4): 155–72. doi:10.1007/s00114-005-0078-6.
PMID 16525788.
46.^ Peretó, J. (2005). "Controversies on the origin of life" (PDF). Int. Microbiol. 8 (1): 23–31.
PMID 15906258. http://www.im.microbios.org/0801/0801023.pdf. Retrieved 2007-10-07.
47.^ Szathmáry, E. (February 2005). "Life: In search of the simplest cell". Nature 433 (7025):
469–470. doi:10.1038/433469a. PMID 15690023.
http://www.nature.com/nature/journal/v433/n7025/full/433469a.html. Retrieved 2008-09-01.
48.^ Luisi, P. L., Ferri, F. and Stano, P. (2006). "Approaches to semi-synthetic minimal cells: a
review". Naturwissenschaften 93 (1): 1–13. doi:10.1007/s00114-005-0056-z. PMID 16292523.
 49.^ Joyce, G.F. (2002). "The antiquity of RNA-based evolution". Nature 418 (6894): 214–21.
doi:10.1038/418214a. PMID 12110897.
50.^ a b Hoenigsberg, H. (December 2003)). "Evolution without speciation but with selection:
LUCA, the Last Universal Common Ancestor in Gilbert’s RNA world". Genetic and Molecular
Research 2 (4): 366–375. PMID 15011140. http://www.funpecrp.com.br/gmr/year2003/vol4-
2/gmr0070_full_text.htm. Retrieved 2008-08-30. (also available as PDF)
51.^ Trevors, J. T. and Abel, D. L. (2004). "Chance and necessity do not explain the origin of
life". Cell Biol. Int. 28 (11): 729–39. doi:10.1016/j.cellbi.2004.06.006. PMID 15563395.
52.^ Forterre, P., Benachenhou-Lahfa, N., Confalonieri, F., Duguet, M., Elie, C. and Labedan, B.
(1992). "The nature of the last universal ancestor and the root of the tree of life, still open questions".
BioSystems 28 (1-3): 15–32. doi:10.1016/0303-2647(92)90004-I. PMID 1337989.
53.^ Cech, T.R. (August 2000). "The ribosome is a ribozyme". Science 289 (5481): 878–9.
doi:10.1126/science.289.5481.878. PMID 10960319.
http://www.sciencemag.org/cgi/content/short/289/5481/878. Retrieved 2008-09-01.
54.^ Johnston, W. K. et al (2001). "RNA-Catalyzed RNA Polymerization: Accurate and General
RNA-Templated Primer Extension". Science 292 (5520): 1319–1325. doi:10.1126/science.1060786.
PMID 11358999.
55.^ a b
• Levy, M. and Miller, S.L. (July 1998). "The stability of the RNA bases: implications for
the origin of life". Proc. Natl. Acad. Sci. U.S.A. 95 (14): 7933–8. doi:10.1073/pnas.95.14.7933.
PMID 9653118. PMC 20907. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=9653118.
• Larralde, R., Robertson, M. P. and Miller, S. L. (August 1995). "Rates of
decomposition of ribose and other sugars: implications for chemical evolution". Proc. Natl.
 Acad. Sci. U.S.A. 92 (18): 8158–60. doi:10.1073/pnas.92.18.8158. PMID 7667262.
PMC 41115. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=7667262.
• Lindahl, T. (April 1993). "Instability and decay of the primary structure of DNA".
Nature 362 (6422): 709–15. doi:10.1038/362709a0. PMID 8469282.
56.^ Orgel, L. (November 2000). "Origin of life. A simpler nucleic acid". Science (journal) 290
(5495): 1306–7. PMID 11185405.
57.^ Nelson, K.E., Levy, M., and Miller, S.L. (April 2000). "Peptide nucleic acids rather than RNA
may have been the first genetic molecule". Proc. Natl. Acad. Sci. U.S.A. 97 (8): 3868–71.
doi:10.1073/pnas.97.8.3868. PMID 10760258. PMC 18108. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=10760258.
58.^ Martin, W. and Russell, M.J. (2003). "On the origins of cells: a hypothesis for the
evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from
prokaryotes to nucleated cells". Philosophical Transactions of the Royal Society: Biological 358
(1429): 59–85. doi:10.1098/rstb.2002.1183. PMID 12594918.
59.^ Wächtershäuser, G. (August 2000). "Origin of life. Life as we don't know it". Science
(journal) 289 (5483): 1307–8. PMID 10979855.
60.^ Trevors, J.T. and Psenner, R. (2001). "From self-assembly of life to present-day bacteria: a
possible role for nanocells". FEMS Microbiol. Rev. 25 (5): 573–82. doi:10.1111/j.1574-
6976.2001.tb00592.x. PMID 11742692.
61.^ Segré, D., Ben-Eli, D., Deamer, D. and Lancet, D. (February-April 2001). "The Lipid World"
(PDF). Origins of Life and Evolution of Biospheres 2001 31 (1-2): 119–45.
doi:10.1023/A:1006746807104. PMID 11296516. http://ool.weizmann.ac.il/Segre_Lipid_World.pdf.
Retrieved 2008-09-01.
 62.^ Cairns-Smith, A.G. (1968). "An approach to a blueprint for a primitive organism". in
Waddington, C,H.. Towards a Theoretical Biology. 1. Edinburgh University Press. pp. 57–66
63.^ Ferris, J.P. (June 1999). "Prebiotic Synthesis on Minerals: Bridging the Prebiotic and RNA
Worlds". Biological Bulletin. Evolution: A Molecular Point of View (Biological Bulletin, Vol. 196, No. 3)
196 (3): 311–314. doi:10.2307/1542957. PMID 10390828. http://www.jstor.org/pss/1542957.
Retrieved 2008-09-01.
64.^ Hanczyc, M.M., Fujikawa, S.M. and Szostak, Jack W. (October 2003). "Experimental
Models of Primitive Cellular Compartments: Encapsulation, Growth, and Division". Science 302
(5645): 618–622. doi:10.1126/science.1089904. PMID 14576428.
http://www.sciencemag.org/cgi/content/abstract/302/5645/618. Retrieved 2008-09-01.
65.^ Hartman, H. (October 1998). "Photosynthesis and the Origin of Life". Origins of Life and
Evolution of Biospheres 28 (4–6): 512–521. http://www.springerlink.com/content/t1n325268n01217k/.
Retrieved 2008-09-01.
66.^ a b Krumbein, W.E., Brehm, U., Gerdes, G., Gorbushina, A.A., Levit, G. and Palinska, K.A.
(2003). "Biofilm, Biodictyon, Biomat Microbialites, Oolites, Stromatolites, Geophysiology, Global
Mechanism, Parahistology". in Krumbein, W.E., Paterson, D.M., and Zavarzin, G.A. (PDF). Fossil
and Recent Biofilms: A Natural History of Life on Earth . Kluwer Academic. pp. 1–28.
ISBN 1402015976. Archived from the original on January 06, 2007.
http://web.archive.org/web/20070106201614/http://134.106.242.33/krumbein/htdocs/Archive/397/Kru
mbein_397.pdf. Retrieved 2008-07-09
67.^ a b Risatti, J. B., Capman, W. C. and Stahl, D. A. (October 11, 1994). "Community structure
of a microbial mat: the phylogenetic dimension" (PDF). Proceedings of the National Academy of
Sciences 91 (21): 10173–10177. doi:10.1073/pnas.91.21.10173. PMID 7937858. PMC 44980.
http://www.pnas.org/content/91/21/10173.full.pdf. Retrieved 2008-07-09.
 68.^ (the editor) (June 2006)). "Editor's Summary: Biodiversity rocks". Nature 441.
http://www.nature.com/nature/journal/v441/n7094/edsumm/e060608-01.html. Retrieved 2009-01-10.
69.^ Allwood, A. C., Walter, M. R., Kamber, B. S., Marshall, C. P. and Burch, I. W. (June 2006)).
"Stromatolite reef from the Early Archaean era of Australia". Nature 441 (7094): 714–718.
doi:10.1038/nature04764. PMID 16760969.
http://www.nature.com/nature/journal/v441/n7094/abs/nature04764.html. Retrieved 2008-08-31.
70.^ Blankenship, R.E.; Bebout, BM; Des Marais, DJ (1 January 2001). "Molecular evidence for
the evolution of photosynthesis". Trends in Plant Science 6 (1): 4–6. doi:10.1038/35085554.
PMID 11460161. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6TD1-424KK4J-
3&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_url
Version=0&_userid=10&md5=6f38f9f1d29b24fc90d0145837338b9e. Retrieved 2008-07-14.
71.^ Hoehler, T.M., Bebout, B.M. and Des Marais, D.J. (19 July 2001). "The role of microbial
mats in the production of reduced gases on the early Earth". Nature 412 (6844): 324–327.
doi:10.1038/35085554. PMID 11460161.
http://www.nature.com/nature/journal/v412/n6844/full/412324a0.html. Retrieved 2008-07-14.
72.^ Abele, D. (7 November 2002). "Toxic oxygen: The radical life-giver". Nature 420 (27): 27.
doi:10.1038/420027a. PMID 12422197.
http://www.nature.com/nature/journal/v420/n6911/full/420027a.html. Retrieved 2008-07-14.
73.^ "Introduction to Aerobic Respiration". University of California, Davis. Archived from the
original on October 29, 2007.
http://web.archive.org/web/20071029120120/http://trc.ucdavis.edu/biosci10v/bis10v/week3/06aerobi
crespirintro.html. Retrieved 2008-07-14.
74.^ Goldblatt, C., Lenton, T.M. and Watson, A.J. (2006). "The Great Oxidation at ~2.4 Ga as a
bistability in atmospheric oxygen due to UV shielding by ozone" (PDF). Geophysical Research
Abstracts 8 (00770). http://www.cosis.net/abstracts/EGU06/00770/EGU06-J-00770.pdf. Retrieved
2008-09-01.
75.^ a b Glansdorff, N., Xu, Y. and Labedan, B. (2008). "The Last Universal Common Ancestor:
emergence, constitution and genetic legacy of an elusive forerunner". Biology Direct 3 (29): 29.
doi:10.1186/1745-6150-3-29. PMID 18613974.
76.^ a b Brocks, J. J., Logan, G. A., Buick, R. and Summons, R. E. (1999). "Archaean molecular
fossils and the rise of eukaryotes". Science 285 (5430): 1033–1036.
doi:10.1126/science.285.5430.1033. PMID 10446042.
http://www.sciencemag.org/cgi/content/abstract/285/5430/1033. Retrieved 2008-09-02.
77.^ a b Hedges, S. B., Blair, J. E., Venturi, M. L. and Shoe, J. L (January 2004). "A molecular
timescale of eukaryote evolution and the rise of complex multicellular life". BMC Evolutionary Biology
4 (2): 2. doi:10.1186/1471-2148-4-2. PMID 15005799. PMC 341452.
http://www.biomedcentral.com/1471-2148/4/2/abstract/. Retrieved 2008-07-14.
78.^ Burki, F., Shalchian-Tabrizi, K., Minge, M., Skjæveland, Å., Nikolaev, S. I. et al. (2007).
"Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE 2 (8): e790.
doi:10.1371/journal.pone.0000790. PMID 17726520.
79.^ Parfrey, L. W., Barbero, E., Lasser, E., Dunthorn, M., Bhattacharya, D., Patterson, D.J. and
Katz, L.A. (December 2006). "Evaluating Support for the Current Classification of Eukaryotic
Diversity". PLoS Genetics 2 (12): e220. doi:10.1371/journal.pgen.0020220. PMID 17194223.
80.^ Margulis, L. (1981). Symbiosis in cell evolution. San Francisco: W.H. Freeman.
ISBN 0716712563.
81.^ Vellai, T. and Vida, G. (1999). "The origin of eukaryotes; the difference between eukaryotic
and prokaryotic cells". Proceedings of the Royal Society: Biology 266 (1428): 1571–1577.
doi:10.1098/rspb.1999.0817. PMID 10467746.
 82.^ Selosse, M-A., Abert, B., and Godelle, B. (2001). "Reducing the genome size of organelles
favours gene transfer to the nucleus". Trends in ecology & evolution 16 (3): 135–141.
doi:10.1016/S0169-5347(00)02084-X. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6VJ1-429XTFM-
H&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=8370ca16bcde45bfa1c050068a2d6e19. Retrieved 2008-09-02.
83.^ Pisani, D., Cotton, J.A. and McInerney, J.O. (2007). "Supertrees disentangle the chimerical
origin of eukaryotic genomes". Mol Biol Evol. 24 (8): 1752–60. doi:10.1093/molbev/msm095.
PMID 17504772.
84.^ Gray, M.W., Burger, G., and Lang, B.F. (1999). "Mitochondrial evolution". Science 283
(5407): 1476–1481. doi:10.1126/science.283.5407.1476. PMID 10066161.
http://www.sciencemag.org/cgi/content/abstract/283/5407/1476. Retrieved 2008-09-02.
85.^ Rasmussen, B., Fletcher, I.R., Brocks, J.R. and Kilburn, M.R. (October 2008).
"Reassessing the first appearance of eukaryotes and cyanobacteria". Nature 455 (7216): 1101–1104.
doi:10.1038/nature07381. PMID 18948954.
86.^ Han, T.M. and Runnegar, B. (July 1992). "Megascopic eukaryotic algae from the 2.1-billion-
year-old negaunee iron-formation, Michigan". Science 257 (5067): 232–235.
doi:10.1126/science.1631544. PMID 1631544.
http://www.sciencemag.org/cgi/content/abstract/257/5067/232. Retrieved 2008-09-02.
87.^ Javaux, E. J., Knoll, A. H. and Walter, M. R. (September 2004). "TEM evidence for
eukaryotic diversity in mid-Proterozoic oceans". Geobiology 2 (3): 121–132. doi:10.1111/j.1472-
4677.2004.00027.x. http://www3.interscience.wiley.com/journal/118814335/abstract. Retrieved
2008-09-02.
 88.^ a b Butterfield, N. J. (2005). "Probable Proterozoic fungi". Paleobiology 31 (1): 165–182.
doi:10.1666/0094-8373(2005)031<0165:PPF>2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/content/abstract/31/1/165. Retrieved 2008-09-02.
89.^ a b Nakagaki, T., Yamada, H. and Tóth, Á. (September 2000). "Intelligence: Maze-solving
by an amoeboid organism". Nature 407 (6803): 470. doi:10.1038/35035159. PMID 11028990.
http://www.nature.com/nature/journal/v407/n6803/abs/407470a0.html. Retrieved 2008-09-03.
90.^ Bell, G. and Mooers, A.O. (1968). "Size and complexity among multicellular organisms".
Biological Journal of the Linnean Society 60 (3): 345–363. doi:10.1111/j.1095-8312.1997.tb01500.x.
http://www3.interscience.wiley.com/journal/119168103/abstract. Retrieved 2008-09-03.
91.^ Kaiser, D. (2001). "Building a multicellular organism". Annual Review of Genetics 35: 103–
123. doi:10.1146/annurev.genet.35.102401.090145. PMID 11700279.
92.^ a b Bonner, J. T. (January 1999). "The Origins of Multicellularity". Integrative Biology 1 (1):
27–36. doi:10.1002/(SICI)1520-6602(1998)1:1<27::AID-INBI4>3.0.CO;2-6.
http://doi.wiley.com/10.1002/(SICI)1520-6602(1998)1:1%3C27::AID-INBI4%3E3.0.CO;2-6. Retrieved
2008-09-03.
93.^ a b c d e Butterfield, N. J. (September 2000). "Bangiomorpha pubescens n. gen., n. sp.:
implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic
radiation of eukaryotes". Paleobiology 26 (3): 386–404. doi:10.1666/0094-
8373(2000)026<0386:BPNGNS>2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/content/abstract/26/3/386. Retrieved 2008-09-02.
94.^ a b c d e Jokela, J. (2001). "Sex: Advantage". Encyclopedia of Life Sciences. John Wiley &
Sons, Ltd.. doi:10.1038/npg.els.0001716
 95.^ a b Holmes, R.K. and Jobling, M.G. (1996). "Genetics: Exchange of Genetic Information". in
Baron, S.. Baron's Medical Microbiology (4th ed.). Galveston: University of Texas Medical Branch.
ISBN 0-9631172-1-1. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?
highlight=conjugation&rid=mmed.section.468. Retrieved 2008-09-02
96.^ Christie, P. J. (April 2001). "Type IV secretion: intercellular transfer of macromolecules by
systems ancestrally related to conjugation machines". Molecular Microbiology 40 (22): 294–305.
doi:10.1046/j.1365-2958.2001.02302.x. http://lib.bioinfo.pl/meid:10183. Retrieved 2008-09-02.
97.^ Ramesh, M. A., Malik, S-B. and Logsdon, J. M. Jr. (January 2005). "A phylogenomic
inventory of meiotic genes; evidence for sex in Giardia and an early eukaryotic origin of meiosis"
(PDF). Current Biology 15 (2): 185–91. doi:10.1016/j.cub.2005.01.003. PMID 15668177.
http://euplotes.biology.uiowa.edu/web/jmlpubls/rml05.pdf. Retrieved 2008-12-22.
98.^ a b Otto, S. P., and Gerstein, A. C. (2006). "Why have sex? The population genetics of sex
and recombination". Biochemical Society Transactions 34 (Pt 4): 519–522. doi:10.1042/BST0340519.
PMID 16856849. http://www.biochemsoctrans.org/bst/034/0519/bst0340519.htm. Retrieved 2008-12-
22.
99.^ Bernstein, H., Byerly, H., Hopf, F. and Michod, R. (1984). "Origin of sex". Journal of
Theoretical Biology 110 (3): 323–351. doi:10.1016/S0022-5193(84)80178-2. PMID 6209512.
100.^ a b c Judson, O. (2002). Dr. Tatiana's sex advice to all creation. New York: Metropolitan
Books. pp. 233–4. ISBN 0-8050-6331-5.
101.^ Hickey, D. (1982). "Selfish DNA: a sexually-transmitted nuclear parasite". Genetics 101
(3–4): 519–31. PMID 6293914.
102.^ DasSarma, S. (2007). Extreme Microbes. "Extreme Microbes". American Scientist 95:
224–231.
 103.^ Sterrer, W. (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology
216 (4): 387–96. doi:10.1006/jtbi.2002.3008. PMID 12151256. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6WMD-46DM0JD-
1&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=3f5b76ba0b7fff999ee670479f06e5af. Retrieved 2008-08-02.
104.^ Christie, P. J. (April 2001). "Type IV secretion: intercellular transfer of macromolecules by
systems ancestrally related to conjugation machines". Molecular Microbiology 40 (22): 294–305.
doi:10.1046/j.1365-2958.2001.02302.x. http://lib.bioinfo.pl/meid:10183. Retrieved 2008-09-02.
105.^ a b Dong, L., Xiao, S., Shen, B. and Zhou, C. (January 2008). "Silicified Horodyskia and
Palaeopascichnus from upper Ediacaran cherts in South China: tentative phylogenetic interpretation
and implications for evolutionary stasis". Journal of the Geological Society 165: 367–378.
doi:10.1144/0016-76492007-074.
http://findarticles.com/p/articles/mi_qa3721/is_200801/ai_n24394476/pg_1?tag=artBody;col1.
Retrieved 2008-09-02.
106.^ Dickey, Gwyneth. "African fossils suggest complex life arose early", Science News,
Washington, D.C., Wednesday, June 30th, 2010. Retrieved on 2010-07-02.
107.^ Gaidos, E., Dubuc, T., Dunford, M., McAndrew, P., Padilla-gamiño, J., Studer, B.,
Weersing, K. and Stanley, S. (2007). "The Precambrian emergence of animal life: a geobiological
perspective" (PDF). Geobiology 5: 351. doi:10.1111/j.1472-4669.2007.00125.x.
http://www.soest.hawaii.edu/GG/FACULTY/GAIDOS/geobiology2007.pdf. Retrieved 2008-09-03.
108.^ Davidson, M.W.. "Animal Cell Structure". Florida State University.
http://micro.magnet.fsu.edu/cells/animalcell.html. Retrieved 2008-09-03.
109.^ Saupe, S.G. "Concepts of Biology". College of St. Benedict / St. John's University.
http://employees.csbsju.edu/SSAUPE/biol116/Zoology/digestion.htm. Retrieved 2008-09-03.
 110.^ Hinde, R. T. (1998). "The Cnidaria and Ctenophora". in Anderson, D.T.,. Invertebrate
Zoology. Oxford University Press. pp. 28–57. ISBN 0195513681.
111.^ Chen, J.-Y., Oliveri, P., Gao, F., Dornbos, S.Q., Li, C-W., Bottjer, D.J. and Davidson, E.H.
(August 2002). "Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from
Southwest China" (PDF). Developmental Biology 248 (1): 182–196. doi:10.1006/dbio.2002.0714.
PMID 12142030. http://www.uwm.edu/~sdornbos/PDF's/Chen%20et%20al.%202002.pdf. Retrieved
2008-09-03.
112.^ Grazhdankin, D. (2004). "Patterns of distribution in the Ediacaran biotas: facies versus
biogeography and evolution". Paleobiology 30: 203. doi:10.1666/0094-
8373(2004)030<0203:PODITE>2.0.CO;2. ISSN 0094–8373.
113.^ Seilacher, A. (1992). "Vendobionta and Psammocorallia: lost constructions of
Precambrian evolution" (abstract). Journal of the Geological Society, London 149 (4): 607–613.
doi:10.1144/gsjgs.149.4.0607. ISSN 0016–7649.
http://jgs.lyellcollection.org/cgi/content/abstract/149/4/607. Retrieved 2007-06-21.
114.^ Martin, M.W.; Grazhdankin, D. V., Bowring, S. A., Evans, D. A. D., Fedonkin, M. A. and
Kirschvink, J. L. (2000-05-05). "Age of Neoproterozoic Bilaterian Body and Trace Fossils, White Sea,
Russia: Implications for Metazoan Evolution" (abstract). Science 288 (5467): 841.
doi:10.1126/science.288.5467.841. PMID 10797002.
http://www.scienceonline.org/cgi/content/abstract/288/5467/841. Retrieved 2008-07-03.
115.^ Fedonkin, M. A. and Waggoner, B. (1997). "The late Precambrian fossil Kimberella is a
mollusc-like bilaterian organism" (abstract). Nature 388: 868–871. doi:10.1038/42242.
http://www.nature.com/nature/journal/v388/n6645/abs/388868a0.html. Retrieved 2008-07-03.
 116.^ Mooi, R. and Bruno, D. (1999). "Evolution within a bizarre phylum: Homologies of the first
echinoderms" (PDF). American Zoologist 38: 965–974.
http://icb.oxfordjournals.org/cgi/reprint/38/6/965.pdf. Retrieved 2007-11-24.
117.^ McMenamin, M. A. S (2003). "Spriggina is a trilobitoid ecdysozoan" (abstract). Abstracts
with Programs (Geological Society of America) 35 (6): 105.
http://gsa.confex.com/gsa/2003AM/finalprogram/abstract_62056.htm. Retrieved 2007-11-24.
118.^ Lin, J. P.; Gon, S.M.; Gehling, J.G.; Babcock, L.E.; Zhao, Y.L.; Zhang, X.L.; Hu, S.X.;
Yuan, J.L.; Yu, M.Y.; Peng, J. (2006). "A Parvancorina-like arthropod from the Cambrian of South
China". Historical Biology 18 (1): 33–45. doi:10.1080/08912960500508689. edit
119.^ Butterfield, N. J. (2006). "Hooking some stem-group "worms": fossil lophotrochozoans in
the Burgess Shale". Bioessays 28 (12): 1161–6. doi:10.1002/bies.20507. PMID 17120226.
120.^ a b c Bengtson, S. (2004). "Early skeletal fossils". in Lipps, J.H., and Waggoner, B.M.
(PDF). Neoproterozoic - Cambrian Biological Revolutions. 10. 67–78.
http://www.cosmonova.org/download/18.4e32c81078a8d9249800021554/Bengtson2004ESF.pdf.
Retrieved 2008-07-18
121.^ Gould, S. J. (1989). Wonderful Life. Hutchinson Radius. pp. 124–136 and many others.
ISBN 0091742714.
122.^ Gould, S. J. (1989). Wonderful Life: The Burgess Shale and the Nature of History . W.W.
Norton & Company. ISBN 039330700X.
123.^ Budd, G. E. (2003). "The Cambrian Fossil Record and the Origin of the Phyla" (Free full
text). Integrative and Comparative Biology 43 (1): 157–165. doi:10.1093/icb/43.1.157. http://intl-
icb.oxfordjournals.org/cgi/content/abstract/43/1/157. Retrieved 2008-07-15.
124.^ Budd, G. E. (1996). "The morphology of Opabinia regalis and the reconstruction of the
arthropod stem-group". Lethaia 29 (1): 1–14. doi:10.1111/j.1502-3931.1996.tb01831.x.
 125.^ Marshall, C. R. (2006). "Explaining the Cambrian “Explosion” of Animals". Annu. Rev.
Earth Planet. Sci. 34: 355–384. doi:10.1146/annurev.earth.33.031504.103001.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.33.031504.103001?
journalCode=earth. Retrieved 2007-11-06.
126.^ Janvier, P. (2001). "Vertebrata (Vertebrates)". Encyclopedia of Life Sciences. Wiley
InterScience. doi:10.1038/npg.els.0001531
127.^ Conway Morris, S. (August 2, 2003). "Once we were worms". New Scientist 179 (2406):
34. http://cas.bellarmine.edu/tietjen/Evolution/once_we_were_worms.htm. Retrieved 2008-09-05.
128.^ Shu, D-G., Luo, H-L., Conway Morris, S., Zhang, X-L., Hu, S-X., Chen, L., J. Han, J., Zhu,
M., Li, Y. and Chen, L-Z. (November 1999). "Lower Cambrian vertebrates from south China" (PDF).
Nature 402: 42–46. doi:10.1038/46965. http://www.bios.niu.edu/davis/bios458/Shu1.pdf. Retrieved
2008-09-05.
129.^ Shu, D.-G., Conway Morris, S., Han, J., Zhang, Z.-F., Yasui, K., Janvier, P., Chen, L.,
Zhang, X.-L., Liu, J.-N., Li, Y. and Liu, H.-Q. (January 2003). "Head and backbone of the Early
Cambrian vertebrate Haikouichthys". Nature 421 (6922): 526–529. doi:10.1038/nature01264.
PMID 12556891. http://www.nature.com/nature/journal/v421/n6922/abs/nature01264.html. Retrieved
2008-09-05.
130.^ Sansom I. J., Smith, M. M. and Smith, M. P. (2001). "The Ordovician radiation of
vertebrates". in Ahlberg, P.E.. Major Events in Early Vertebrate Evolution. Taylor and Francis.
pp. 156–171. ISBN 0-415-23370-4
131.^ a b Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. pp. 120–122.
ISBN 0632044446.
132.^ a b c Selden, P. A. (2001). ""Terrestrialization of Animals"". in Briggs, D.E.G., and
Crowther, P.R.. Palaeobiology II: A Synthesis. Blackwell. pp. 71–74. ISBN 0632051493.
http://books.google.com/?id=AHsrhGOTRM4C&pg=PA71&lpg=PA71&dq=
%22Terrestrialization+of+Animals%22+selden. Retrieved 2008-09-05
133.^ a b c d e f g Shear, W.A. (2000). "The Early Develpoment of Terrestrial Ecosystems". in
Gee, H.. Shaking the Tree: Readings from Nature in the History of Life . University of Chicago Press.
pp. 169–184. ISBN 0226284964. http://books.google.com/?id=ZJe_Dmdbm-
QC&pg=PA233&dq=evolution+flowering+plant+angiosperm. Retrieved 2008-09-09
134.^ a b Hawksworth, D.L. (2001). "Lichens". Encyclopedia of Life Sciences. John Wiley &
Sons, Ltd.. doi:10.1038/npg.els.0000368
135.^ Retallack, G.J.; Feakes, C.R. (1987). "Trace Fossil Evidence for Late Ordovician Animals
on Land". Science 235 (4784): 61–63. doi:10.1126/science.235.4784.61. PMID 17769314.
136.^ a b Kenrick, P. and Crane, P. R. (September 1997). "The origin and early evolution of
plants on land" (PDF). Nature 389: 33. doi:10.1038/37918.
http://botit.botany.wisc.edu/courses/botany_940/06EvidEvol/papers/KendrickCrane1997.pdf.
Retrieved 2008-09-05.
137.^ Scheckler, S. E. (2001). ""Afforestation – the First Forests"". in Briggs, D.E.G., and
Crowther, P.R.. Palaeobiology II: A Synthesis. Blackwell. pp. 67–70. ISBN 0632051493.
http://books.google.com/?
id=AHsrhGOTRM4C&pg=PA69&lpg=PA69&dq=devonian+meandering+plants+trees. Retrieved
2008-09-05
138.^ The phrase "Late Devonian wood crisis" is used at "Palaeos – Tetrapoda: Acanthostega".
PALAEOS: The Trace of Life on Earth.
http://www.palaeos.com/Vertebrates/Units/150Tetrapoda/150.150.html. Retrieved 2008-09-05.
139.^ a b Algeo, T. J. and Scheckler, S. E. (1998). "Terrestrial-marine teleconnections in the
Devonian: links between the evolution of land plants, weathering processes, and marine anoxic
events". Philosophical Transactions of the Royal Society: Biology 353: 113–130.
doi:10.1098/rstb.1998.0195.
140.^ Taylor T. N. and Osborn J. M. (1996). "The importance of fungi in shaping the
paleoecosystem". Review of Paleobotany and Palynology 90: 249–262. doi:10.1016/0034-
6667(95)00086-0. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6V6W-454YDFK-
7&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_url
Version=0&_userid=10&md5=9d5d008d99d044684e947ad74b05514d. Retrieved 2008-09-05.
141.^ MacNaughton, R. B., Cole, J. M., Dalrymple, R. W., Braddy, S. J., Briggs, D. E. G. and
Lukie, T. D. (May 2002). "First steps on land: Arthropod trackways in Cambrian-Ordovician eolian
sandstone, southeastern Ontario, Canada". Geology 30 (5): 391–394. doi:10.1130/0091-
7613(2002)030<0391:FSOLAT>2.0.CO;2.
http://geology.geoscienceworld.org/cgi/content/abstract/30/5/391. Retrieved 2008-09-05.
142.^ Vaccari, N. E., Edgecombe, G. D. and Escudero, C. (2004). "Cambrian origins and
affinities of an enigmatic fossil group of arthropods". Nature 430 (6999): 554–557.
doi:10.1038/nature02705. PMID 15282604.
143.^ Buatois, L. A., Mangano, M. G., Genise, J. F. and Taylor, T. N. (June 1998). "The
ichnologic record of the continental invertebrate invasion; evolutionary trends in environmental
expansion, ecospace utilization, and behavioral complexity". PALAIOS (PALAIOS, Vol. 13, No. 3) 13
(3): 217–240. doi:10.2307/3515447. http://palaios.sepmonline.org/cgi/content/abstract/13/3/217.
Retrieved 2008-09-05.
144.^ Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. p. 126. ISBN 0632044446.
145.^ Grimaldi, D. and Engel, M. (2005). "Insects Take to the Skies". Evolution of the Insects.
Cambridge University Press. pp. 155–160. ISBN 0521821495. http://books.google.com/?
id=Ql6Jl6wKb88C&dq=%22Evolution+of+the+Insects%22+grimaldi&printsec=frontcover. Retrieved
2009-01-11.
146.^ Grimaldi, D. and Engel, M. (2005). "Diversity of evolution". Evolution of the Insects.
Cambridge University Press. p. 12. ISBN 0521821495. http://books.google.com/?
id=Ql6Jl6wKb88C&dq=%22Evolution+of+the+Insects%22+grimaldi&printsec=frontcover. Retrieved
2009-01-11.
147.^ a b c Clack, J. A. (November, 2005). "Getting a Leg Up on Land". Scientific American.
http://www.sciam.com/article.cfm?id=getting-a-leg-up-on-land. Retrieved 2008-09-06.
148.^ a b Ahlberg, P. E. and Milner, A. R. (April 1994). "The Origin and Early Diversification of
Tetrapods". Nature 368: 507–514. doi:10.1038/368507a0.
http://www.nature.com/nature/journal/v368/n6471/abs/368507a0.html. Retrieved 2008-09-06.
149.^ Gordon, M. S., Graham, J. B. and Wang, T. (September/October 2004). "Revisiting the
Vertebrate Invasion of the Land". Physiological and Biochemical Zoology 77 (5): 697–699.
doi:10.1086/425182.
150.^ Daeschler, E. B., Shubin, N. H. and Jenkins, F. A. (April 2006). "A Devonian tetrapod-like
fish and the evolution of the tetrapod body plan" (PDF). Nature 440 (7085): 757–763.
doi:10.1038/nature04639. PMID 16598249. http://www.com.univ-
mrs.fr/~boudouresque/Publications_DOM_2006_2007/Daeschler_et_al_2006_Nature.pdf. Retrieved
2008-09-06.
151.^ Debraga, M. and Rieppel, O. (July 1997). "Reptile phylogeny and the interrelationships of
turtles". Zoological Journal of the Linnean Society 120 (3): 281–354. doi:10.1111/j.1096-
3642.1997.tb01280.x. http://www3.interscience.wiley.com/journal/119830935/abstract. Retrieved
2008-09-07.
 152.^ a b Benton M. J. and Donoghue, P. C. J. (2007). "Paleontological Evidence to Date the
Tree of Life". Molecular Biology and Evolution 24 (1): 26–53. doi:10.1093/molbev/msl150.
PMID 17047029. http://mbe.oxfordjournals.org/cgi/content/full/24/1/26. Retrieved 2008-09-07.
153.^ a b Benton, M. J. (May 1990). "Phylogeny of the Major Tetrapod Groups: Morphological
Data and Divergence Dates". Journal of Molecular Evolution 30 (5): 409–424.
doi:10.1007/BF02101113. PMID 2111854. http://www.springerlink.com/content/k152294003652458/.
Retrieved 2008-09-07.
154.^ Sidor, C. A., O'Keefe, F. R., Damiani, R., Steyer, J. S., Smith, R. M. H., Larsson, H. C. E.,
Sereno, P. C., Ide, O., and Maga, A. (April 2005). "Permian tetrapods from the Sahara show climate-
controlled endemism in Pangaea". Nature 434 (7035): 886–889. doi:10.1038/nature03393.
PMID 15829962. http://www.nature.com/nature/journal/v434/n7035/full/nature03393.html. Retrieved
2008-09-08.
155.^ Smith, R. and Botha, J. (September-October 2005). "The recovery of terrestrial vertebrate
diversity in the South African Karoo Basin after the end-Permian extinction". Comptes Rendus
Palevol 4 (6-7): 623–636. doi:10.1016/j.crpv.2005.07.005. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6X1G-4GYH7VN-
1&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=add24b0622f2aff0b41e7c42a3160fa7. Retrieved 2008-09-08.
156.^ Benton, M. J. (2005). When Life Nearly Died: The Greatest Mass Extinction of All Time .
Thames & Hudson. ISBN 978-0500285732.
157.^ Sahney, S. and Benton, M.J. (2008). "Recovery from the most profound mass extinction of
all time" (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759.
doi:10.1098/rspb.2007.1370. PMID 18198148. PMC 2596898.
http://journals.royalsociety.org/content/qq5un1810k7605h5/fulltext.pdf.
 158.^ Gauthier, J., Cannatella, D. C., de Queiroz, K., Kluge, A. G. and Rowe, T. (1989).
"Tetrapod Phylogeny". in B. Fernholm, B., Bremer K., and Jörnvall, H. (PDF). The Hierarchy of Life.
Elsevier Science. p. 345. http://si-
pddr.si.edu/dspace/bitstream/10088/4689/1/VZ_1989GauthieretalHierLife.pdf. Retrieved 2008-09-08
159.^ Benton, M. J. (March 1983). "Dinosaur Success in the Triassic: a Noncompetitive
Ecological Model" (PDF). Quarterly Review of Biology 58 (1).
http://palaeo.gly.bris.ac.uk/Benton/reprints/1983success.pdf. Retrieved 2008-09-08
160.^ a b Padian, K. (2004). "Basal Avialae". in Weishampel, David B.; Dodson, Peter; &
Osmólska, Halszka (eds.). The Dinosauria (Second ed.). Berkeley: University of California Press.
pp. 210–231. ISBN 0-520-24209-2.
161.^ Hou, L., Zhou, Z., Martin, L. D. and Feduccia, A. (October 2002). "A beaked bird from the
Jurassic of China". Nature 377: 616–618. doi:10.1038/377616a0.
http://www.nature.com/nature/journal/v377/n6550/abs/377616a0.html. Retrieved 2008-09-08.
162.^ Clarke, J. A., Zhou, Z. and Zhang, F. (2006). "Insight into the evolution of avian flight from
a new clade of Early Cretaceous ornithurines from China and the morphology of Yixianornis grabaui".
Journal of Anatomy 208 (3): 287–308. doi:10.1111/j.1469-7580.2006.00534.x. PMID 16533313.
PMC 2100246. http://www3.interscience.wiley.com/journal/118559634/abstract?
CRETRY=1&SRETRY=0. Retrieved 2008-09-08.
163.^ Ruben, J. A. and Jones, T. D. (2000). "Selective Factors Associated with the Origin of Fur
and Feathers". American Zoologist 40 (4): 585–596. doi:10.1093/icb/40.4.585.
http://icb.oxfordjournals.org/cgi/content/full/40/4/585.
164.^ Luo, Z-X., Crompton, A. W. and Sun, A-L. (May 2001). "A New Mammaliaform from the
Early Jurassic and Evolution of Mammalian Characteristics". Science 292 (5521): 1535–1540.
doi:10.1126/science.1058476. PMID 11375489.
http://www.sciencemag.org/cgi/content/full/292/5521/1535. Retrieved 2008-09-08.
165.^ Cifelli, R.L. (November 2001). "Early mammalian radiations". Journal of Paleontology 75:
1214. doi:10.1666/0022-3360(2001)075<1214:EMR>2.0.CO;2.
http://findarticles.com/p/articles/mi_qa3790/is_200111/ai_n8958762/pg_6.
166.^ Flynn, J. J., Parrish, J. M. Rakotosamimanana, B., Simpson, W. F. and Wyss, A.R.
(September 1999). "A Middle Jurassic mammal from Madagascar". Nature 401: 57–60.
doi:10.1038/43420. http://www.nature.com/nature/journal/v401/n6748/abs/401057a0.html. Retrieved
2008-09-08.
167.^ MacLeod, N., Rawson, P. F., Forey, P. L., Banner. F. T., Boudagher-Fadel, M. K., Bown,
P. R., Burnett, J. A., Chambers, P., Culver, S., Evans, S. E., Jeffery, C., Kaminski, M. A., Lord, A. R.,
Milner, A. C., Milner, A. R., Morris, N., Owen, E., Rosen, B. R., ,Smith, A. B., Taylor, P. D., Urquhart,
E. and Young, J. R. (1997). "The Cretaceous–Tertiary biotic transition". Journal of the Geological
Society 154 (2): 265–292. doi:10.1144/gsjgs.154.2.0265.
http://findarticles.com/p/articles/mi_qa3721/is_199703/ai_n8738406/print.
168.^ Alroy, J. (March 1999). "The fossil record of North American mammals: evidence for a
Paleocene evolutionary radiation". Systematic biology 48 (1): 107–18.
doi:10.1080/106351599260472. PMID 12078635.
169.^ Archibald, J. D. and Deutschman, D. H. (June 2001). "Quantitative Analysis of the Timing
of the Origin and Diversification of Extant Placental Orders". Journal of Mammalian Evolution 8 (2):
107–124. doi:10.1023/A:1011317930838.
http://www.ingentaconnect.com/content/klu/jomm/2001/00000008/00000002/00342277. Retrieved
2008-09-24.
 170.^ Simmons, N. B., Seymour, K. L., Habersetzer, J. and Gunnell, G. F. (February 2008).
"Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation". Nature 451
(7180): 818–821. doi:10.1038/nature06549. PMID 18270539.
171.^ Thewissen, J. G. M., Madar, S. I. and Hussain, S. T. (1996). " Ambulocetus natans, an
Eocene cetacean (Mammalia) from Pakistan". Courier Forschungsinstitut Senckenberg 191: 1–86.
ISBN 978-3-510-61084-6.
172.^ a b c d Crane, P. R., Friis, E. M. and Pedersen, K. R. (2000). "The Origin and Early
Diversification of Angiosperms". in Gee, H.. Shaking the Tree: Readings from Nature in the History of
Life. University of Chicago Press. pp. 233–250. ISBN 0226284964. http://books.google.com/?
id=ZJe_Dmdbm-QC&pg=PA233&dq=evolution+flowering+plant+angiosperm. Retrieved 2008-09-09
173.^ a b c d Crepet, W. L. (November 2000). "Progress in understanding angiosperm history,
success, and relationships: Darwin’s abominably "perplexing phenomenon"". Proceedings of the
National Academy of Sciences 97 (24): 12939–12941. doi:10.1073/pnas.97.24.12939.
PMID 11087846. PMC 34068. http://www.pnas.org/content/97/24/12939.full.pdf+html. Retrieved
2008-09-09.
174.^ a b Wilson, E. O. and Hölldobler, B. (September 2005). "Eusociality: Origin and
consequences". Proceedings of the National Academy of Sciences 102 (38): 13367–13371.
doi:10.1073/pnas.0505858102. PMID 16157878. PMC 1224642.
http://www.pnas.org/content/102/38/13367.full.pdf+html. Retrieved 2008-09-07.
175.^ Hughes, W. O. H., Oldroyd, B. P., Beekman, M. and Ratnieks, F. L. W. (2008-05-30).
"Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality" (html). Science
(American Association for the Advancement of Science) 320 (5880): 1213–1216.
doi:10.1126/science.1156108. PMID 18511689.
http://www.sciencemag.org/cgi/content/abstract/320/5880/1213. Retrieved 2008-08-04.
 176.^ Lovegrove, B. G. (January 1991). "The evolution of eusociality in molerats (Bathyergidae):
a question of risks, numbers, and costs". Behavioral Ecology and Sociobiology 28 (1): 37–45.
doi:10.1007/BF00172137. http://www.springerlink.com/content/k4n52v522l816125/. Retrieved 2008-
09-07.
177.^ Labandeira, C. and Eble, G. J. (2000). "The Fossil Record of Insect Diversity and
Disparity". in Anderson, J., Thackeray, F., van Wyk, B., and de Wit, M. (PDF). Gondwana Alive:
Biodiversity and the Evolving Biosphere. Witwatersrand University Press.
http://www.santafe.edu/research/publications/workingpapers/00-08-044.pdf. Retrieved 2008-09-07
178.^ Brunet, M., Guy, F., Pilbeam, D., Mackaye, H. T. et al (July 2002). "A new hominid from
the Upper Miocene of Chad, Central Africa". Nature 418 (6894): 145–151. doi:10.1038/nature00879.
PMID 12110880. http://www.nature.com/nature/journal/v418/n6894/abs/nature00879.html. Retrieved
2008-09-09.
179.^ de Heinzelin, J., Clark, J. D., White, T. et al (April 1999). "Environment and Behavior of
2.5-Million-Year-Old Bouri Hominids". Science 284 (5414): 625–629.
doi:10.1126/science.284.5414.625. PMID 10213682.
http://www.sciencemag.org/cgi/content/full/sci;284/5414/625. Retrieved 2008-09-09.
180.^ De Miguel, C. and Henneberg, M. (2001). "Variation in hominid brain size: How much is
due to method?". HOMO - Journal of Comparative Human Biology 52 (1): 3–58. doi:10.1078/0018-
442X-00019. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B7GW4-4DPCHXC-
2&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=aef79dbca1f189c885cfe9f36636b131. Retrieved 2008-09-09.
181.^ Leakey, Richard (1994). The Origin of Humankind. Science Masters Series. New York,
NY: Basic Books. pp. 87–89. ISBN 0465053130.
 182.^ Mellars, Paul (2006). "Why did modern human populations disperse from Africa ca.
60,000 years ago?". Proceedings of the National Academy of Sciences 103 (25): 9381.
doi:10.1073/pnas.0510792103. PMID 16772383. PMC 1480416.
http://www.pnas.org/cgi/reprint/0510792103v1.
183.^ Benton, M. J. (2004). "6. Reptiles Of The Triassic". Vertebrate Palaeontology (3rd ed.).
Blackwell. ISBN 978-0-632-05637-8. http://www.blackwellpublishing.com/book.asp?
ref=0632056371.
184.^ Van Valkenburgh, B. (1999). "Major patterns in the history of xarnivorous mammals".
Annual Review of Earth and Planetary Sciences 26: 463–493. doi:10.1146/annurev.earth.27.1.463.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.27.1.463.
185.^ a b MacLeod, N. (2001-01-06). "Extinction!".
http://www.firstscience.com/home/articles/earth/extinction-page-2-1_1258.html. Retrieved 2008-09-
11.
186.^ Martin, R. E. (1995). "Cyclic and secular variation in microfossil biomineralization: clues to
the biogeochemical evolution of Phanerozoic oceans". Global and Planetary Change 11 (1): 1.
doi:10.1016/0921-8181(94)00011-2.
187.^ Martin, R.E. (1996). "Secular increase in nutrient levels through the Phanerozoic:
Implications for productivity, biomass, and diversity of the marine biosphere". PALAIOS (PALAIOS,
Vol. 11, No. 3) 11 (3): 209–219. doi:10.2307/3515230. http://jstor.org/stable/3515230.
188.^ a b Rohde, R. A. and Muller, R. A. (March 2005). "Cycles in fossil diversity" (PDF). Nature
434 (7030): 208–210. doi:10.1038/nature03339. PMID 15758998. http://muller.lbl.gov/papers/Rohde-
Muller-Nature.pdf. Retrieved 2008-09-22.
189.^ Field, C. B., Behrenfeld, M. J., Randerson, J. T. and Falkowski, P. (July 1998). "Primary
Production of the Biosphere: Integrating Terrestrial and Oceanic Components". Science 281 (5374):
 237–240. doi:10.1126/science.281.5374.237. PMID 9657713.
http://www.sciencemag.org/cgi/content/full/sci;281/5374/237. Retrieved 2008-09-13.
190.^ Grant, B. S., and Wiseman, L. L. (2002). "Recent History of Melanism in American
Peppered Moths". Journal of Heredity 93 (2): 86–90. doi:10.1093/jhered/93.2.86. ISSN 1465-7333.
PMID 12140267. http://jhered.oxfordjournals.org/cgi/content/abstract/93/2/86. Retrieved 2008-09-11.
191.^ Levin, B. R., Perrot, V. and Walker, N. (March 1, 2000). "Compensatory Mutations,
Antibiotic Resistance and the Population Genetics of Adaptive Evolution in Bacteria". Genetics 154
(3): 985–997. PMID 10757748. PMC 1460977.
http://www.genetics.org/cgi/content/abstract/154/3/985. Retrieved 2008-09-11.
192.^ Hawks, J., Wang, E. T., Cochran, G. M., Harpending, H. C. and Moyzis, R. K. (December
2007). "Recent acceleration of human adaptive evolution". Proceedings of the National Academy of
Sciences 104 (52): 20753–20758. doi:10.1073/pnas.0707650104. PMID 18087044. PMC 2410101.
http://www.pnas.org/content/104/52/20753.full. Retrieved 2008-09-11.

[edit] Further reading

• Cowen, R. (2004). History of Life (4th ed.). Blackwell Publishing Limited. ISBN 978-
1405117562.
• The Ancestor's Tale, A Pilgrimage to the Dawn of Life . Boston: Houghton Mifflin Company.
2004. ISBN 0-618-00583-8.
• Richard Dawkins. (1990). The Selfish Gene. Oxford University Press. ISBN 0192860925.
• Smith, John Maynard; Eörs Szathmáry (1997). The Major Transitions in Evolution.
Oxfordshire: Oxford University Press. ISBN 0-198-50294-X.
[edit] External links
General information
• General information on evolution- Fossil Museum nav.
• Understanding Evolution from University of California, Berkeley
• National Academies Evolution Resources
• Evolution poster- PDF format "tree of life"
• Everything you wanted to know about evolution by New Scientist
• Howstuffworks.com — How Evolution Works
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and
Theories
History of evolutionary thought
• The Complete Work of Charles Darwin Online
• Understanding Evolution: History, Theory, Evidence, and Implications
Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_life"

Categories: History by topic | Paleontology | Evolutionary biology

Hidden categories: Good articles
 W000

Histoire évolutive du vivant

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Cet article est une ébauche concernant l'origine et évolution de la vie.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
En biologie, l'évolution est le processus par lequel des populations d'organismes vivants acquièrent
et transmettent des traits biologiques nouveaux de génération en génération. Sur une grande échelle de
temps, la répétition de ce processus explique l'origine de nouvelles espèces et la vaste diversité du monde
vivant. Les espèces biologiques contemporaines sont reliées entre elles par une descendance commune et
sont le produit de l'évolution et de la spécialisation sur plusieurs milliards d'années.

Sommaire
[masquer]
• 1 Chronologie simple de l'évolution
• 2 L'origine de la vie
• 3 L'archéen et l'évolution des premières cellules
• 4 Le protérozoïque et l'apparition de pluricellularité
• 5 L'explosion cambrienne
• 6 Le paléozoïque et la colonisation du milieu terrestre
• 7 Le mésosoïque
• 8 Le cénozoïque : histoire évolutive récente
• 9 L'évolution aujourd'hui
• 10 Dans l'enseignement,
• 11 Notes et références
• 12 Voir aussi
• 12.1 Articles connexes
Chronologie simple de l'évolution [modifier]
• - 4 600 millions d'années : formation de la planète Terre,
• - 4 000 millions d'années : apparition des cellules procaryotes,
• - 3 000 millions d'années : apparition de la photosynthèse,
• - 2 100 millions d'années : apparition d'une vie multicellulaire,
• - 2 000 millions d'années : apparition des cellules eucaryotes,
• - 600 millions d'années : apparition d'animaux simples,
• - 570 millions d'années : apparition des arthropodes,
• - 550 millions d'années : apparition d'animaux complexes
• - 500 millions d'années : apparition des poissons et des proto-amphibiens,
• - 475 millions d'années : apparition des plantes terrestres,
• - 400 millions d'années : apparition des insectes et des graines,
• - 360 millions d'années : apparition des amphibiens,
• - 300 millions d'années : apparition des reptiles,
• - 200 millions d'années : apparition des mammifères,
• - 150 millions d'années : apparition des oiseaux,
• - 100 millions d'années : apparition des fleurs
• - 65 millions d'années : disparition des dinosaures non-aviaires (voir article Extinction du
Crétacé).
 L'origine de la vie [modifier]
Article détaillé : Origine de la vie.

L'archéen et l'évolution des premières cellules [modifier]

Il y a 3,5 milliards d'années les cyanobactéries apparaissent et constituent les stromatolithes
actuelles. Ces procaryotes (unicellulaires) immobiles (sans flagelle) longtemps appelés algues bleue-vertes
sont les premiers organismes à posséder les 2 photosystèmes leur permettant de pratiquer la
photosynthèse. Elles participent de ce fait à la réduction de la concentration en CO 2 atmosphérique et à
l'augmentation de la proportion en oxygène. Selon la théorie de l'endosymbiose plastidiale, elles seraient à
l'origine des plastes dans les végétaux supérieurs intégrés par symbiose dans leurs cellules.

Le protérozoïque et l'apparition de pluricellularité [modifier]

Article détaillé : Évolution de la pluricellularité.
L'existence d'ères glaciaires généralisées il y a de 850 à 630 millions d'années environ, une ère que
l'on appelle le cryogénien, aurait favorisé le regroupement des colonies bactériennes (ou des animalcules les
plus primitifs) autour « d'oasis » de vies dans des sources hydrothermales sous-marines ou affleurant en
surface. La planète étant plongée dans un manteau de glaces, ce qui rend les comportements
« individualistes » des micro-organismes défavorables.

L'explosion cambrienne [modifier]

Article détaillé : Explosion cambrienne.
Le paléozoïque et la colonisation du milieu terrestre [modifier]
• Histoire évolutive des poissons cartilagineux
Lignées passées au milieu terrestre :
• Histoire évolutive des végétaux
• Histoire évolutive des gastéropodes

Le mésosoïque [modifier]
• Histoire évolutive des mammifères
• Histoire évolutive des dinosaures
• Histoire évolutive des oiseaux

Le cénozoïque : histoire évolutive récente [modifier]

• Histoire évolutive des équidés
• Histoire évolutive des cétacés
• Histoire évolutive des hippopotamidés
• Histoire évolutive des siréniens
• Histoire évolutive des primates
• Histoire évolutive des homininés
 L'évolution aujourd'hui [modifier]
L'évolution, bien que lente et rarement observable à l'échelle humaine, peut parfois être décelée
chez les espèces à reproduction rapide. Ainsi, les moustiques sont parfois localement devenus résistants à
des insecticides. Il en va de même pour les bactéries responsables de pathologies humaines, souvent
multirésistantes aux antibiotiques.
Enfin, on peut constater le résultat d'une évolution divergente récente dans le cas de la colonie de crabes
découverte sous Rome[1]. Ainsi qu'un changement de régime alimentaire et l'apparition de sous-espèces
chez certains lézards, comme Podarcis sicula, avec modifications anatomiques internes et externes.

Dans l'enseignement, [modifier]

L'histoire évolutive est en Europe généralement, comme au collège de France [2] à Paris abordée
sous un double angle (Biologie historique et Évolutionnisme), en tentant d'à la fois enseigner ce qu'on sait de
l’histoire de la vie sur Terre et des mécanismes sous-jacents à cette histoire, et ce que l’on peut penser des
différentes interprétations humaines relatives à cette histoire et de ces mécanismes, interprétations qui ont
varié dans l'espace et dans le temps selon les contexte et les cultures ou religions.

Notes et références [modifier]

1. ↑ Sous les vestiges de Rome, une colonie de crabes prospère. [archive]
2. ↑ Leçon inaugurale d'Armand de Ricqlès au Collège de France [archive], Chaire de Biologie
historique et évolutionnisme, Lundi 6 mai 1996
 Voir aussi [modifier]
Articles connexes [modifier]
• Évolution
• Histoire de la Terre
• Biodiversité
• Biosphère

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle
 Adaptation · Convergence évolutive · Exaptation · Extinction des
Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux
 [Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_du_vivant ».

Catégorie : Histoire évolutive | [+]

 W000

Evidence of common descent

From Wikipedia, the free encyclopedia

Jump to: navigation, search

Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
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Biology portal · v • d • e
Scientists working in a variety of fields over many years have discovered a large body of evidence of
common descent of living things that demonstrates the occurrence of evolution and provides a wealth of
information on the natural processes by which the variety and diversity of life on Earth developed. This
evidence supports the modern evolutionary synthesis, which is the scientific theory that explains how and
why life changes over time. Evolutionary biologists document the fact of common descent; making
predictions, testing hypotheses, and developing theories that illustrate and describe its causes.
Comparison of the genetic sequence of organisms has revealed that organisms that are
phylogenetically close have a higher degree of sequence similarity than organisms that are phylogenetically
distant. Further evidence for common descent comes from genetic detritus such as pseudogenes, regions of
DNA that are orthologous to a gene in a related organism, but are no longer active and appear to be
undergoing a steady process of degeneration.
Fossils are important for estimating when various lineages developed in geologic time. As
fossilization is an uncommon occurrence, usually requiring hard body parts and death near a site where
sediments are being deposited, the fossil record only provides sparse and intermittent information about the
evolution of life. Evidence of organisms prior to the development of hard body parts such as shells, bones
and teeth is especially scarce, but exists in the form of ancient microfossils, as well as impressions of various
soft-bodied organisms. The comparative study of the anatomy of groups of animals reveals structural
features that are fundamentally similar or homologous, clearly demonstrating phylogenetic and ancestral
relationships with other organism, most especially when compared with fossils of ancient extinct organisms.
Vestigial structures and comparisons in embryonic development are largely a contributing factor in
anatomical resemblance in concordance with common descent. Since metabolic processes do not leave
fossils, research into the evolution of the basic cellular processes is done largely by comparison of existing
organisms’ physiology and biochemistry. Many lineages diverged at different stages of development, so it is
possible to determine when certain metabolic processes appeared by comparing the traits of the
descendants of a common ancestor. Universal biochemical organization and molecular variance patterns in
all organisms also show a direct correlation with common descent.
 Further evidence comes from the field of biogeography because evolution with common descent
provides the best and most thorough explanation for a variety of facts concerning the geographical
distribution of plants and animals across the world. This is especially obvious in the field of island
biogeography. Combined with the theory of plate tectonics common descent provides a way to combine facts
about the current distribution of species with evidence from the fossil record to provide a logically consistent
explanation of how the distribution of living organisms has changed over time.
The development and spread of antibiotic resistant bacteria, like the spread of pesticide resistant
forms of plants and insects provides evidence that evolution due to natural selection is an ongoing process in
the natural world. Alongside this, are observed instances of the separation of populations of species into sets
of new species (speciation). Speciation has been observed directly and indirectly in the lab and in nature.
Multiple forms of such have been described and documented as examples for individual modes of speciation.
Furthermore, evidence of common descent extends from direct laboratory and experimentation with the
artificial selection of organisms—historically and currently—and other controlled experiments involving many of
the topics in the article. This article explains the different types of evidence for evolution with common
descent along with many specialized examples of each.
Contents
[hide]
• 1 Evidence from comparative physiology and biochemistry
• 1.1 Genetics
• 1.1.1 Universal biochemical organisation
and molecular variance patterns
• 1.1.2 DNA sequencing
• 1.1.3 Proteins
• 1.1.4 Pseudogenes
• 1.1.5 Other mechanisms
• 1.2 Examples
• 1.2.1 Chromosome 2 in humans
• 1.2.2 Cytochrome c
• 1.2.3 Out of Africa hypothesis of human
evolution
• 2 Evidence from comparative anatomy
• 2.1 Homologous structures and divergent (adaptive)
evolution
• 2.2 Vestigial structures
• 2.3 Evolutionary developmental biology and
embryonic development
[edit] Evidence from comparative physiology and biochemistry
See also: Archaeogenetics, Common descent, Last universal ancestor, and Most recent common
ancestor, Timeline of evolution, Timeline of human evolution, and Universal Code (Biology)

[edit] Genetics

While on board HMS Beagle, Charles Darwin collected numerous specimens, many new to science,
which supported his later theory of evolution by natural selection.
Although it has only recently become available, one of the strongest evidences for common descent
comes from the study of gene sequences. Comparative sequence analysis examines the relationship
between the DNA sequences of different species[1], producing several lines of evidence that confirm
Darwin's original hypothesis of common descent. If the hypothesis of common descent is true, then species
that share a common ancestor will have inherited that ancestor's DNA sequence. Notably they will have
inherited mutations unique to that ancestor. More closely-related species will have a greater fraction of
identical sequence and will have shared substitutions when compared to more distantly-related species.
The simplest and most powerful evidence is provided by phylogenetic reconstruction. Such
reconstructions, especially when done using slowly-evolving protein sequences, are often quite robust and
can be used to reconstruct a great deal of the evolutionary history of modern organisms (and even in some
instances such as the recovered gene sequences of mammoths, Neanderthals or T. rex, the evolutionary
history of extinct organisms). These reconstructed phylogenies recapitulate the relationships established
through morphological and biochemical studies. The most detailed reconstructions have been performed on
the basis of the mitochondrial genomes shared by all eukaryotic organisms, which are short and easy to
sequence; the broadest reconstructions have been performed either using the sequences of a few very
ancient proteins or by using ribosomal RNA sequence.
This evidence does not support the rival hypothesis that genetic similarity of two species is the
product of common functional or structural requirements, and not common descent[ citation needed] (for
example, if there is one best way to produce a hoof, all hoofed creatures will share a genetic basis even if
they are not related). However, phylogenetic relationships also extend to a wide variety of nonfunctional
sequence elements, including repeats, transposons, pseudogenes, and mutations in protein-coding
sequences that do not result in changes in amino-acid sequence. While a minority of these elements might
later be found to harbor function, in aggregate they demonstrate that identity must be the product of common
descent rather than common function.
 [edit] Universal biochemical organisation and molecular variance patterns
All known extant organisms are based on the same fundamental biochemical organisation: genetic
information encoded as nucleic acid (DNA, or RNA for viruses), transcribed into RNA, then translated into
proteins (that is, polymers of amino acids) by highly conserved ribosomes. Perhaps most tellingly, the
Genetic Code (the "translation table" between DNA and amino acids) is the same for almost every organism,
meaning that a piece of DNA in a bacterium codes for the same amino acid as in a human cell. ATP is used
as energy currency by all extant life. A deeper understanding of developmental biology shows that common
morphology is, in fact, the product of shared genetic elements.[ citation needed] For example, although
camera-like eyes are believed to have evolved independently on many separate occasions,[ citation needed]
they share a common set of light-sensing proteins (opsins), suggesting a common point of origin for all
sighted creatures.[2] Another noteworthy example is the familiar vertebrate body plan, whose structure is
controlled by the homeobox (Hox) family of genes.

[edit] DNA sequencing

Comparison of the DNA sequences allows organisms to be grouped by sequence similarity, and the
resulting phylogenetic trees are typically congruent with traditional taxonomy, and are often used to
strengthen or correct taxonomic classifications. Sequence comparison is considered a measure robust
enough to be used to correct erroneous assumptions in the phylogenetic tree in instances where other
evidence is scarce. For example, neutral human DNA sequences are approximately 1.2% divergent (based
on substitutions) from those of their nearest genetic relative, the chimpanzee, 1.6% from gorillas, and 6.6%
from baboons.[3] Genetic sequence evidence thus allows inference and quantification of genetic relatedness
between humans and other apes.[4][5] The sequence of the 16S ribosomal RNA gene, a vital gene encoding
a part of the ribosome, was used to find the broad phylogenetic relationships between all extant life. The
analysis, originally done by Carl Woese, resulted in the three-domain system, arguing for two major splits in
the early evolution of life. The first split led to modern Bacteria and the subsequent split led to modern
Archaea and Eukaryotes.

[edit] Proteins
The proteomic evidence also supports the universal ancestry of life. Vital proteins, such as the
ribosome, DNA polymerase, and RNA polymerase, are found in everything from the most primitive bacteria
to the most complex mammals. The core part of the protein is conserved across all lineages of life, serving
similar functions. Higher organisms have evolved additional protein subunits, largely affecting the regulation
and protein-protein interaction of the core. Other overarching similarities between all lineages of extant
organisms, such as DNA, RNA, amino acids, and the lipid bilayer, give support to the theory of common
descent. The chirality of DNA, RNA, and amino acids is conserved across all known life. As there is no
functional advantage to right- or left-handed molecular chirality, the simplest hypothesis is that the choice
was made randomly by early organisms and passed on to all extant life through common descent. Further
evidence for reconstructing ancestral lineages comes from junk DNA such as pseudogenes, "dead" genes
which steadily accumulate mutations.[6]

[edit] Pseudogenes
Pseudogenes also known, as Junk DNA, or dead DNA is the extra DNA that is in our genome but
does not get transcribed into RNA to produce protein. This is an example of a vestige since reproducing
these genes uses energy and is a waste, yet 99% of our genome is these (1% working DNA)[7]. The way a
pseudogene is produced is a gene that is no longer used will accumulate mutations that make it non-
functional but since it is not used; when it disappears, it does not affect fitness.
 [edit] Other mechanisms
There is also a large body of molecular evidence for a number of different mechanisms for large
evolutionary changes, among them: genome and gene duplication, which facilitates rapid evolution by
providing substantial quantities of genetic material under weak or no selective constraints; horizontal gene
transfer, the process of transferring genetic material to another cell that is not an organism's offspring,
allowing for species to acquire beneficial genes from each other; and recombination, capable of reassorting
large numbers of different alleles and of establishing reproductive isolation. The Endosymbiotic theory
explains the origin of mitochondria and plastids (e.g. chloroplasts), which are organelles of eukaryotic cells,
as the incorporation of an ancient prokaryotic cell into ancient eukaryotic cell. Rather than evolving
eukaryotic organelles slowly, this theory offers a mechanism for a sudden evolutionary leap by incorporating
the genetic material and biochemical composition of a separate species. Evidence supporting this
mechanism has recently been found in the protist Hatena: as a predator it engulfs a green algae cell, which
subsequently behaves as an endosymbiont, nourishing Hatena, which in turn loses its feeding apparatus and
behaves as an autotroph.[8][9]
Since metabolic processes do not leave fossils, research into the evolution of the basic cellular
processes is done largely by comparison of existing organisms. Many lineages diverged when new metabolic
processes appeared, and it is theoretically possible to determine when certain metabolic processes
appeared by comparing the traits of the descendants of a common ancestor or by detecting their physical
manifestations. As an example, the appearance of oxygen in the earth's atmosphere is linked to the evolution
of photosynthesis.
[edit] Examples

[edit] Chromosome 2 in humans

Main article: Chromosome 2 (human)
Further information: Genes of the Chromosome 2 fusion site in chimpanzees

Fusion of ancestral chromosomes left distinctive remnants of telomeres, and a vestigial centromere
Clear evidence for the evolution of Homo sapiens from a common ancestor with chimpanzees is the
number of chromosomes in human as compared to all other members of Hominidae. All Hominidae with the
exception of humans have 24 pairs of chromosomes. Humans have only 23 pairs. Human chromosome 2 is
widely accepted to be a result of an end-to-end fusion of two ancestral chromosomes.[10][11]
The evidence for this includes:
 • The correspondence of chromosome 2 to two ape chromosomes. The closest human
relative, the common chimpanzee, has near-identical DNA sequences to human chromosome 2, but
they are found in two separate chromosomes. The same is true of the more distant gorilla and
orangutan.[12][13]
• The presence of a vestigial centromere. Normally a chromosome has just one centromere,
but in chromosome 2 there are remnants of a second centromere.[14]
• The presence of vestigial telomeres. These are normally found only at the ends of a
chromosome, but in chromosome 2 there are additional telomere sequences in the middle.[15]
Chromosome 2 thus presents very strong evidence in favour of the common descent of humans and
other apes. According to researcher J. W. IJdo, "We conclude that the locus cloned in cosmids c8.1 and
c29B is the relic of an ancient telomere-telomere fusion and marks the point at which two ancestral ape
chromosomes fused to give rise to human chromosome 2."[15]

[edit] Cytochrome c
Main article: Cytochrome c
A classic example of biochemical evidence for evolution is the variance of the ubiquitous (i.e. all
living organisms have it, because it performs very basic life functions) protein Cytochrome c in living cells.
The variance of cytochrome c of different organisms is measured in the number of differing amino acids,
each differing amino acid being a result of a base pair substitution, a mutation. If each differing amino acid is
assumed to be the result of one base pair substitution, it can be calculated how long ago the two species
diverged by multiplying the number of base pair substitutions by the estimated time it takes for a substituted
base pair of the cytochrome c gene to be successfully passed on. For example, if the average time it takes
for a base pair of the cytochrome c gene to mutate is N years, the number of amino acids making up the
cytochrome c protein in monkeys differ by one from that of humans, this leads to the conclusion that the two
species diverged N years ago.
The primary structure of cytochrome c consists of a chain of about 100 amino acids. Many higher
order organisms possess a chain of 104 amino acids.[16]
The cytochrome c molecule has been extensively studied for the glimpse it gives into evolutionary
biology. Both chicken and turkeys have identical sequence homology (amino acid for amino acid), as do pigs,
cows and sheep. Both humans and chimpanzees share the identical molecule, while rhesus monkeys share
all but one of the amino acids:[17] the 66th amino acid is isoleucine in the former and threonine in the latter.
[16]
What makes these homologous similarities particularly suggestive of common ancestry in the case of
cytochrome C, in addition to the fact that the phylogenies derived from them match other phylogenies very
well, is the high degree of functional redundancy of the cytochrome C molecule. The different existing
configurations of amino acids do not significantly affect the functionality of the protein, which indicates that
the base pair substitutions are not part of a directed design, but the result of random mutations that aren't
subject to selection.[18]

[edit] Out of Africa hypothesis of human evolution

Main article: Recent single-origin hypothesis
See also: Human mitochondrial DNA haplogroup and Human Y-chromosome DNA haplogroup
 Mathematical models of evolution, pioneered by the likes of Sewall Wright, Ronald Fisher and J. B.
S. Haldane and extended via diffusion theory by Motoo Kimura, allow predictions about the genetic structure
of evolving populations. Direct examination of the genetic structure of modern populations via DNA
sequencing has recently allowed verification of many of these predictions. For example, the Out of Africa
theory of human origins, which states that modern humans developed in Africa and a small sub-population
migrated out (undergoing a population bottleneck), implies that modern populations should show the
signatures of this migration pattern. Specifically, post-bottleneck populations (Europeans and Asians) should
show lower overall genetic diversity and a more uniform distribution of allele frequencies compared to the
African population. Both of these predictions are borne out by actual data from a number of studies.[19]

[edit] Evidence from comparative anatomy

Comparative study of the anatomy of groups of animals or plants reveals that certain structural
features are basically similar. For example, the basic structure of all flowers consists of sepals, petals,
stigma, style and ovary; yet the size, colour, number of parts and specific structure are different for each
individual species.

[edit] Homologous structures and divergent (adaptive) evolution

If widely separated groups of organisms are originated from a common ancestry, they are expected
to have certain basic features in common. The degree of resemblance between two organisms should
indicate how closely related they are in evolution:
• Groups with little in common are assumed to have diverged from a common ancestor much
earlier in geological history than groups which have a lot in common;
 • In deciding how closely related two animals are, a comparative anatomist looks for structures
that are fundamentally similar, even though they may serve different functions in the adult. Such
structures are described as homologous and suggest a common origin.
• In cases where the similar structures serve different functions in adults, it may be necessary
to trace their origin and embryonic development. A similar developmental origin suggests they are
the same structure, and thus likely to be derived from a common ancestor.
When a group of organisms share a homologous structure which is specialized to perform a variety
of functions in order to adapt different environmental conditions and modes of life are called adaptive
radiation. The gradual spreading of organisms with adaptive radiation is known as divergent evolution.

[edit] Vestigial structures

Main article: Vestigiality
See also: Human vestigiality
A strong and direct evidence for common descent comes from vestigial structures.[20] Rudimentary
body parts, those that are smaller and simpler in structure than corresponding parts in the ancestral species,
are called vestigial organs. They are usually degenerated or underdeveloped. The existence of vestigial
organs can be explained in terms of changes in the environment or modes of life of the species. Those
organs are thought to be functional in the ancestral species but are now either nonfunctional or repurposed.
Examples are the pelvic girdles of whales, haltere (hind wings) of flies and mosquitos, wings of flightless
birds such as ostriches, and the leaves of some xerophytes (e.g. cactus) and parasitic plants (e.g. dodder). It
must be noted, however, that vestigial structures may have had their original function replaced with another.
For example the halteres in dipterists help balance the insect while in flight and the wings of ostriches are
used in mating rituals.
“ The most reasonable conclusion to draw is that these creatures descended
from creatures in which these parts were functional, which in turn indicates that most (or
indeed all) creatures descended from common ancenstors. ”

—Natan Slifkin, The Challenge of Creation, page 262

[edit] Evolutionary developmental biology and embryonic development

This section requires expansion.

Main article: Evolutionary developmental biology

See also: Embryogenesis
Evolutionary developmental biology is the biological field that compares the developmental process
of different organisms to determine ancestral relationships between species. A large variety of organism’s
genomes contain a small fraction of genes that control the organisms development. Hox genes are an
example of these types of nearly universal genes in organisms pointing to an origin of common ancestry.
Embryological evidence stems from the development of organisms at the embryological level with the
comparison of different organisms embryos similarity. Remains of ancestral traits often appear and disappear
in different stages of the embryological development process. Examples such as hair growth and loss
(lanugo) during development[21]; the appearance of transitions from fish to amphibians to reptiles and then
to mammals in all mammal embryos; the appearance of gill-like structures (pharyngeal arch) in vertebrate
embryo development including. Note that in fish the arches become gills while in humans for example,
become the pharynx.

[edit] Atavisms
This section requires expansion.

Main article: Atavism

An atavism is an evolutionary throwback, such as traits reappearing which had disappeared
generations ago, some examples of this are chicken's teeth, Hind-legged snakes or whales, the extra toes of
ungulates that do not even reach the ground as well as Human tails.[22][23]
[edit] Examples
 Figure 5a: The principle of homology illustrated by the adaptive radiation of the forelimb of mammals.
All conform to the basic pentadactyl pattern but are modified for different usages. The third metacarpal is
shaded throughout; the shoulder is crossed-hatched.

Figure 5b: Illustration of the Eoraptor lunensis pelvis of the saurischian order and the Lesothosaurus
diagnosticus pelvis of the ornithischian order in the Dinosauria superorder. The parts of the pelvis show
modification over time. The cladogram is shown to illustrate the distance of divergence between the two
species.
 Figure 5c: Adaptation of insect mouthparts: a, antennae; c, compound eye; lb, labrium; lr, labrum;
md, mandibles; mx, maxillae.

[edit] Pentadactyl limb

Further information: Evolution of mammals
The pattern of limb bones called pentadactyl limb is an example of homologous structures (Fig. 5a).
It is found in all classes of tetrapods (i.e. from amphibians to mammals). It can even be traced back to the
fins of certain fossil fishes from which the first amphibians are thought to have evolved such as tiktaalik. The
limb has a single proximal bone (humerus), two distal bones (radius and ulna), a series of carpals (wrist
bones), followed by five series of metacarpals (palm bones) and phalanges (digits). Throughout the
tetrapods, the fundamental structures of pentadactyl limbs are the same, indicating that they originated from
a common ancestor. But in the course of evolution, these fundamental structures have been modified. They
have become superficially different and unrelated structures to serve different functions in adaptation to
different environments and modes of life. This phenomenon is clearly shown in the forelimbs of mammals.
For example:
• In the monkey, the forelimbs are much elongated to form a grasping hand for climbing and
swinging among trees.
• In the pig, the first digit is lost, and the second and fifth digits are reduced. The remaining two
digits are longer and stouter than the rest and bear a hoof for supporting the body.
• In the horse, the forelimbs are adapted for support and running by great elongation of the
third digit bearing a hoof.
• The mole has a pair of short, spade-like forelimbs for burrowing.
 • The anteater uses its enlarged third digit for tearing down ant hills and termite nests.
• In the whale, the forelimbs become flippers for steering and maintaining equilibrium during
swimming.
• In the bat, the forelimbs have turned into wings for flying by great elongation of four digits,
while the hook-like first digit remains free for hanging from trees.

[edit] Pelvic structure of dinosaurs

See also: Evolution of dinosaurs and Evolution of birds
Similar to the pentadactyl limb in mammals, the earliest dinosaurs split into two distinct orders—the
saurischia and ornithischia. They are classified as one or the other in accordance with what the fossils show.
In Figure 5b, it is clear that early saurischians resembled early ornithischians. The pattern of the pelvis in all
species of dinosaurs is an example of homologous structures. Each order of dinosaur has slightly differing
pelvis bones providing evidence of common descent. Additionally, modern birds show a similarity to ancient
saurischian pelvic structures therefore indicating the evolution of birds from dinosaurs.

[edit] Insect mouthparts

The basic structures are the same, including a labrum (upper lip), a pair of mandibles, a hypopharynx
(floor of mouth), a pair of maxillae, and a labium. These structures are enlarged and modified; others are
reduced and lost. The modifications enable the insects to exploit a variety of food materials (Fig. 5c):
(A) Primitive state — biting and chewing: e.g. grasshopper. Strong mandibles and maxillae for
manipulating food. (B) Ticking and biting: e.g. honey bee. Labium long to lap up nectar; mandibles chew
pollen and mould wax. (C) Sucking: e.g. butterfly. Labrum reduced; mandibles lost; maxillae long forming
sucking tube. (D) Piercing and sucking, e.g.. female mosquito. Labrum and maxillae form tube; mandibles
form piercing stylets; labrum grooved to hold other parts.

[edit] Other arthropod appendages

Insect mouthparts and antennae are considered homologues of insect legs. Parallel developments
are seen in some arachnids: The anterior pair of legs may be modified as analogues of antennae, particularly
in whip scorpions, which walk on six legs. These developments provide support for the theory that complex
modifications often arise by duplication of components, with the duplicates modified in different directions.
 [edit] Recurrent laryngeal nerve in giraffes

The path of the recurrent laryngeal nerve in giraffes. This is an excellent illustration of how inefficient
the evolution can be. The laryngeal nerve is compensated for by subsequent tinkering from natural selection.
The recurrent laryngeal nerve is a fourth branch of the vagus nerve, which is a cranial nerve. In
mammals its path is extraordinary: as a part of the vagus nerve, it comes from the brain, passes through the
neck down to heart, rounds the dorsal aorta and returns up to the larynx, again through the neck.
 This path looks not too optimal even for human, but for giraffes it becomes quite suboptimal. Due to
the lengths of their necks, the recurrent laryngeal nerve may be up to 4m long (13 ft), despite its optimal route
being a distance of just several inches.
Its route is explained by the evolution of mammals from fishes, which had no neck. Therefore the
nerve's route was quite reasonable: it was a relatively short nerve that innervated one gill slit, and passed
near the gill arch. Since then gills have evolved and the gill arch has became a dorsal aorta.[24][25]
[edit] Route of the vas deferens

Route of the vas deferens from the testis to the penis

 Similar to the laryngeal nerve in giraffes, the vas deferens is part of the male anatomy of many
vertebrates; it transports sperm from the epididymis in anticipation of ejaculation. In humans, the vas
deferens routes up from the testicle, looping over the ureter, and back down to the urethra and penis. It has
been suggested that this is due to the descent of the testicles during the course of human evolution—likely
associated with temperature. As the testicles descended, the vas deferens lengthened to accommodate the
accidental “hook” over the ureter.[25][26]
[edit] Evidence from paleontology

An insect trapped in amber.

When organisms die, they often decompose rapidly or are consumed by scavengers, leaving no
permanent evidences of their existence. However, occasionally, some organisms are preserved. The
remains or traces of organisms from a past geologic age embedded in rocks by natural processes are called
fossils. They are extremely important for understanding the evolutionary history of life on Earth, as they
provide direct evidence of evolution and detailed information on the ancestry of organisms. Paleontology is
the study of past life based on fossil records and their relations to different geologic time periods.
For fossilization to take place, the traces and remains of organisms must be quickly buried so that
weathering and decomposition do not occur. Skeletal structures or other hard parts of the organisms are the
most commonly occurring form of fossilized remains (Paul, 1998), (Behrensmeyer, 1980) and (Martin, 1999).
There are also some trace "fossils" showing moulds, cast or imprints of some previous organisms.
As an animal dies, the organic materials gradually decay, such that the bones become porous. If the
animal is subsequently buried in mud, mineral salts will infiltrate into the bones and gradually fill up the pores.
The bones will harden into stones and be preserved as fossils. This process is known as petrification. If dead
animals are covered by wind-blown sand, and if the sand is subsequently turned into mud by heavy rain or
floods, the same process of mineral infiltration may occur. Apart from petrification, the dead bodies of
organisms may be well preserved in ice, in hardened resin of coniferous trees (amber), in tar, or in anaerobic,
acidic peat. Fossilization can sometimes be a trace, an impression of a form. Examples include leaves and
footprints, the fossils of which are made in layers that then harden.
[edit] Fossil record

Fossil trilobite. Trilobites were hard-shelled arthropods, related to living horseshoe crabs and spiders,
that first appeared in significant numbers around 540 mya, dying out 250 mya.
It is possible to find out how a particular group of organisms evolved by arranging its fossil records in
a chronological sequence. Such a sequence can be determined because fossils are mainly found in
sedimentary rock. Sedimentary rock is formed by layers of silt or mud on top of each other; thus, the resulting
rock contains a series of horizontal layers, or strata. Each layer contains fossils which are typical for a
specific time period during which they were made. The lowest strata contain the oldest rock and the earliest
fossils, while the highest strata contain the youngest rock and more recent fossils.
A succession of animals and plants can also be seen from fossil records. By studying the number
and complexity of different fossils at different stratigraphic levels, it has been shown that older fossil-bearing
rocks contain fewer types of fossilized organisms, and they all have a simpler structure, whereas younger
rocks contain a greater variety of fossils, often with increasingly complex structures.[ citation needed]
In the past, geologists could only roughly estimate the ages of various strata and the fossils found.
They did so, for instance, by estimating the time for the formation of sedimentary rock layer by layer. Today,
by measuring the proportions of radioactive and stable elements in a given rock, the ages of fossils can be
more precisely dated by scientists. This technique is known as radiometric dating.
Throughout the fossil record, many species that appear at an early stratigraphic level disappear at a
later level. This is interpreted in evolutionary terms as indicating the times at which species originated and
became extinct. Geographical regions and climatic conditions have varied throughout the Earth's history.
Since organisms are adapted to particular environments, the constantly changing conditions favoured
species which adapted to new environments through the mechanism of natural selection.

[edit] Extent of the fossil record

See also: Transitional fossil and List of transitional fossils
 Charles Darwin collected fossils in South America, and found fragments of armor which he thought
were like giant versions of the scales on the modern armadillos living nearby. The anatomist Richard Owen
showed him that the fragments were from gigantic extinct glyptodons, related to the armadillos. This was one
of the patterns of distribution that helped Darwin to develop his theory.[27]
 Cynognathus, a Eucynodont, one of a grouping of Therapsids ("mammal-like reptiles") that is
ancestral to all modern mammals.
Despite the relative rarity of suitable conditions for fossilization, approximately 250,000 fossil species
are known.[28] The number of individual fossils this represents varies greatly from species to species, but
many millions of fossils have been recovered: for instance, more than three million fossils from the last Ice
Age have been recovered from the La Brea Tar Pits in Los Angeles.[29] Many more fossils are still in the
ground, in various geological formations known to contain a high fossil density, allowing estimates of the total
fossil content of the formation to be made. An example of this occurs in South Africa's Beaufort Formation
(part of the Karoo Supergroup, which covers most of South Africa), which is rich in vertebrate fossils,
including therapsids (reptile/mammal transitional forms).[30] It has been estimated that this formation
contains 800 billion vertebrate fossils.[31]

[edit] Limitations
The fossil record is an important source for scientists when tracing the evolutionary history of
organisms. However, because of limitations inherent in the record, there are not fine scales of intermediate
forms between related groups of species. This lack of continuous fossils in the record is a major limitation in
tracing the descent of biological groups. Furthermore, there are also much larger gaps between major
evolutionary lineages.[citation needed] When transitional fossils are found that show intermediate forms in
what had previously been a gap in knowledge, they are often popularly referred to as "missing links".
There is a gap of about 100 million years between the beginning of the Cambrian period and the end
of the Ordovician period. The early Cambrian period was the period from which numerous fossils of sponges,
cnidarians (e.g., jellyfish), echinoderms (e.g., eocrinoids), molluscs (e.g., snails) and arthropods (e.g.,
trilobites) are found. The first animal that possessed the typical features of vertebrates, the Arandaspis, was
dated to have existed in the later Ordovician period. Thus few, if any, fossils of an intermediate type between
invertebrates and vertebrates have been found, although likely candidates include the Burgess Shale animal,
Pikaia gracilens, and its Maotianshan shales relatives, Myllokunmingia, Yunnanozoon, Haikouella lanceolata,
and Haikouichthys.[citation needed]
Some of the reasons for the incompleteness of fossil records are:[ citation needed]
• In general, the probability that an organism becomes fossilized is very low;
• Some species or groups are less likely to become fossils because they are soft-bodied;
• Some species or groups are less likely to become fossils because they live (and die) in
conditions that are not favourable for fossilization;
• Many fossils have been destroyed through erosion and tectonic movements;
• Most fossils are fragmentary;
• Some evolutionary change occurs in populations at the limits of a species' ecological range,
and as these populations are likely to be small, the probability of fossilization is lower (see
punctuated equilibrium);
• Similarly, when environmental conditions change, the population of a species is likely to be
greatly reduced, such that any evolutionary change induced by these new conditions is less likely to
be fossilized;
• Most fossils convey information about external form, but little about how the organism
functioned;
• Using present-day biodiversity as a guide, this suggests that the fossils unearthed represent
only a small fraction of the large number of species of organisms that lived in the past.
[edit] Examples
This section requires expansion.

[edit] Evolution of the horse

Main article: Evolution of the horse
 Evolution of the horse showing reconstruction of the fossil species obtained from successive rock
strata. The foot diagrams are all front views of the left forefoot. The third metacarpal is shaded throughout.
The teeth are shown in longitudinal section.
Due to an almost-complete fossil record found in North American sedimentary deposits from the
early Eocene to the present, the horse provides one of the best examples of evolutionary history (phylogeny).
This evolutionary sequence starts with a small animal called Hyracotherium (commonly referred to as
Eohippus) which lived in North America about 54 million years ago, then spread across to Europe and Asia.
Fossil remains of Hyracotherium show it to have differed from the modern horse in three important respects:
it was a small animal (the size of a fox), lightly built and adapted for running; the limbs were short and
slender, and the feet elongated so that the digits were almost vertical, with four digits in the forelimbs and
three digits in the hindlimbs; and the incisors were small, the molars having low crowns with rounded cusps
covered in enamel.
The probable course of development of horses from Hyracotherium to Equus (the modern horse)
involved at least 12 genera and several hundred species. The major trends seen in the development of the
horse to changing environmental conditions may be summarized as follows:
• Increase in size (from 0.4 m to 1.5 m — from 15in to 60in);
• Lengthening of limbs and feet;
• Reduction of lateral digits;
• Increase in length and thickness of the third digit;
• Increase in width of incisors;
• Replacement of premolars by molars; and
 • Increases in tooth length, crown height of molars.
Fossilized plants found in different strata show that the marshy, wooded country in which
Hyracotherium lived became gradually drier. Survival now depended on the head being in an elevated
position for gaining a good view of the surrounding countryside, and on a high turn of speed for escape from
predators, hence the increase in size and the replacement of the splayed-out foot by the hoofed foot. The
drier, harder ground would make the original splayed-out foot unnecessary for support. The changes in the
teeth can be explained by assuming that the diet changed from soft vegetation to grass. A dominant genus
from each geological period has been selected to show the slow alteration of the horse lineage from its
ancestral to its modern form.

[edit] Evidence from geographical distribution

Data about the presence or absence of species on various continents and islands (biogeography)
can provide evidence of common descent and shed light on patterns of speciation.

[edit] Continental distribution

All organisms are adapted to their environment to a greater or lesser extent. If the abiotic and biotic
factors within a habitat are capable of supporting a particular species in one geographic area, then one might
assume that the same species would be found in a similar habitat in a similar geographic area, e.g. in Africa
and South America. This is not the case. Plant and animal species are discontinuously distributed throughout
the world:
• Africa has Old World monkeys, apes, elephants, leopards, giraffes, and hornbills.
• South America has New World monkeys, cougars, jaguars, sloths, llamas, and toucans.
 • Deserts in North and South America have native cacti, but deserts in Africa, Asia, and
Australia have succulent native euphorbs that resemble cacti but are very different, even though in
some cases cacti have done very well (for example in Australian deserts) when introduced by
humans.[32]
Even greater differences can be found if Australia is taken into consideration, though it occupies the
same latitude as much of South America and Africa. Marsupials like kangaroos, bandicoots, and quolls make
up about half of Australia's indigenous mammal species.[33] By contrast, marsupials are today totally absent
from Africa and form a smaller portion of the mammalian fauna of South America, where opossums, shrew
opossums, and the monito del monte occur. The echidnas and platypus, the only living representatives of
primitive egg-laying mammals (monotremes), can be found only in Australia and are totally absent in the rest
of the world. On the other hand, Australia is missing many groups of placental mammals that are common on
other continents (carnivorans, artiodactyls, shrews, squirrels, lagomorphs), although it does have indigenous
bats and murine rodents; many other placentals, such as rabbits and foxes, have been introduced there by
humans.
Other animal distribution examples include bears, located on all continents excluding Africa and
Australia, and the polar bear only located nearest to the North Pole. Penguins are located only around the
South Pole despite similar weather conditions at the North Pole. Families of sirenians are distributed
exclusively around the earth’s waters, where manatees are located in western Africa waters, northern South
American waters, and West Indian waters only while the related family, the Dugongs, are located only in
Oceanic waters north of Australia, and the coasts surrounding the Indian Ocean.
The same kinds of fossils are found from areas known to be adjacent to one another in the past but
which, through the process of continental drift, are now in widely divergent geographic locations. For
example, fossils of the same types of ancient amphibians, arthropods and ferns are found in South America,
Africa, India, Australia and Antarctica, which can be dated to the Paleozoic Era, at which time these regions
were united as a single landmass called Gondwana.[34] Sometimes the descendants of these organisms can
be identified and show unmistakable similarity to each other, even though they now inhabit very different
regions and climates.

[edit] Island biogeography

Four of the 13 finch species found on the Galápagos Archipelago, are thought to have evolved by an
adaptive radiation that diversified their beak shapes to adapt them to different food sources.
 [edit] Types of species found on islands
Evidence from island biogeography has played an important and historic role in the development of
evolutionary biology. For purposes of biogeography, islands are divided into two classes. Continental islands
are islands like Great Britain, and Japan that have at one time or another been part of a continent. Oceanic
islands, like the Hawaiian islands, the Galapagos islands and St. Helena, on the other hand are islands that
have formed in the ocean and never been part of any continent. Oceanic islands have distributions of native
plants and animals that are unbalanced in ways that make them distinct from the biotas found on continents
or continental islands. Oceanic islands do not have native terrestrial mammals (they do sometimes have bats
and seals), amphibians, or fresh water fish. In some cases they have terrestrial reptiles (such as the iguanas
and giant tortoises of the Galapagos islands) but often (for example Hawaii) they do not. This despite the fact
that when species such as rats, goats, pigs, cats, mice, and cane toads, are introduced to such islands by
humans they often thrive. Starting with Charles Darwin, many scientists have conducted experiments and
made observations that have shown that the types of animals and plants found, and not found, on such
islands are consistent with the theory that these islands were colonized accidentally by plants and animals
that were able to reach them. Such accidental colonization could occur by air, such as plant seeds carried by
migratory birds, or bats and insects being blown out over the sea by the wind, or by floating from a continent
or other island by sea, as for example by some kinds of plant seeds like coconuts that can survive immersion
in salt water, and reptiles that can survive for extended periods on rafts of vegetation carried to sea by
storms.[35]

[edit] Endemism
Many of the species found on remote islands are endemic to a particular island or group of islands,
meaning they are found nowhere else on earth. Examples of species endemic to islands include many
flightless birds of New Zealand, lemurs of Madagascar, the Komodo dragon of Komodo [36], the Dragon’s
blood tree of Socotra [37], Tuatara of New Zealand[38][39], and others. However many such endemic
species are related to species found on other nearby islands or continents; the relationship of the animals
found on the Galapagos Islands to those found in South America is a well-known example.[35] All of these
facts, the types of plants and animals found on oceanic islands, the large number of endemic species found
on oceanic islands, and the relationship of such species to those living on the nearest continents, are most
easily explained if the islands were colonized by species from nearby continents that evolved into the
endemic species now found there.[35]
Other types of endemism do not have to include, in the strict sense, islands. Islands can mean
isolated lakes or remote and isolated areas. Examples of these would include the highlands of Ethiopia, Lake
Baikal, Fynbos of South Africa, forests of New Caledonia, and others. Examples of endemic organisms living
in isolated areas include the Kagu of New Caledonia[40], cloud rats of the Luzon tropical pine forests of the
Philippines[41][42], the Baikal Seal[43] and the omul of Lake Baikal.

[edit] Adaptive radiations

Oceanic islands are frequently inhabited by clusters of closely related species that fill a variety of
ecological niches, often niches that are filled by very different species on continents. Such clusters, like the
Finches of the Galapagos, Hawaiian honeycreepers, members of the sunflower family on the Juan
Fernandez Archipelago and wood weevils on St. Helena are called adaptive radiations because they are best
explained by a single species colonizing an island (or group of islands) and then diversifying to fill available
ecological niches. Such radiations can be spectacular; 800 species of the fruit fly family Drosophila, nearly
half the world's total, are endemic to the Hawaiian islands. Another illustrative example from Hawaii is the
Silversword alliance, which is a group of thirty species found only on those islands. Members range from the
Silverswords that flower spectacularly on high volcanic slopes to trees, shrubs, vines and mats that occur at
various elevations from mountain top to sea level, and in Hawaiian habitats that vary from deserts to
rainforests. Their closest relatives outside Hawaii, based on molecular studies, are tarweeds found on the
west coast of North America. Interestingly, these tarweeds have sticky seeds that facilitate distribution by
migrant birds. Continental islands have less distinct biota, but those that have been long separated from any
continent also have endemic species and adaptive radiations, such as the 75 lemur species of Madagascar,
and the eleven extinct moa species of New Zealand.[35][44]

[edit] Ring Species

Main article: Ring species
In biology, a ring species is a connected series of neighboring populations that can interbreed with
relatively closely related populations, but for which there exist at least two "end" populations in the series that
are too distantly related to interbreed. Often such non-breeding-though-genetically-connected populations
co-exist in the same region thus creating a "ring". Ring species provide important evidence of evolution in
that they illustrate what happens over time as populations genetically diverge, and are special because they
represent in living populations what normally happens over time between long deceased ancestor
populations and living populations.[45][46]
[edit] Examples

Figure 6a: A dymaxion map of the world showing the distribution of present species of camelid. The
solid black lines indicate migration routes and the blue represents current camel locations.
 Figure 6b: Current distribution of Glossopteris placed on a Permian map showing the connection of
the continents. (1, South America; 2, Africa; 3, Madagascar; 4, India; 5, Antarctica; and 6, Australia)
 Figure 6c: Present day distribution of marsupials. (Distribution shown in blue. Introduced areas
shown in green.)

[edit] Migration, isolation, and distribution of the Camel

The history of the camel provides an example of how fossil evidence can be used to reconstruct
migration and subsequent evolution. The fossil record indicates that the evolution of camelids started in
North America (see figure 6a), from which 6 million years ago they migrated across the Bering Strait into Asia
and then to Africa, and 3.5 million years ago through the Isthmus of Panama into South America. Once
isolated, they evolved along their own lines, giving rise to the Bactrian camel and Dromedary in Asia and
Africa and the llama and its relatives in South America. Camelids then went extinct in North America at the
end of the last ice age.[47]
 [edit] Distribution of Glossopteris
The combination of continental drift and evolution can sometimes be used to make predictions about
what will be found in the fossil record. Glossopteris is an extinct species of seed fern plants from the
Permian. Glossopteris appears in the fossil record around the beginning of the Permian on the ancient
continent of Gondwana. Continental drift explains the current biogeography of the tree. Present day
Glossopteris fossils are found in Permian strata in southeast South America, southeast Africa, all of
Madagascar, northern India, all of Australia, all of New Zealand, and scattered on the southern and northern
edges of Antarctica. During the Permian, these continents were connected as Gondwana (see figure 6b) in
agreement with magnetic striping, other fossil distributions, and glacial scratches pointing away from the
temperate climate of the South Pole during the Permian.[35]

[edit] Distribution of marsupials

The history of marsupials also provides an example of how the theories of evolution and continental
drift can be combined to make predictions about what will be found in the fossil record. The earliest marsupial
fossils are about 80 million years old and found in North America; by 40 million years ago fossils show that
they could be found throughout South America, but there is no evidence of them in Australia, where they now
predominate, until about 30 million years ago. The theory of evolution predicts that the Australian marsupials
must be descended from the older ones found in the Americas. The theory of continental drift says that
between 30 and 40 million years ago South America and Australia were still part of the Southern hemisphere
super continent of Gondwana and that they were connected by land that is now part of Antarctica. Therefore
combining the two theories scientists predicted that marsupials migrated from what is now South America
across what is now Antarctica to what is now Australia between 40 and 30 million years ago. This hypothesis
led paleontologists to Antarctica to look for marsupial fossils of the appropriate age. After years of searching
they found, starting in 1982, fossils on Seymour Island off the coast of the Antarctic Peninsula of more than a
dozen marsupial species that lived 35-40 million years ago.[32]

[edit] Evidence from observed natural selection

This section requires expansion.

Examples for the evidence for evolution often stems from direct observation of natural selection in
the field and the laboratory. Scientists have observed and documented a multitude of events where natural
selection is in action. The most well known examples are antibiotic resistance in the medical field along with
better-known laboratory experiments documenting evolution's occurrence. Natural selection is tantamount to
common descent in the fact that long-term occurrence and selection pressures can lead to the diversity of life
on earth as found today. All adaptations—documented and undocumented changes concerned—are caused
by natural selection and genetic drift. The examples below are only a small fraction of the actual experiments
and observations.

[edit] Examples of natural selection in the lab and in the field

[edit] Antibiotic and pesticide resistance

Main article: Antibiotic resistance
The development and spread of antibiotic resistant bacteria, like the spread of pesticide resistant
forms of plants and insects is evidence for evolution of species, and of change within species. Thus the
appearance of vancomycin resistant Staphylococcus aureus, and the danger it poses to hospital patients is a
direct result of evolution through natural selection. The rise of Shigella strains resistant to the synthetic
antibiotic class of sulfonamides also demonstrates the generation of new information as an evolutionary
process.[48] Similarly, the appearance of DDT resistance in various forms of Anopheles mosquitoes, and the
appearance of myxomatosis resistance in breeding rabbit populations in Australia, are all evidence of the
existence of evolution in situations of evolutionary selection pressure in species in which generations occur
rapidly.

[edit] E. coli long-term evolution experiment

Main article: E. coli long-term evolution experiment
See also: Experimental evolution
Experimental evolution uses controlled experiments to test hypotheses and theories of evolution. In
one early example, William Dallinger set up an experiment shortly before 1880, subjecting microbes to heat
with the aim of forcing adaptive changes. His experiment ran for around seven years, and his published
results were acclaimed, but he did not resume the experiment after the apparatus failed.[49]
The E. coli long-term evolution experiment begun in 1988 under the leadership of Richard Lenski is
still in progress, and has shown adaptations including the evolution of a strain of E. coli that was able to grow
on citric acid in the growth media.

[edit] Lactose intolerance in humans

Lactose intolerance is the inability to metabolize lactose, because of a lack of the required enzyme
lactase in the digestive system. The normal mammalian condition is for the young of a species to experience
reduced lactase production at the end of the weaning period (a species-specific length of time). In humans, in
non-dairy consuming societies, lactase production usually drops about 90% during the first four years of life,
although the exact drop over time varies widely.[50] However, certain human populations have a mutation on
chromosome 2 which eliminates the shutdown in lactase production, making it possible for members of these
populations to continue consumption of fresh milk and other dairy products throughout their lives without
difficulty. This appears to be an evolutionarily recent adaptation to dairy consumption, and has occurred
independently in both northern Europe and east Africa in populations with a historically pastoral lifestyle.[51]

[edit] Nylon-eating bacteria

Nylon-eating bacteria are a strain of Flavobacterium that is capable of digesting certain byproducts of
nylon 6 manufacture. There is scientific consensus that the capacity to synthesize nylonase most probably
developed as a single-step mutation that survived because it improved the fitness of the bacteria possessing
the mutation. This is seen as a good example of evolution through mutation and natural selection that has
been observed as it occurs.[52][53][54][55]

[edit] Peppered moth

Main article: Peppered moth evolution
One classic example of adaptation in response to selection pressure is the case of the peppered
moth. The color of the moth has gone from light to dark to light again over the course of a few hundred years
due to the appearance and later disappearance of pollution from the Industrial Revolution in England.
 [edit] Radiotrophic fungus
Recently evolved Radiotrophic fungi are fungi which appear to use the pigment melanin to convert
gamma radiation into chemical energy for growth[56][57] and were first discovered in 2007 as black molds
growing inside and around the Chernobyl Nuclear Power Plant.[56] Research at the Albert Einstein College
of Medicine showed that three melanin-containing fungi, Cladosporium sphaerospermum, Wangiella
dermatitidis, and Cryptococcus neoformans, increased in biomass and accumulated acetate faster in an
environment in which the radiation level was 500 times higher than in the normal environment.

[edit] Urban wildlife

Urban wildlife is wildlife that is able to live or thrive in urban environments. These types of
environments can exert selection pressures on organism, often leading to new adaptations. For example, the
weed Crepis sancta, found in France, has two types of seed, heavy and fluffy. The heavy ones land nearby to
the parent plant, whereas the fluffy seeds float further away on the wind. In urban environments, seeds that
float far will often land on infertile concrete. Within about 5-12 generations, the weed has been found to
evolve to produce significantly more heavy seeds than its rural relatives do.[58][59]

[edit] Evidence from observed speciation

Speciation is the evolutionary process by which new biological species arise. Speciation can occur
from a variety of different causes and are classified in various forms (e.g. allopatric, sympatric,
polyploidization, etc). Scientists have observed numerous examples of speciation in the laboratory and in
nature, however, evolution has produced far more species than an observer would consider necessary. For
example, there are well over 350,000 described species of beetles.[60] Great examples of observed
speciation come from the observations of island biogeography and the process of adaptive radiation, both
explained in an earlier section. The examples shown below provide strong evidence for common descent and
are only a small fraction of the instances observed.

[edit] Examples

[edit] Blackcap
The Sylvia atricapillab, commonly referred to as Blackcaps, lives in Germany and flies southwest to
Spain while a smaller group flies northwest to Great Britain during the winter. The smaller blackcap
population only recently rerouted to Spain. Gregor Rolshausen from the University of Freiburg found that the
genetic separation of the two populations is already in progress. The differences found have arisen in about
30 generations. With DNA sequencing, the individuals can be assigned to a correct group with an 85%
accuracy. Stuart Bearhop from the University of Exeter reported that birds wintering in England tend to mate
only among themselves, and not usually with those wintering in the Mediterranean. (Bearhop et al. 2005) It is
still inference to say that the populations will become two different species, but experts deduce that it is
expected due to the continued genetic and geographic separation.[61]
 [edit] Drosophila melanogaster

A common fruit fly (Drosophila melanogaster).

William R. Rice and George W. Salt found experimental evidence of sympatric speciation in the
common fruit fly. They collected a population of Drosophila melanogaster from Davis, California and placed
the pupae into a habitat maze. Newborn flies had to investigate the maze to find food. The flies had three
choices to take in finding food. Light and dark (phototaxis), up and down (geotaxis), and the scent of
acetaldehyde and the scent of ethanol (chemotaxis) were the three options. This eventually divided the flies
into 42 spatio-temporal habitats.
They then cultured two strains that chose opposite habitats. One of the strains emerged early,
immediately flying upward in the dark attracted to the acetaldehyde. The other strain emerged late and
immediately flew downward, attracted to light and ethanol. Pupae from the two strains were then placed
together in the maze and allowed to mate at the food site. They then were collected. A selective penalty was
imposed on the female flies that switched habitats. This entailed that none of their gametes would pass on to
the next generation. After 25 generations of this mating test, it showed reproductive isolation between the two
strains. They repeated the experiment again without creating the penalty against habitat switching and the
result was the same; reproductive isolation was produced.[62][63][64]

[edit] Hawthorn fly

One example of evolution at work is the case of the hawthorn fly, Rhagoletis pomonella, also known
as the apple maggot fly, which appears to be undergoing sympatric speciation.[65] Different populations of
hawthorn fly feed on different fruits. A distinct population emerged in North America in the 19th century some
time after apples, a non-native species, were introduced. This apple-feeding population normally feeds only
on apples and not on the historically preferred fruit of hawthorns. The current hawthorn feeding population
does not normally feed on apples. Some evidence, such as the fact that six out of thirteen allozyme loci are
different, that hawthorn flies mature later in the season and take longer to mature than apple flies; and that
there is little evidence of interbreeding (researchers have documented a 4-6% hybridization rate) suggests
that this is occurring.[66]

[edit] London Underground mosquito

The London Underground mosquito is a species of mosquito in the genus Culex found in the London
Underground. It is thought to have evolved from the overground species Culex pipiens recently.
This mosquito, although first discovered in the London Underground system, has been found in
underground systems around the world. It is suggested that it may have adapted to human-made
underground systems since the last century from local above-ground Culex pipiens,[67] although more recent
evidence suggests that it is a southern mosquito variety related to Culex pipiens that has adapted to the
warm underground spaces of northern cities.[68]
The evidence for this mosquito being a different species from Culex pipiens comes from research by
Kate Byne and Richard Nichols. The species have very different behaviours,[69] are extremely difficult to
mate,[67] and with different allele frequency, consistent with genetic drift during a founder event.[70] More
specifically, this mosquito, Culex pipiens molestus, breeds all-year round, is cold intolerant, and bites rats,
mice, and humans, in contrast to the above ground species Culex pipiens that is cold tolerant, hibernates in
the winter, and bites only birds. When the two varieties were cross-bred the eggs were infertile suggesting
reproductive isolation.[67][69]
The fundamental results still stands: the genetic data indicate that the molestus form in the London
Underground mosquito appeared to have a common ancestry, rather than the population at each station
being related to the nearest above-ground population (i.e. the pipiens form). Byrne and Nichols' working
hypothesis was that adaptation to the underground environment had occurred locally in London once only.
These widely separated populations are distinguished by very minor genetic differences, which
suggest that the molestus form developed recently: a single mtDNA difference shared among the
underground populations of ten Russian cities;[71] a single fixed microsatellite difference in populations
spanning Europe, Japan, Australia, the middle East and Atlantic islands.[68]

[edit] Madeira House Mouse

This section requires expansion.
 The Madeira House mouse is a new species of mouse descended from the house mouse ( Mus
Musculus) that went through a recent speciation event.[72]

[edit] Mollies
The Shortfin Molly—Poecilia Mexicana—is a small fish that lives in the Sulfur Caves of Mexico.
Michael Tobler from the Texas A&M University has studied the fish for years and found that two distinct
populations of mollies—the dark interior fish and the bright surface water fish—are becoming more genetically
divergent. (Tobler et al. 2009) The populations have no obvious barrier separating the two; however, it was
found that the mollies are hunted by a large water bug (Belostoma spp). Tobler collected the bug and both
types of mollies, placed them in large plastic bottles, and put them back in the cave. After a day, it was found
that, in the light, the cave-adapted fish endured the most damage, with four out of every five stab-wounds
from the water bugs sharp mouthparts. In the dark, the situation was the opposite. The mollies’ senses can
detect a predator’s threat in their own habitats, but not in the other ones. Moving from one habitat to the other
significantly increases the risk of dying. Tobler plans on further experiments, but it is believed that it is a good
example of the rise of a new species.[73]

[edit] Thale cress

Kirsten Bomblies et al. from the Max Planck Institute for Developmental Biology discovered that two
genes passed down by each parent of the thale cress plant, Arabidopsis thaliana. When the genes are
passed down, it ignites a reaction in the hybrid plant that turns its own immune system against it. In the
parents, the genes were not detrimental, but they evolved separately to react defectively when combined.
(Bomlies et al. 2007)
 To test this, Bomblies crossed 280 genetically different strains of Arabidopsis in 861 distinct ways
and found that 2 per cent of the resulting hybrids were necrotic. Along with allocating the same indicators, the
20 plants also shared a comparable collection of genetic activity in a group of 1,080 genes. In almost all of
the cases, Bomblies discovered that only two genes were required to cause the autoimmune response.
Bomblies looked at one hybrid in detail and found that one of the two genes belonged to the NB-LRR class, a
common group of disease resistance genes involved in recognizing new infections. When Bomblies removed
the problematic gene, the hybrids developed normally. (Bomlies et al. 2007)
Over successive generations, these incompatibilities could create divisions between different plant
strains, reducing their chances of successful mating and turning distinct strains into separate species.[74]

[edit] Interspecies fertility or hybridization

Understood from laboratory studies and observed instances of speciation in nature, finding species
that are able reproduce successfully or create hybrids between two different species infers that their
relationship is close. In conjunction with this, hybridization has been found to be a precursor to the creation of
new species by being a source of new genes for a species. The examples provided are only a small fraction
of the observed instances of speciation through hybridization. It is worth noting that plants are often subject
to the creation of a new species though hybridization.

[edit] Polar bear

See also: Polar bear#Taxonomy and evolution
A specific example of large-scale evolution is the polar bear (Ursus maritimus), which though clearly
related to the brown bear (Ursus arctos) by virtue of the fact that though separate species they can still
interbreed and produce fertile offspring[75] it has also obviously acquired significant physiological differences
from the brown bear. These differences allow the polar bear to comfortably survive in conditions that the
brown could not including the ability to swim sixty miles or more at a time in freezing waters, blend in—the
change in fur color—and to stay warm in the arctic environment. Additionally, the elongation of the neck
makes it easier to keep their heads above water while swimming and the oversized webbed feet that act as
paddles when swimming. The polar bear has also evolved small papillae and vacuole-like suction cups on
the soles to make them less likely to slip on the ice, feet covered with heavy matting to protect the bottoms
from intense cold and provide traction, smaller ears to reduce the loss of heat, eyelids that act like
sunglasses, to accommodate their all-meat diet, a large stomach capacity to enable opportunistic feeding,
and the ability to fast for up to nine months while recycling their urea.[76][77]

[edit] Raphanobrassica
Raphanobrassica includes all intergeneric hybrids between the genera Raphanus (radish) and
Brassica (cabbages, etc).
The Raphanobrassica is an allopolyploid cross between the radish (Raphanus sativus) and cabbage
(Brassica oleracea). Plants of this parentage are now known as radicole. Two other fertile forms of
Raphanobrassica are known. Raparadish, an allopolyploid hybrid between Raphanus sativus and Brassica
rapa is grown as a fodder crop. "Raphanofortii" is the allopolyploid hybrid between Brassica tournefortii and
Raphanus caudatus.
The Raphanobrassica is a fascinating plant, because—in spite of its hybrid nature—it is not sterile.
This has led some botanists to propose that the accidental hybridization of a flower by pollen of another
species in nature could be a mechanism of speciation common in higher plants.
 [edit] Salsify
Salsifies are one example where hybrid speciation has been observed. In the early 1900s, humans
introduced three species of goatsbeard into North America. These species, the western salsify ( T. dubius),
the meadow salsify (T. pratensis), and the oyster plant (T. porrifolius), are now common weeds in urban
wastelands. In the 1950s, botanists found two new species in the regions of Idaho and Washington, where
the three already known species overlapped. One new species, Tragopogon miscellus, is a tetraploid hybrid
of T. dubius and T. pratensis. The other species, Tragopogon mirus, is also an allopolyploid, but its ancestors
were T. dubius and T. porrifolius. These new species are usually referred to as "the Ownbey hybrids" after
the botanist who first described them. The T. mirus population grows mainly by reproduction of its own
members, but additional episodes of hybridization continue to add to the T. mirus population.[citation needed]

[edit] Welsh groundsel

Welsh groundsel is an allopolyploid, a plant which contains sets of chromosomes originating from
two different species. Its ancestor was Senecio × baxteri, an infertile hybrid which can arise spontaneously
when the closely-related groundsel (Senecio vulgaris) and Oxford ragwort (Senecio squalidus) grow
alongside each other. Sometime in the early 20th century, an accidental doubling of the number of
chromosomes in an S. × baxteri plant led to the formation of a new fertile species.[78][79]

[edit] York groundsel

The York groundsel (Senecio eboracensis) is a hybrid species of the self-incompatible Senecio
squalidus (also known as Oxford ragwort) and the self-compatible Senecio vulgaris (also known as Common
groundsel). Like S. vulgaris, S. eboracensis is self-compatible, however, it shows little or no natural crossing
with its parent species, and is therefore reproductively isolated, indicating that strong breed barriers exist
between this new hybrid and its parents.
It is thought to have resulted from a backcrossing of the F1 hybrid of its parents to S. vulgaris. S.
vulgaris is native to Britain, while S. squalidus was introduced from Sicily in the early 1700s; therefore, S.
eboracensis has speciated from those two species within the last 300 years.
Other hybrids descended from the same two parents are known. Some are infertile, such as S. x
baxteri. Other fertile hybrids are also known, including S. vulgaris var. hibernicus, now common in Britain,
and the allohexaploid S. cambrensis, which according to molecular evidence probably originated
independently at least three times in different locations. Morphological and genetic evidence support the
status of S. eboracensis as separate from other known hybrids.[80]

[edit] Evidence from artificial selection

 The Chihuahua mix and Great Dane illustrate the range of sizes among dog breeds.
Artificial selection demonstrates the diversity that can exist among organisms that share a relatively
recent common ancestor. In artificial selection, one species is bred selectively at each generation, allowing
only those organisms that exhibit desired characteristics to reproduce. These characteristics become
increasingly well-developed in successive generations. Artificial selection was successful long before science
discovered the genetic basis. Examples of artificial selection would be dog breeding, genetically modified
food, flower breeding, cultivation of foods such as wild cabbage[81], and others.

[edit] Evidence from computation and mathematical iteration

This section requires expansion.

"It has taken more than five decades, but the electronic computer is now powerful enough to simulate
evolution"[82] assisting bioinformatics in its attempt to solve biological problems.
Computer science allows the iteration of self changing complex systems to be studied, allowing a
mathematical understanding of the nature of the processes behind evolution; providing evidence for the
hidden causes of known evolutionary events. The evolution of specific cellular mechanisms like
spliceosomes that can turn the cell's genome into a vast workshop of billions of interchangeable parts that
can create tools that create tools that create tools that create us can be studied for the first time in an exact
way.
For example, Christoph Adami et al. make this point in Evolution of biological complexity:
 To make a case for or against a trend in the evolution of complexity in biological evolution,
complexity needs to be both rigorously defined and measurable. A recent information-theoretic (but
intuitively evident) definition identifies genomic complexity with the amount of information a sequence
stores about its environment. We investigate the evolution of genomic complexity in populations of
digital organisms and monitor in detail the evolutionary transitions that increase complexity. We show
that, because natural selection forces genomes to behave as a natural "Maxwell Demon," within a
fixed environment, genomic complexity is forced to increase.[83]
For example, David J. Earl and Michael W. Deem make this point in Evolvability is a selectable trait:
Not only has life evolved, but life has evolved to evolve. That is, correlations within protein
structure have evolved, and mechanisms to manipulate these correlations have evolved in tandem.
The rates at which the various events within the hierarchy of evolutionary moves occur are not random
or arbitrary but are selected by Darwinian evolution. Sensibly, rapid or extreme environmental change
leads to selection for greater evolvability. This selection is not forbidden by causality and is strongest
on the largest-scale moves within the mutational hierarchy. Many observations within evolutionary
biology, heretofore considered evolutionary happenstance or accidents, are explained by selection for
evolvability. For example, the vertebrate immune system shows that the variable environment of
antigens has provided selective pressure for the use of adaptable codons and low-fidelity polymerases
during somatic hypermutation. A similar driving force for biased codon usage as a result of
productively high mutation rates is observed in the hemagglutinin protein of influenza A.[84]
"Computer simulations of the evolution of linear sequences have demonstrated the importance of
recombination of blocks of sequence rather than point mutagenesis alone. Repeated cycles of point
mutagenesis, recombination, and selection should allow in vitro molecular evolution of complex sequences,
such as proteins."[85] Evolutionary molecular engineering, also called directed evolution or in vitro molecular
evolution involves the iterated cycle of mutation, multiplication with recombination, and selection of the fittest
of individual molecules (proteins, DNA, and RNA). Natural evolution can be relived showing us possible
paths from catalytic cycles based on proteins to based on RNA to based on DNA.[85][86][87][88]

[edit] Works
• Bearhop, Stuart; Fiedler, Wolfgang; Furness, Robert W.; Votier, Stephen C.; Waldron,
Susan; Newton, Jason; Bowen, Gabriel J.; Berthold, Peter & Farnsworth, Keith (2005): Assortative
mating as a mechanism for rapid evolution of a migratory divide. Science 310(5747): 502-504.
doi:10.1126/science.1115661 (HTML abstract) Supporting Online Material
• Bomblies, Lempe, Epple, Warthmann, Lanz, Dangl & Weigel. 2007. Autoimmune response
as a mechanism for a Dobhansky-Muller-Type incompatibility syndrome in plants. PLoS Biol 5: e236
doi:10.1371/journal.pbio.0050236
• Davis, Paul and Kenrick, Paul. 2004. Fossil Plants. Smithsonian Books (in association with
the Natural History Museum of London), Washington, D.C. ISBN 1-58834-156-9
• Karpechenko, G.D., Polyploid hybrids of Raphanus sativus X Brassica oleracea L., Bull.
Appl. Bot. 17:305-408 (1927).
• Terasawa, Y. Crossing between Brassico-raphanus and B. chinensis and Raphanus sativus.
Japanese Journal of Genetics. 8(4): 229–230 (1933).
• Tobler, Micheal (2009). Does a predatory insect contribute to the divergence between cave-
and surface-adapted fish populations? Biology Letters doi:10.1098/rsbl.2009.0272
[edit] See also
• Nothing in Biology Makes Sense Except in the Light of Evolution

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

[edit] References
1. ^ Mount DM. (2004). Bioinformatics: Sequence and Genome Analysis (2nd ed.). Cold Spring
Harbor Laboratory Press: Cold Spring Harbor, NY.. ISBN 0-87969-608-7.
2. ^ http://www.imls.uzh.ch/research/noll/publ/Dev_Cell_2003_5_773_785.pdf
3. ^ Two sources:
Chen FC, Li WH (February 2001). "Genomic divergences between humans and other hominoids and
the effective population size of the common ancestor of humans and chimpanzees". Am J Hum
Genet. 68 (2): 444–56. doi:10.1086/318206. PMID 11170892.
Cooper GM, Brudno M, Green ED, Batzoglou S, Sidow A (May 2003). "Quantitative estimates of
sequence divergence for comparative analyses of mammalian genomes". Genome Res. 13 (5): 813–
20. doi:10.1101/gr.1064503. PMID 12727901. PMC 430923.
http://www.genome.org/cgi/content/full/13/5/813.
4. ^ The picture labeled "Human Chromosome 2 and its analogs in the apes" in the article
Comparison of the Human and Great Ape Chromosomes as Evidence for Common Ancestry is
literally a picture of a link in humans that links two separate chromosomes in the nonhuman apes
creating a single chromosome in humans. It is considered a missing link, and the ape-human
connection is of particular interest. Also, while the term originally referred to fossil evidence, this too
is a trace from the past corresponding to some living beings which when alive were the physical
embodiment of this link.
5. ^ The New York Times report Still Evolving, Human Genes Tell New Story , based on A Map
of Recent Positive Selection in the Human Genome , states the International HapMap Project is
"providing the strongest evidence yet that humans are still evolving" and details some of that
evidence.
6. ^ Petrov DA, Hartl DL (2000). "Pseudogene evolution and natural selection for a compact
genome". J Hered. 91 (3): 221–7. doi:10.1093/jhered/91.3.221. PMID 10833048.
http://jhered.oxfordjournals.org/cgi/pmidlookup?view=long&pmid=10833048.
7. ^ http://www.genomesize.com/
8. ^ Okamoto N, Inouye I (2005). "A secondary symbiosis in progress". Science 310 (5746):
287. doi:10.1126/science.1116125. PMID 16224014.
9. ^ Okamoto N, Inouye I (October 2006). "Hatena arenicola gen. et sp. nov., a katablepharid
undergoing probable plastid acquisition". Protist 157 (4): 401–19. doi:10.1016/j.protis.2006.05.011.
PMID 16891155.
10.^ Human Chromosome 2 is a fusion of two ancestral chromosomes by Alec MacAndrew;
accessed 18 May 2006.
11.^ Evidence of Common Ancestry: Human Chromosome 2 (video) 2007
12.^ Yunis and Prakash; Prakash, O (1982). "The origin of man: a chromosomal pictorial
legacy". Science 215 (4539): 1525–1530. doi:10.1126/science.7063861. PMID 7063861.
13.^ Human and Ape Chromosomes; accessed 8 September 2007.
 14.^ Avarello et al.; Pedicini, A; Caiulo, A; Zuffardi, O; Fraccaro, M (1992). "Evidence for an
ancestral alphoid domain on the long arm of human chromosome 2". Human Genetics 89 (2): 247–9.
doi:10.1007/BF00217134. PMID 1587535.
15.^ a b IJdo et al.; Baldini, A; Ward, DC; Reeders, ST; Wells, RA (1991). "Origin of human
chromosome 2: an ancestral telomere-telomere fusion". Proceedings of the National Academy of
Sciences 88 (20): 9051–5. doi:10.1073/pnas.88.20.9051. PMID 1924367.
16.^ a b Amino acid sequences in cytochrome c proteins from different species, adapted from
Strahler, Arthur; Science and Earth History, 1997. page 348.
17.^ Lurquin PF, Stone L (2006). Genes, Culture, and Human Evolution: A Synthesis. Blackwell
Publishing, Incorporated. pp. 79. ISBN 1-4051-5089-0. http://books.google.com/?
id=zdeWdF_NQhEC&pg=PA79&lpg=PA79&dq=chimpanzee+rhesus+cytochrome+c.
18.^ 29+ Evidences for Macroevolution; Protein functional redundancy, Douglas Theobald,
Ph.D.
19.^ Pallen, Mark (2009). Rough Guide to Evolution. Rough Guides. pp. 200–206. ISBN 978-1-
85828-946-5.
20.^ Natan Slifkin (2006). The Challenge of Creation.... Zoo Torah. pp. 258–9. ISBN 1-933143-
15-0.
21.^ http://www.springerlink.com/content/mr73x80w2x7744jp/fulltext.pdf
22.^ http://www.edwardtbabinski.us/whales/atavisms.html
23.^ http://animals.howstuffworks.com/animal-facts/atavism.htm
24.^ Mark Ridley (2004). Evolution (3rd ed.). Blackwell Publishing. pp. 282. ISBN 1405103450.
http://books.google.com/?id=b-HGB9PqXCUC&lpg=RA1-PA281&pg=RA1-PA282#v=onepage&q=. .
25.^ a b Dawkins, Richard (2009). The Greatest Show on Earth: The Evidence for Evolution .
Bantam Press. pp. 364–365. ISBN 978-1-4165-9478-9.
 26.^ Williams, G.C. (1992). Natural selection: domains, levels, and challenges. Oxford Press.
ISBN 0-19-506932-3.
27.^ "Confessions of a Darwinist". Niles Eldredge.
http://www.vqronline.org/articles/2006/spring/eldredge-confessions-darwinist/. Retrieved 2010-06-
22.
28.^ Laboratory 11 - Fossil Preservation, by Pamela J. W. Gore, Georgia Perimeter College
29.^ [1]
30.^ [2]
31.^ Six Flood Arguments Creationists Can't Answer, by Robert J. Schadewald, 1982
32.^ a b Coyne, Jerry A. (2009). Why Evolution is True. Viking. pp. 91–99. ISBN 978-0-670-
02053-9.
33.^ Menkhorst, Peter; Knight, Frank (2001). A Field Guide to the Mammals of Australia. Oxford
Uniersity Press. p. 14. ISBN 0-19-550870-X.
34.^ http://biology.clc.uc.edu/courses/bio303/contdrift.htm
35.^ a b c d e Coyne, Jerry A. (2009). Why Evolution is True. Viking. pp. 99–110. ISBN 978-0-670-
02053-9.
36.^ Trooper Walsh; Murphy, James Jerome; Claudio Ciofi; Colomba De LA Panouse (2002).
Komodo Dragons: Biology and Conservation (Zoo and Aquarium Biology and Conservation Series) .
Washington, D.C.: Smithsonian Books. ISBN 1-58834-073-2.
37.^ "The Wonder Land of Socotra, Yemen". ALAN BURDICK.
http://travel.nytimes.com/2007/03/25/travel/tmagazine/03well.socotra.t.html. Retrieved 2010-07-08.
38.^ "Tuatara". New Zealand Ecology: Living Fossils. TerraNature Trust. 2004.
http://www.terranature.org/tuatara.htm. Retrieved 2006-11-10.
 39.^ "Facts about tuatara". Conservation: Native Species. Threatened Species Unit, Department
of Conservation, Government of New Zealand. http://www.doc.govt.nz/templates/page.aspx?
id=33163. Retrieved 2007-02-10.
40.^ "New Caledonia's most wanted".
http://www.birdlife.org/news/features/2006/05/new_caledonia.html. Retrieved 2010-07-08.
41.^ "Giant bushy-tailed cloud rat (Crateromys schadenbergi)". http://www.arkive.org/giant-
bushy-tailed-cloud-rat/crateromys-schadenbergi/info.html. Retrieved 2010-07-08.
42.^ Rabor, D.S. (1986). Guide to Philippine Flora and Fauna.. Natural Resources Management
Centre, Ministry of Natural Resources and University of the Philippines..
43.^ Schofield, James (27 July 2001). "Lake Baikal’s Vanishing Nerpa Seal". The Moscow
Times. http://www.themoscowtimes.com/stories/2001/07/27/106.html. Retrieved 2007-09-27.
44.^ Pallen, Mark (2009). Rough Guide to Evolution. Rough Guides. pp. 87. ISBN 978-1-85828-
946-5.
45.^ http://www.blackwellpublishing.com/ridley/a-z/Ring_species.asp
46.^ http://evolution.berkeley.edu/evolibrary/article/0_0_0/devitt_02
47.^ Prothero, Donald R.; Schoch, Robert M. (2002). Horns, tusks, and flippers: the evolution of
hoofed mammals. JHU press. pp. 45. ISBN 0801871352.
48.^ Tanaka T, Hashimoto H. (1989). "Drug-resistance and its transferability of Shigella strains
isolated in 1986 in Japan". Kansenshogaku Zasshi 63 (1): 15–26. PMID 2501419.
49.^ J. W. Haas, Jr. (June 2000). "The Rev. Dr. William H. Dallinger F.R.S.: Early Advocate of
Theistic Evolution and Foe of Spontaneous Generation". Perspectives on Science and Christian
Faith 52. pp. 107–117. http://www.asa3.org/ASA/PSCF/2000/PSCF6-00Haas.html. Retrieved 2010-
06-15.
50.^ Soy and Lactose Intolerance Wayback: Soy Nutrition
 51.^ Coles Harriet (2007-01-20). "The lactase gene in Africa: Do you take milk?". The Human
Genome, Wellcome Trust. http://genome.wellcome.ac.uk/doc_WTX038968.html. Retrieved 2008-07-
18.
52.^ Thwaites WM (Summer 1985). "New Proteins Without God's Help". Creation Evolution
Journal (National Center for Science Education (NCSE)) 5 (2): 1–3. http://ncse.com/cej/5/2/new-
proteins-without-gods-help.
53.^ Evolution and Information: The Nylon Bug
54.^ Why scientists dismiss 'intelligent design', Ker Than, MSNBC, Sept. 23, 2005
55.^ Miller, Kenneth R. Only a Theory: Evolution and the Battle for America's Soul (2008) pp.
80-82
56.^ a b Science News, Dark Power: Pigment seems to put radiation to good use, Week of May
26, 2007; Vol. 171, No. 21 , p. 325 by Davide Castelvecchi
57.^ Dadachova E, Bryan RA, Huang X, Moadel T, Schweitzer AD, Aisen P, Nosanchuk JD,
Casadevall A. (2007). "Ionizing radiation changes the electronic properties of melanin and enhances
the growth of melanized fungi". PLoS ONE 2 (5): e457. doi:10.1371/journal.pone.0000457.
PMID 17520016. PMC 1866175. http://www.plosone.org/article/fetchArticle.action?articleURI=info
%3Adoi%2F10.1371%2Fjournal.pone.0000457.
58.^ Cheptou, P., Carrue, O., Rouifed, S., Cantarel, A. (2008). "Rapid evolution of seed
dispersal in an urban environment in the weed Crepis sancta.". Proceedings of the National
Academy of Sciences. http://www.pnas.org/content/105/10/3796.abstract.
59.^ "Evolution in the urban jungle". http://theoystersgarter.com/2008/03/12/evolution-in-the-
urban-jungle/. Retrieved 2010-07-08.
60.^ James K. Liebherr and Joseph V. McHugh in Resh, V. H. & R. T. Cardé (Editors) 2003.
Encyclopedia of Insects. Academic Press.
 61.^ Ed Yong (December 3, 2009). "British birdfeeders split blackcaps into two genetically
distinct groups : Not Exactly Rocket Science". ScienceBlogs.
http://scienceblogs.com/notrocketscience/2009/12/british_birdfeeders_split_blackcaps_into_two_gen
etically_dis.php. Retrieved 2010-05-21.
62.^ William R. Rice, George W. Salt (1990). "The Evolution of Reproductive Isolation as a
Correlated Character Under Sympatric Conditions: Experimental Evidence". Evolution, Society for
the Study of Evolution 44.
63.^ "he Evolution of Reproductive Isolation as a Correlated Character Under Sympatric
Conditions: Experimental Evidence". William R. Rice, George W. Salt.
http://www.lifesci.ucsb.edu/eemb/faculty/rice/publications/pdf/25.pdf. Retrieved 2010-05-23.
64.^ "Observed Instances of Speciation, 5.3.5 Sympatric Speciation in Drosophila
melanogaster". Joseph Boxhorn. http://www.talkorigins.org/faqs/faq-speciation.html. Retrieved 2010-
05-23.
65.^ Feder JL, Roethele JB, Filchak K, Niedbalski J, Romero-Severson J (1 March 2003).
"Evidence for inversion polymorphism related to sympatric host race formation in the apple maggot
fly, Rhagoletis pomonella". Genetics 163 (3): 939–53. PMID 12663534. PMC 1462491.
http://www.genetics.org/cgi/pmidlookup?view=long&pmid=12663534.
66.^ Berlocher SH, Bush GL (1982). "An electrophoretic analysis of Rhagoletis (Diptera:
Tephritidae) phylogeny". Systematic Zoology 31 (2): 136–55. doi:10.2307/2413033.
http://jstor.org/stable/2413033.
Berlocher SH, Feder JL (2002). "Sympatric speciation in phytophagous insects: moving beyond
controversy?". Annu Rev Entomol. 47: 773–815. doi:10.1146/annurev.ento.47.091201.145312.
PMID 11729091.
Bush GL (1969). "Sympatric host race formation and speciation in frugivorous flies of the genus
Rhagoletis (Diptera: Tephritidae)". Evolution 23 (2): 237–51. doi:10.2307/2406788.
http://jstor.org/stable/2406788.
Prokopy RJ, Diehl SR, Cooley SS (June 1988). "Behavioral evidence for host races in Rhagoletis
pomonella flies". Oecologia 76 (1): 138–47. doi:10.1007/BF00379612 (inactive 2010-01-08).
http://www.springerlink.com/content/p1716r36n2164855/?
p=d8018d5a59294c2984f253b7152445b7&pi=20.
Feder JL, Roethele JB, Wlazlo B, Berlocher SH (October 1997). "Selective maintenance of allozyme
differences among sympatric host races of the apple maggot fly". Proc Natl Acad Sci USA. 94 (21):
11417–21. doi:10.1073/pnas.94.21.11417. PMID 11038585. PMC 23485.
http://www.pnas.org/cgi/content/full/94/21/11417.
67.^ a b c "London underground source of new insect forms". The Times. 1998-08-26.
http://www.gene.ch/gentech/1998/Jul-Sep/msg00188.html.
68.^ a b Fonseca DM, Keyghobadi N, Malcolm CA, et al. (March 2004). "Emerging vectors in the
Culex pipiens complex". Science 303 (5663): 1535–8. doi:10.1126/science.1094247.
PMID 15001783. http://www.mosquitocatalog.org/files/pdfs/wr380.pdf.
69.^ a b Alan Burdick (2001). "Insect From the Underground — London, England Underground
home to different species of mosquitos". Natural History.
http://findarticles.com/p/articles/mi_m1134/is_1_110/ai_70770157.
70.^ Byrne K, Nichols RA (January 1999). "Culex pipiens in London Underground tunnels:
differentiation between surface and subterranean populations". Heredity 82 (1): 7–15.
doi:10.1038/sj.hdy.6884120. PMID 10200079.
71.^ Vinogradova EB and Shaikevich EV Morphometric, physiological and molecular
characteristics of underground populations of the urban mosquito Culex pipiens Linnaeus f. molestus
Forskål (Diptera: Culicidae) from several areas of Russia European Mosquito Bulletin, 22(2007), 17-
24. Journal of the European Mosquito Control Association ISSN1460-6127
72.^ http://www.nature.com/nature/journal/v403/n6766/full/403158a0.html
73.^ "Giant insect splits cavefish into distinct populations". Ed Yong.
http://scienceblogs.com/notrocketscience/2009/05/giant_insect_splits_cavefish_into_distinct_populat
ions.php. Retrieved 2010-05-22.
74.^ "New plant species arise from conflicts between immune system genes". Ed Yong.
http://scienceblogs.com/notrocketscience/2009/08/new_plant_species_arise_from_conflicts_betwee
n_immune_system.php. Retrieved 2010-05-22.
75.^ Adaptive Traits of the Polar Bear (Ursus Maritimus)
76.^ Polar Bear Evolution
77.^ Ron Rayborne Accepts Hovind's Challenge
78.^ Andrew J. Lowe, Richard J. Abbott (1996). "Origins of the New Allopolyploid Species
‘’Senecio camrensis (asteracea)’’ and its Relationship to the Canary Islands Endemic ‘’Senecio
tenerifae’’". American Journal of Botany 83 (10): 1365–1372. http://www.jstor.org/pss/2446125.
79.^ Jerry A. Coyne (2009). Why Evolution is True. Penguin Group. pp. 187 – 189. ISBN 978-0-
670-02053-9.
80.^ Missouri Botanical Garden. "TROPICOS Web display Senecio vulgaris L.". Nomenclatural
and Specimen Data Base. Missouri State Library. http://mobot.mobot.org/cgi-bin/search_pick?
name=Senecio+vulgaris. Retrieved 2008-02-01.
81.^ Raven, Peter H., et al. (2005). Biology of Plants (7th rev. ed.). New York: W.H. Freeman.
ISBN 0716762846. OCLC 183148564.
82.^
http://www.trnmag.com/Stories/2003/052103/Simulated_evolution_gets_complex_052103.html
 83.^ Adami C, Ofria C, Collier TC (April 2000). "Evolution of biological complexity". Proc Natl
Acad Sci USA. 97 (9): 4463–8. doi:10.1073/pnas.97.9.4463. PMID 10781045. PMC 18257.
http://www.pnas.org/cgi/content/full/97/9/4463.
84.^ Earl DJ, Deem MW (August 2004). "Evolvability is a selectable trait". Proc Natl Acad Sci
USA. 101 (32): 11531–6. doi:10.1073/pnas.0404656101. PMID 15289608. PMC 511006.
http://www.pnas.org/cgi/content/full/101/32/11531.
85.^ a b Stemmer WP (October 1994). "DNA shuffling by random fragmentation and reassembly:
in vitro recombination for molecular evolution". Proc Natl Acad Sci USA. 91 (22): 10747–51.
doi:10.1073/pnas.91.22.10747. PMID 7938023. PMC 45099. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=7938023.
86.^ Sauter E (March 27, 2006). ""Accelerated Evolution" Converts RNA Enzyme to DNA
Enzyme In Vitro". TSRI – News & Views 6 (11).
http://www.scripps.edu/newsandviews/e_20060327/evo.html.
87.^ [3]
88.^ In Vitro Molecular Evolution

[edit] Further reading

• Biological science, Oxford, 2002.
• Clegg CJ (1998). Genetics & evolution. London: J. Murray. ISBN 0-7195-7552-4.
• Coyne, Jerry A. (2009). Why Evolution is True. New York: Oxford University Press.
ISBN 978-0-19-923084-6.
 • Darwin, Charles November 24, 1859. On the Origin of Species by means of Natural
Selection or the Preservation of Favoured Races in the Struggle for Life . London: John Murray,
Albemarle Street. 502 pages. Reprinted: Gramercy (May 22, 1995). ISBN 0-517-12320-7
• Dawkins, Richard (2009). The Greatest Show on Earth: The Evidence for Evolution . Bantam
Press. ISBN 978-1-4165-9478-9.
• Endler, John A. (1986). Natural selection in the wild. New Jersey: Princton University Press.
ISBN 0-691-08387-8.
• Futuyma, D.J. 1998. Evolutionary Biology. 3rd ed. Sinauer Associates, Sunderland,
Massachusetts. (dated 1998, published 1997) ISBN 0-87893-189-9
• Gigerenzer, Gerd (1989). The Empire of chance: how probability changed science and
everyday life. Cambridge, UK: Cambridge University Press. ISBN 0-521-33115-3.
• Hill A, Behrensmeyer AK (1980). Fossils in the making: vertebrate taphonomy and
paleoecology. Chicago: University of Chicago Press. ISBN 0-226-04169-7.
• Ho, YK (2004). Advanced-level Biology for Hong Kong, Manhattan Press. ISBN 962-990-
635-X
• Martin RE (1999). Taphonomy: a process approach. Cambridge, UK: Cambridge University
Press. ISBN 0-521-59833-8.
• Mayr, Ernst (2001). What evolution is. New York: Basic Books. ISBN 0-465-04426-3.
• Paul CRC, Donovan SK (1998). The adequacy of the fossil record. New York: John Wiley.
ISBN 0-471-96988-5.
• Sober, Elliott (2008). Evidence and Evolution: The logic behind the science . Cambridge
University Press. ISBN 978-0-521-87188-4.
[edit] External links
• National Academies Evolution Resources
• TalkOrigins Archive - 29+ Evidences for Macroevolution: The Scientific Case for Common
Descent
• TalkOrigins Archive - Transitional Vertebrate Fossils FAQ
• Understanding Evolution: Your one-stop source for information on evolution
• National Academy Press: Teaching About Evolution and the Nature of Science
• Evolution — Provided by PBS.
• Evolution News from Genome News Network (GNN)
• Evolution by Natural Selection — An introduction to the logic of the theory of evolution by
natural selection
• Howstuffworks.com — How Evolution Works
• 15 Evolutionary Gems

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 W000

Microevolution
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Biology portal · v • d • e
Microevolution is the occurrence of small-scale changes in allele frequencies in a population, over a
few generations, also known as "change below the species level".[1] These changes are due to four different
processes: mutation, selection (natural and artificial), gene flow and genetic drift.
Population genetics is the branch of biology that provides the mathematical structure for the study of
the process of microevolution. Ecological genetics concerns itself with observing microevolution in the wild.
Typically, observable instances of evolution are examples of microevolution; for example, bacterial strains
that have antibiotic resistance.
Microevolution can be contrasted with macroevolution, which is the occurrence of large-scale
changes in gene frequencies in a population over a geological time period (i.e. consisting of extended
microevolution). The difference is largely one of approach. Microevolution is reductionist, but macroevolution
is holistic. Each approach offers different insights into the evolution process. Macroevolution can be seen as
the sum of long periods of microevolution, and thus the two are qualitatively identical while being
quantitatively different.
Contents
[hide]
• 1 Origin of the term
• 2 The four processes
• 2.1 Selection
• 2.2 Genetic drift
• 2.3 Mutation
• 2.4 Gene flow
• 3 Misuse
• 4 See also
• 5 References
• 6 External links

[edit] Origin of the term

The term was first used by Harvard-educated botanist Robert Greenleaf Leavitt in the journal
Botanical Gazette in 1909, addressing what he called the "mystery" of how formlessness gives rise to form.
[2]
..The production of form from formlessness in the egg-derived individual, the multiplication of
parts and the orderly creation of diversity among them, in an actual evolution, of which anyone may
 ascertain the facts, but of which no one has dissipated the mystery in any significant measure. This
microevolution forms an integral part of the grand evolution problem and lies at the base of it, so that
we shall have to understand the minor process before we can thoroughly comprehend the more
general one...
However, Leavitt was using the term to describe what we would now call developmental biology, it
was not until Russian Entomologist Yuri Filipchenko used the terms "macroevolution" and "microevolution" in
1927 in his German language work, "Variabilität und Variation", that it attained its modern usage. The term
was later brought into the English-speaking world by Theodosius Dobzhansky in his book Genetics and the
Origin of Species (1937).[1]

[edit] The four processes

[edit] Selection
Main articles: Natural selection and Artificial selection
Selection is the process by which heritable traits that make it more likely for an organism to survive
and successfully reproduce become more common in a population over successive generations.
It is sometimes valuable to distinguish between naturally occurring selection, natural selection, and
selection that is a manifestation of choices made by humans, artificial selection. This distinction is rather
diffuse. Natural selection is nevertheless the dominant part of selection.
 Natural selection of a population for dark coloration.
The natural genetic variation within a population of organisms means that some individuals will
survive more successfully than others in their current environment. Factors which affect reproductive success
are also important, an issue which Charles Darwin developed in his ideas on sexual selection.
 Natural selection acts on the phenotype, or the observable characteristics of an organism, but the
genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common
in a population (see allele frequency). Over time, this process can result in adaptations that specialize
organisms for particular ecological niches and may eventually result in the emergence of new species.
Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin
in his groundbreaking 1859 book On the Origin of Species,[3] in which natural selection was described by
analogy to artificial selection, a process by which animals and plants with traits considered desirable by
human breeders are systematically favored for reproduction. The concept of natural selection was originally
developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of
modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and
molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary
explanation for adaptive evolution.

[edit] Genetic drift

Main article: Genetic drift
Genetic drift is the change in the relative frequency in which a gene variant (allele) occurs in a
population due to random sampling and chance. That is, the alleles in the offspring in the population are a
random sample of those in the parents. And chance has a role in determining whether a given individual
survives and reproduces. A population's allele frequency is the fraction or percentage of its gene copies
compared to the total number of gene alleles that share a particular form.[4]
Genetic drift is an important evolutionary process which leads to changes in allele frequencies over
time. It may cause gene variants to disappear completely, and thereby reduce genetic variability. In contrast
to natural selection, which makes gene variants more common or less common depending on their
reproductive success,[5] the changes due to genetic drift are not driven by environmental or adaptive
pressures, and may be beneficial, neutral, or detrimental to reproductive success.
The effect of genetic drift is larger in small populations, and smaller in large populations. Vigorous
debates wage among scientists over the relative importance of genetic drift compared with natural selection.
Ronald Fisher held the view that genetic drift plays at the most a minor role in evolution, and this remained
the dominant view for several decades. In 1968 Motoo Kimura rekindled the debate with his neutral theory of
molecular evolution which claims that most of the changes in the genetic material are caused by genetic drift.
[6]

[edit] Mutation
Main article: Mutation
 Duplication of part of a chromosome
Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation, viruses,
transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA replication.[7][8][9]
Errors are introduced particularly often in the process of DNA replication, in the polymerization of the second
strand. These errors can also be induced by the organism itself, by cellular processes such as
hypermutation.
Mutations can have an impact on the phenotype of an organism, especially if they occur within the
protein coding sequence of a gene. Error rates are usually very low—1 error in every 10 million–100 million
bases—due to the "proofreading" ability of DNA polymerases.[10][11] Without proofreading, error rates are a
thousandfold higher. Chemical damage to DNA occurs naturally as well, and cells use DNA repair
mechanisms to repair mismatches and breaks in DNA. Nevertheless, the repair sometimes fails to return the
DNA to its original sequence.
In organisms that use chromosomal crossover to exchange DNA and recombine genes, errors in
alignment during meiosis can also cause mutations.[12] Errors in crossover are especially likely when similar
sequences cause partner chromosomes to adopt a mistaken alignment; this makes some regions in
genomes more prone to mutating in this way. These errors create large structural changes in DNA sequence
—duplications, inversions or deletions of entire regions, or the accidental exchanging of whole parts between
different chromosomes (called translocation).
Mutation can result in several different types of change in DNA sequences; these can either have no
effect, alter the product of a gene, or prevent the gene from functioning. Studies in the fly Drosophila
melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably be harmful,
with about 70 percent of these mutations having damaging effects, and the remainder being either neutral or
weakly beneficial.[13] Due to the damaging effects that mutations can have on cells, organisms have evolved
mechanisms such as DNA repair to remove mutations.[7] Therefore, the optimal mutation rate for a species
is a trade-off between costs of a high mutation rate, such as deleterious mutations, and the metabolic costs
of maintaining systems to reduce the mutation rate, such as DNA repair enzymes.[14] Viruses that use RNA
as their genetic material have rapid mutation rates,[15] which can be an advantage since these viruses will
evolve constantly and rapidly, and thus evade the defensive responses of e.g. the human immune system.
[16]
Mutations can involve large sections of DNA becoming duplicated, usually through genetic
recombination.[17] These duplications are a major source of raw material for evolving new genes, with tens
to hundreds of genes duplicated in animal genomes every million years.[18] Most genes belong to larger
families of genes of shared ancestry.[19] Novel genes are produced by several methods, commonly through
the duplication and mutation of an ancestral gene, or by recombining parts of different genes to form new
combinations with new functions.[20][21]
Here, domains act as modules, each with a particular and independent function, that can be mixed
together to produce genes encoding new proteins with novel properties.[22] For example, the human eye
uses four genes to make structures that sense light: three for color vision and one for night vision; all four
arose from a single ancestral gene.[23] Another advantage of duplicating a gene (or even an entire genome)
is that this increases redundancy; this allows one gene in the pair to acquire a new function while the other
copy performs the original function.[24][25] Other types of mutation occasionally create new genes from
previously noncoding DNA.[26][27]

[edit] Gene flow

Main article: Gene flow
Gene flow is the exchange of genes between populations, which are usually of the same species.[28]
Examples of gene flow within a species include the migration and then breeding of organisms, or the
exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and
horizontal gene transfer.
 Migration into or out of a population can change allele frequencies, as well as introducing genetic
variation into a population. Immigration may add new genetic material to the established gene pool of a
population. Conversely, emigration may remove genetic material. As barriers to reproduction between two
diverging populations are required for the populations to become new species, gene flow may slow this
process by spreading genetic differences between the populations. Gene flow is hindered by mountain
ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has
hindered the flow of plant genes.[29]
Depending on how far two species have diverged since their most recent common ancestor, it may
still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.[30]
Such hybrids are generally infertile, due to the two different sets of chromosomes being unable to pair up
during meiosis. In this case, closely related species may regularly interbreed, but hybrids will be selected
against and the species will remain distinct. However, viable hybrids are occasionally formed and these new
species can either have properties intermediate between their parent species, or possess a totally new
phenotype.[31] The importance of hybridization in creating new species of animals is unclear, although cases
have been seen in many types of animals,[32] with the gray tree frog being a particularly well-studied
example.[33]
Hybridization is, however, an important means of speciation in plants, since polyploidy (having more
than two copies of each chromosome) is tolerated in plants more readily than in animals.[34][35] Polyploidy is
important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to
pair with an identical partner during meiosis.[36] Polyploids also have more genetic diversity, which allows
them to avoid inbreeding depression in small populations.[37]
Horizontal gene transfer is the transfer of genetic material from one organism to another organism
that is not its offspring; this is most common among bacteria.[38] In medicine, this contributes to the spread
of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other
species.[39] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces
cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred.[40][41] An
example of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have received a range
of genes from bacteria, fungi, and plants.[42] Viruses can also carry DNA between organisms, allowing
transfer of genes even across biological domains.[43] Large-scale gene transfer has also occurred between
the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.
[44]
Gene flow is the transfer of alleles from one population to another.
Migration into or out of a population may be responsible for a marked change in allele frequencies.
Immigration may also result in the addition of new genetic variants to the established gene pool of a
particular species or population.
There are a number of factors that affect the rate of gene flow between different populations. One of
the most significant factors is mobility, as greater mobility of an individual tends to give it greater migratory
potential. Animals tend to be more mobile than plants, although pollen and seeds may be carried great
distances by animals or wind.
Maintained gene flow between two populations can also lead to a combination of the two gene pools,
reducing the genetic variation between the two groups. It is for this reason that gene flow strongly acts
against speciation, by recombining the gene pools of the groups, and thus, repairing the developing
differences in genetic variation that would have led to full speciation and creation of daughter species.
For example, if a species of grass grows on both sides of a highway, pollen is likely to be transported
from one side to the other and vice versa. If this pollen is able to fertilise the plant where it ends up and
produce viable offspring, then the alleles in the pollen have effectively been able to move from the population
on one side of the highway to the other.

[edit] Misuse
Main article: Speciation
The term 'microevolution' has recently become popular among the anti-evolution movement, and in
particular among young Earth creationists, to distinguish between evolutionary changes among populations
which can interbreed (microevolution) and changes among populations which cannot interbreed
(macroevolution). The main argument is that change beyond the species level would require similar enough
changes in both a male and female in the same place and time, and that those two would have to
successfully interbreed in order to create a sustainable population. Although theoretically possible, the
combination of factors are seen to be so improbable as to become implausible.
The claim that microevolution is qualitatively different from macroevolution is fallacious, as the main
difference between the two processes is that one occurs within a few generations, whilst the other takes
place over thousands of years (i.e. a quantitative difference).[45][not in citation given] Essentially they
describe the same process; although evolution beyond the species level results in beginning and ending
generations which could not interbreed, the intermediate generations could. Even changes in the number of
chromosomes can be accounted for by intermediate stages in which a single chromosome divides in
generational stages, or multiple chromosomes fuse. A well documented example is the chromosome
difference between humans and great apes.[46]
The attempt to differentiate between microevolution and macroevolution is considered to have no
scientific basis by any mainstream scientific organization, including the American Association for the
Advancement of Science.[47] Contrary to belief among the anti-evolution movement proponents, evolution of
life forms beyond the species level ("macroevolution", i.e. speciation) has indeed been observed and
documented by scientists on many occasions.[48]

[edit] See also

• Macroevolution

[edit] References
1. ^ a b Macroevolution: Its definition, Philosophy and History
2. ^ Leavitt, Robert Botanical Gazette 1909 vol.47 no.1 Jan. A Vegetative Mutant, and the
Principle of Homoeosis in Plants http://www.jstor.org/pss/2466778
3. ^ Darwin C (1859) On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life John Murray, London; modern reprint
Charles Darwin, Julian Huxley (2003). The Origin of Species. Signet Classics. ISBN 0-451-52906-5.
Published online at The complete work of Charles Darwin online: On the origin of species by means
of natural selection, or the preservation of favoured races in the struggle for life.
4. ^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. Glossary. ISBN 0-
87893-189-9.
5. ^ Avers, Charlotte (1989), Process and Pattern in Evolution, Oxford University Press
6. ^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. 320. ISBN 0-
87893-189-9.
 7. ^ a b Bertram J (2000). "The molecular biology of cancer". Mol. Aspects Med. 21 (6): 167–223.
doi:10.1016/S0098-2997(00)00007-8. PMID 11173079.
8. ^ Aminetzach YT, Macpherson JM, Petrov DA (2005). "Pesticide resistance via transposition-
mediated adaptive gene truncation in Drosophila". Science 309 (5735): 764–7.
doi:10.1126/science.1112699. PMID 16051794.
9. ^ Burrus V, Waldor M (2004). "Shaping bacterial genomes with integrative and conjugative
elements". Res. Microbiol. 155 (5): 376–86. doi:10.1016/j.resmic.2004.01.012. PMID 15207870.
10.^ Griffiths, Anthony J. F.; Miller, Jeffrey H.; Suzuki, David T. et al., eds (2000). "Spontaneous
mutations". An Introduction to Genetic Analysis (7th ed.). New York: W. H. Freeman. ISBN 0-7167-
3520-2. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=iga.section.2706.
11.^ Freisinger, E; Grollman; Miller; Kisker (2004). "Lesion (in)tolerance reveals insights into
DNA replication fidelity.". The EMBO journal 23 (7): 1494–505. doi:10.1038/sj.emboj.7600158.
PMID 15057282.
12.^ Griffiths, Anthony J. F.; Miller, Jeffrey H.; Suzuki, David T. et al., eds (2000). "Chromosome
Mutation I: Changes in Chromosome Structure: Introduction". An Introduction to Genetic Analysis
(7th ed.). New York: W. H. Freeman. ISBN 0-7167-3520-2.
http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=iga.section.2844.
13.^ Sawyer SA, Parsch J, Zhang Z, Hartl DL (2007). "Prevalence of positive selection among
nearly neutral amino acid replacements in Drosophila". Proc. Natl. Acad. Sci. U.S.A. 104 (16): 6504–
10. doi:10.1073/pnas.0701572104. PMID 17409186.
14.^ Sniegowski P, Gerrish P, Johnson T, Shaver A (2000). "The evolution of mutation rates:
separating causes from consequences". Bioessays 22 (12): 1057–66. doi:10.1002/1521-
1878(200012)22:12<1057::AID-BIES3>3.0.CO;2-W. PMID 11084621.
 15.^ Drake JW, Holland JJ (1999). "Mutation rates among RNA viruses". Proc. Natl. Acad. Sci.
U.S.A. 96 (24): 13910–3. doi:10.1073/pnas.96.24.13910. PMID 10570172. PMC 24164.
http://www.pnas.org/content/96/24/13910.long.
16.^ Holland J, Spindler K, Horodyski F, Grabau E, Nichol S, VandePol S (1982). "Rapid
evolution of RNA genomes". Science 215 (4540): 1577–85. doi:10.1126/science.7041255.
PMID 7041255.
17.^ Hastings, P J; Lupski, JR; Rosenberg, SM; Ira, G (2009). "Mechanisms of change in gene
copy number". Nature Reviews. Genetics 10 (8): 551–564. doi:10.1038/nrg2593. PMID 19597530.
18.^ Carroll SB, Grenier J, Weatherbee SD (2005). From DNA to Diversity: Molecular Genetics
and the Evolution of Animal Design. Second Edition . Oxford: Blackwell Publishing. ISBN 1-4051-
1950-0.
19.^ Harrison P, Gerstein M (2002). "Studying genomes through the aeons: protein families,
pseudogenes and proteome evolution". J Mol Biol 318 (5): 1155–74. doi:10.1016/S0022-
2836(02)00109-2. PMID 12083509.
20.^ Orengo CA, Thornton JM (2005). "Protein families and their evolution-a structural
perspective". Annu. Rev. Biochem. 74: 867–900. doi:10.1146/annurev.biochem.74.082803.133029.
PMID 15954844.
21.^ Long M, Betrán E, Thornton K, Wang W (November 2003). "The origin of new genes:
glimpses from the young and old". Nat. Rev. Genet. 4 (11): 865–75. doi:10.1038/nrg1204.
PMID 14634634.
22.^ Wang M, Caetano-Anollés G (2009). "The evolutionary mechanics of domain organization
in proteomes and the rise of modularity in the protein world". Structure 17 (1): 66–78.
doi:10.1016/j.str.2008.11.008. PMID 19141283.
 23.^ Bowmaker JK (1998). "Evolution of colour vision in vertebrates". Eye (London, England) 12
(Pt 3b): 541–7. PMID 9775215.
24.^ Gregory TR, Hebert PD (1999). "The modulation of DNA content: proximate causes and
ultimate consequences". Genome Res. 9 (4): 317–24. doi:10.1101/gr.9.4.317 (inactive 2009-11-14).
PMID 10207154. http://genome.cshlp.org/content/9/4/317.full.
25.^ Hurles M (July 2004). "Gene duplication: the genomic trade in spare parts". PLoS Biol. 2
(7): E206. doi:10.1371/journal.pbio.0020206. PMID 15252449.
26.^ Liu N, Okamura K, Tyler DM (2008). "The evolution and functional diversification of animal
microRNA genes". Cell Res. 18 (10): 985–96. doi:10.1038/cr.2008.278. PMID 18711447.
PMC 2712117. http://www.nature.com/cr/journal/v18/n10/full/cr2008278a.html.
27.^ Siepel A (October 2009). "Darwinian alchemy: Human genes from noncoding DNA".
Genome Res. 19 (10): 1693–5. doi:10.1101/gr.098376.109. PMID 19797681. PMC 2765273.
http://genome.cshlp.org/content/19/10/1693.full.
28.^ Morjan C, Rieseberg L (2004). "How species evolve collectively: implications of gene flow
and selection for the spread of advantageous alleles". Mol. Ecol. 13 (6): 1341–56. doi:10.1111/j.1365-
294X.2004.02164.x. PMID 15140081.
29.^ Su H, Qu L, He K, Zhang Z, Wang J, Chen Z, Gu H (2003). "The Great Wall of China: a
physical barrier to gene flow?". Heredity 90 (3): 212–9. doi:10.1038/sj.hdy.6800237.
PMID 12634804.
30.^ Short RV (1975). "The contribution of the mule to scientific thought". J. Reprod. Fertil.
Suppl. (23): 359–64. PMID 1107543.
31.^ Gross B, Rieseberg L (2005). "The ecological genetics of homoploid hybrid speciation". J.
Hered. 96 (3): 241–52. doi:10.1093/jhered/esi026. PMID 15618301.
 32.^ Burke JM, Arnold ML (2001). "Genetics and the fitness of hybrids". Annu. Rev. Genet. 35:
31–52. doi:10.1146/annurev.genet.35.102401.085719. PMID 11700276.
33.^ Vrijenhoek RC (2006). "Polyploid hybrids: multiple origins of a treefrog species". Curr. Biol.
16 (7): R245. doi:10.1016/j.cub.2006.03.005. PMID 16581499.
34.^ Wendel J (2000). "Genome evolution in polyploids". Plant Mol. Biol. 42 (1): 225–49.
doi:10.1023/A:1006392424384. PMID 10688139.
35.^ Sémon M, Wolfe KH (2007). "Consequences of genome duplication". Curr Opin Genet Dev
17 (6): 505–12. doi:10.1016/j.gde.2007.09.007. PMID 18006297.
36.^ Comai L (2005). "The advantages and disadvantages of being polyploid". Nat. Rev. Genet.
6 (11): 836–46. doi:10.1038/nrg1711. PMID 16304599.
37.^ Soltis P, Soltis D (June 2000). "The role of genetic and genomic attributes in the success of
polyploids". Proc. Natl. Acad. Sci. U.S.A. 97 (13): 7051–7. doi:10.1073/pnas.97.13.7051.
PMID 10860970.
38.^ Boucher Y, Douady CJ, Papke RT, Walsh DA, Boudreau ME, Nesbo CL, Case RJ, Doolittle
WF (2003). "Lateral gene transfer and the origins of prokaryotic groups". Annu Rev Genet 37: 283–
328. doi:10.1146/annurev.genet.37.050503.084247. PMID 14616063.
39.^ Walsh T (2006). "Combinatorial genetic evolution of multiresistance". Curr. Opin. Microbiol.
9 (5): 476–82. doi:10.1016/j.mib.2006.08.009. PMID 16942901.
40.^ Kondo N, Nikoh N, Ijichi N, Shimada M, Fukatsu T (2002). "Genome fragment of Wolbachia
endosymbiont transferred to X chromosome of host insect". Proc. Natl. Acad. Sci. U.S.A. 99 (22):
14280–5. doi:10.1073/pnas.222228199. PMID 12386340.
41.^ Sprague G (1991). "Genetic exchange between kingdoms". Curr. Opin. Genet. Dev. 1 (4):
530–3. doi:10.1016/S0959-437X(05)80203-5. PMID 1822285.
 42.^ Gladyshev EA, Meselson M, Arkhipova IR (May 2008). "Massive horizontal gene transfer in
bdelloid rotifers". Science 320 (5880): 1210–3. doi:10.1126/science.1156407. PMID 18511688.
43.^ Baldo A, McClure M (1 September 1999). "Evolution and horizontal transfer of dUTPase-
encoding genes in viruses and their hosts". J. Virol. 73 (9): 7710–21. PMID 10438861.
44.^ Poole A, Penny D (2007). "Evaluating hypotheses for the origin of eukaryotes". Bioessays
29 (1): 74–84. doi:10.1002/bies.20516. PMID 17187354.
45.^ Claim CB902: "Microevolution is distinct from macroevolution", TalkOrigins Archive
46.^ http://www.gate.net/~rwms/hum_ape_chrom.html
47.^ Evolution Press Release American Association for the Advancement of Science
48.^ Complete sourced list of observed instances of speciation, TalkOrigins Archive

[edit] External links

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• Microevolution vs Macroevolution

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• This page was last modified on 16 August 2010 at 10:51.

 W000

Microévolution et macroévolution
Un article de Wikipédia, l'encyclopédie libre.
(Redirigé depuis Microevolution)
Aller à : Navigation, rechercher
Cet article est une ébauche concernant la biologie.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
Dans diverses approches scientifiques non darwiniennes ou créationnistes, les termes de
microévolution et macroévolution séparent l'étude des phénomènes d'évolution en fonction de la grandeur
d'échelle d'étude : la microévolution pour les changements au sein d'une même espèce, et la macroévolution
pour ceux qui ont lieu au stade supérieur à l'espèce.
 Le terme de microévolution est généralement utilisée pour signifier que l'axe d'étude porte sur les
modifications ayant lieu au sein d'une population donnée, et observées le plus souvent sur une courte
échelle de temps (typiquement quelques générations)[1]. Dans ce cadre, l'étude s'intéresse surtout à la
modification des fréquences alléliques dans de petites populations isolées. Ces changements peuvent avoir
plusieurs moteurs, qui agissent simultanément, dont notamment les mutations, la sélection naturelle ou
artificielle, et la dérive génétique.
Le terme de macroévolution est généralement utilisé en paléontologie pour indiquer que le niveau
d'étude porte sur des changements au-dessus du niveau de l'espèce, comme le clade, etc, et porte
principalement sur le phénomène de spéciation et l'évolution dans le temps des groupes d'espèces.
Ces deux termes sont attachés à divers courants scientifiques non darwiniens ou créationnistes,
pour tenter de séparer arbitrairement les phénomènes d'évolutions lorsqu'ils ont lieu au sein d'une même
espèce (microévolution), et ceux qui ont lieu à un niveau supérieur à l'espèce (macroévolution), qui opèrent
généralement sur une échelle de temps géologique[1][2]. Cette distinction repose essentiellement sur une
différence d'échelle de temps, les deux phénomènes étant les manifestations du même processus
d'évolution, selon la théorie synthétique de l'évolution.
Sommaire
[masquer]
• 1 Origine des termes
• 2 La microévolution
• 3 La spéciation
• 4 Macroévolution
• 5 La macroévolution en paléontologie
• 6 Usage des termes dans le créationisme
• 7 Notes et références
• 8 Voir aussi

Origine des termes [modifier]

L'entomologiste de russe Yuri Filipchenko est le premier à faire usage des termes
« macroevolution » et « microevolution », en 1927 et en langue allemande dans sa publication Variabilität
und Variation. Filipchenko est un orthogénéticien : il reste persuadé que l'évolution subit une force dirigiste.
Ce n'est pas un darwinien, et il rejette notamment le concept de compétition dans la sélection naturelle. Ces
termes passent à l'anglais à travers la publication de son élève Theodosius Dobzhansky, Genetics and the
Origin of Species, publié en 1937[1]. Ce livre est l'une des principales publications scientifique dans
l'élaboration d'une synthèse théorique de l'évolution, qui conduisit plus tard à la formulation de la Théorie
synthétique de l'évolution.
La microévolution [modifier]
L'usage du terme microévolution indique que le niveau d'étude des phénomènes d'évolutions est
restreint à celui de la population[3]. Une population est un groupe d'individus d'une même espèce sexuée qui
se reproduisent entre eux, comme par exemple un groupe d'insectes isolés sur une île.
À cette échelle, l'étude de l'évolution des caractéristiques des individus due au processus d'évolution
porte essentiellement sur des changements dans la fréquence de certains allèles d'un ou plusieurs gènes.
Un gène est l'emplacement d'un groupe de nucléotides sur un chromosome. L'arrangement de ces
nucléotides peut varier, et chaque combinaison particulière forme un allèle de ce gène. Ces allèles sont
produits par mutation génétique et hérités par la descendance. La proportion d'un type d'allèle au sein de la
population étudiée varie en fonction de certains paramètres :
• La mutation peut être non viable pour l'organisme ou pathogène.
• Des migrations de population : des reproducteurs apportent de nouveaux allèles dans le pool
génétique de la population, d'autres s'en séparent, ou la population est isolée géographiquement des
autres membres de l'espèce.
• Le hasard : c'est la dérive génétique.
• Des paramètres du milieu : c'est la sélection naturelle.
Ces mutations peuvent s'accumuler par évolution progressive jusqu'à empêcher la reproduction avec
les autres descendants de la même espèce : ils ne sont plus interféconds. C'est le phénomène de spéciation,
et c'est ici que qu'une distinction est parfois faite avec la « macroévolution » .
 Cette définition se restreint donc à la reproduction sexuée, mais les phénomènes sont similaires
chez les espèces asexuées. L'un des exemple typiques de « micro évolution » fréquemment cité dans ce cas
là est l'acquisition d'une résistance aux antibiotiques par une bactérie.

La spéciation [modifier]

Macroévolution [modifier]

La macroévolution en paléontologie [modifier]

Le terme de macroévolution est généralement utilisé pour créer une distinction artificielle entre les
phénomènes dits "de microévolution", qui sont actuels, observables et expérimentables, et relèvent de la
génétique ; et les changements dans la morphologie des fossiles et leur classement, en paléontologie. Il est
fréquemment utilisée dans des théories opposées au néo-darwinisme, en présupposant qu'il existe "autre
chose" qui influe sur l'évolution entre les espèces. Le terme se retrouve ainsi dans les théories
orthogenétiques, la théorie des équilibres ponctués, intelligent design, etc. Une partie des néo-darwinien,
comme John Maynard Smith ou Richard Dawkins, en rejette donc l'usage, pour éviter toute confusion
laissant entendre qu'il existe une distinction entre les mécanismes d'évolution au sein d'une espèce, et
l'évolution entre les espèces[1].
Les deux termes sont repris par Richard Goldschmidt et le paléontologue Otto Schindewolf dans
leurs travaux sur l'orthogenèse, une théorie non-darwinienne sur une évolution linéaire qui présuppose un
dirigisme dans les modifications morphologiques des fossiles d'une même lignée.
 Des paléontologues comme Grantham, Rice, et Stidd and Wade ont tenté d'appliquer les
phénomènes de sélection au niveau des espèces, en imaginant une sélection spécifique, où le milieu
sélectionnerait les espèces en fonctions de leur caractères notamment morphologiques, par analogie avec la
façon dont la sélection naturelle sélectionne les gènes.
Le terme de macroévolution est utilisé par des paléontologues comme Steven Stanley, ou Stephen
Jay Gould et Niles Eldredge, les fondateurs de la théorie des équilibres ponctués.

Usage des termes dans le créationisme [modifier]

Notes et références [modifier]

1. ↑ a, b, c et d (en)Macroevolution: Its definition, Philosophy and History [archive], John
Wilkins, 1997-2006
2. ↑ C'est généralement le cas, mais une spéciation peut avoir lieu sur une échelle de temps
restreinte
3. ↑ (en)Understanding evolution, University of California Museum of Paleontology [archive]

Voir aussi [modifier]

• adaptation
• spéciation
• évolution
 [Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
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• Portail de l’origine et de l’évolution du vivant

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 W000

Macroevolution
From Wikipedia, the free encyclopedia

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Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
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Macroevolution is a scale of analysis of evolution in separated gene pools.[1] Macroevolutionary
studies focus on change that occurs at or above the level of species, in contrast with microevolution,[2] which
refers to smaller evolutionary changes (typically described as changes in allele frequencies) within a species
or population.[3]
 The process of speciation may fall within the purview of either, depending on the forces thought to
drive it. Paleontology, evolutionary developmental biology, comparative genomics and genomic
phylostratigraphy contribute most of the evidence for the patterns and processes that can be classified as
macroevolution. An example of macroevolution is the appearance of feathers during the evolution of birds
from theropod dinosaurs.
Abrupt transformations from a biologic system to another, for example the passing of life from water
into land or the transition from invertebrates to vertebrates, are rare. Few major biological types have
emerged during the evolutionary history of life and most of them survive till today. When lifeforms take such
giant leaps, they meet little to no competition and are able to exploit a plethora of available niches, following
a pattern of adaptive radiation. This can lead to convergent evolution, where unrelated populations display
similar adaptations.[4]
The evolutionary course of Equidae (wide family including all horses and related animals) is often
viewed as a typical example of macroevolution. The earliest known genus, Hyracotherium (now reclassified
as a palaeothere), was a herbivore animal resembling a dog that lived in the early cenozoic. As its habitat
transformed into an open arid grassland, selective pressure required that the animal become a fast grazer.
Thus elongation of legs and head as well as reduction of toes gradually occurred, producing the only extant
genus of Equidae, Equus.[4]
Contents
[hide]
• 1 Origin of the term
• 2 Macroevolution and the modern evolutionary synthesis
• 3 Research topics
• 4 Misuse
• 5 See also
• 6 References
• 7 External links

[edit] Origin of the term

Russian entomologist Yuri Filipchenko first coined the terms "macroevolution" and "microevolution"
in 1927 in his German language work, "Variabilität und Variation". Since the inception of the two terms, their
meanings have been revised several times and the term macroevolution fell into limited disfavour when it
was taken over by such writers as the geneticist Richard Goldschmidt (1940) and the paleontologist Otto
Schindewolf to describe their orthogenetic theories.[5]
As a result, apart from Dobzhansky, Bernhard Rensch and Ernst Mayr, very few neo-Darwinian
writers used the term, preferring instead to talk of evolution as changes in allele frequencies without mention
of the level of the changes (above species level or below). Those who did were generally working within the
continental European traditions (as Dobzhansky, Ernst Mayr, Bernhard Rensch, Richard Goldschmidt, and
Otto Schindewolf were) and those who didn't were generally working within the Anglo-American tradition
(such as John Maynard Smith and Richard Dawkins). Hence, use of the term "macroevolution" is sometimes
wrongly used as a litmus test of whether the writer is "properly" neo-Darwinian or not.[5]

[edit] Macroevolution and the modern evolutionary synthesis

Within the Modern Synthesis school of thought, macroevolution is thought of as the compounded
effects of microevolution.[6] Thus, the distinction between micro- and macroevolution is not a fundamental
one – the only difference between them is of time and scale. As Ernst W. Mayr observes, "transspecific
evolution is nothing but an extrapolation and magnification of the events that take place within populations
and species...it is misleading to make a distinction between the causes of micro- and macroevolution”.[6]
However, time is not a necessary distinguishing factor – macroevolution can happen without gradual
compounding of small changes; whole-genome duplication can result in speciation occurring over a single
generation - this is especially common in plants.[7]
Changes in the genes regulating development have also been proposed as being important in
producing speciation through large and relatively sudden changes in animals' morphology.[8][9]

[edit] Research topics

Some examples of subjects whose study falls within the realm of macroevolution:
• Adaptive radiations such as The Cambrian Explosion.
• Changes in biodiversity through time.
 • Genomic evolution, like horizontal gene transfer, genome fusions in endosymbioses, and
adaptive changes in genome size.
• Mass extinctions.
• Speciation and extinction rates.
• The debate between punctuated equilibrium and gradualism.
• The role of development in shaping evolution, particularly such topics as heterochrony and
phenotypic plasticity.

[edit] Misuse
Main article: Objections to evolution
See also: Speciation
The term "macroevolution" frequently arises within the context of the evolution/creation debate,
usually used by creationists alleging a significant difference between the evolutionary changes observed in
field and laboratory studies and the larger scale macroevolutionary changes that scientists believe to have
taken thousands or millions of years to occur. They may accept that evolutionary change is possible within
species ("microevolution"), but deny that one species can evolve into another ("macroevolution").[1] Contrary
to this belief among the anti-evolution movement proponents, evolution of life forms beyond the species level
("macroevolution", i.e. speciation in a specific case) has indeed been observed multiple times under both
controlled laboratory conditions and in nature.[10] The claim that macroevolution does not occur, or is
impossible, is thus demonstrably false and without support in the scientific community.
Such claims are rejected by the scientific community on the basis of ample evidence that
macroevolution is an active process both presently and in the past.[5][11] The terms macroevolution and
microevolution relate to the same processes operating at different scales, but creationist claims misuse the
terms in a vaguely defined way which does not accurately reflect scientific usage, acknowledging well
observed evolution as "microevolution" and denying that "macroevolution" takes place.[5][12] Evolutionary
theory (including macroevolutionary change) remains the dominant scientific paradigm for explaining the
origins of Earth's biodiversity. Its occurrence is not disputed within the scientific community.[13] While details
of macroevolution are continuously studied by the scientific community, the overall theory behind
macroevolution (i.e. common descent) has been overwhelmingly consistent with empirical data. Predictions
of empirical data from the theory of common descent have been so consistent that biologists often refer to it
as the "fact of evolution".[14][15]
Nicholas Matzke and Paul R. Gross have accused creationists of using "strategically elastic"
definitions of micro- and macroevolution when discussing the topic.[1] The actual definition of macroevolution
accepted by scientists is "any change at the species level or above" (phyla, group, etc.) and microevolution is
"any change below the level of species." Matzke and Gross state that many creationist critics define
macroevolution as something that cannot be attained, as these critics describe any observed evolutionary
change as "just microevolution".[1]

[edit] See also

• List of transitional fossils
• Transitional fossil
• Microevolution
• Speciation
[edit] References
1. ^ a b c d Matzke, Nicholas J. and Paul R. Gross. 2006. Analyzing Critical Analysis: The
Fallback Antievolutionist Strategy. In Eugenie Scott and Glenn Branch, Not in Our Classrooms: Why
Intelligent Design is Wrong for Our Schools, Beacon Press, Boston ISBN 0807032786[page needed]
2. ^ Dobzhansky, Theodosius Grigorievich (1937). Genetics and the origin of species. New
York: Columbia Univ. Press. p. 12. LCCN 37-033383.
3. ^ Reznick DN, Ricklefs RE (February 2009). "Darwin's bridge between microevolution and
macroevolution". Nature 457 (7231): 837–42. doi:10.1038/nature07894. PMID 19212402.
4. ^ a b Introduction to Ecology (1983) - J.C. Emberlin, chapter 8
5. ^ a b c d Macroevolution: Its definition, Philosophy and History
6. ^ a b Kutschera U, Niklas KJ (June 2004). "The modern theory of biological evolution: an
expanded synthesis". Die Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y.
PMID 15241603.
7. ^ Rieseberg LH, Willis JH (August 2007). "Plant speciation". Science 317 (5840): 910–4.
doi:10.1126/science.1137729. PMID 17702935.
8. ^ Valentine JW, Jablonski D (2003). "Morphological and developmental macroevolution: a
paleontological perspective". The International Journal of Developmental Biology 47 (7-8): 517–22.
PMID 14756327. http://www.intjdevbiol.com/paper.php?doi=14756327.
9. ^ Johnson NA, Porter AH (2001). "Toward a new synthesis: population genetics and
evolutionary developmental biology". Genetica 112-113: 45–58. doi:10.1023/A:1013371201773.
PMID 11838782.
10.^ Rice, W.R.; Hostert (1993). "Laboratory experiments on speciation: what have we learned
in 40 years". Evolution 47 (6): 1637–1653. doi:10.2307/2410209. http://links.jstor.org/sici?sici=0014-
 3820(199312)47%3A6%3C1637%3ALEOSWH%3E2.0.CO%3B2-T. Retrieved 2008-05-19.
*Jiggins CD, Bridle JR (2004). "Speciation in the apple maggot fly: a blend of vintages?". Trends
Ecol. Evol. (Amst.) 19 (3): 111–4. doi:10.1016/j.tree.2003.12.008. PMID 16701238.
*Boxhorn, J (1995). "Observed Instances of Speciation". TalkOrigins Archive.
http://www.talkorigins.org/faqs/faq-speciation.html. Retrieved 2008-12-26.
*Weinberg JR, Starczak VR, Jorg, D (1992). "Evidence for Rapid Speciation Following a Founder
Event in the Laboratory". Evolution 46 (4): 1214–20. doi:10.2307/2409766.
http://jstor.org/stable/2409766.
*Kirkpatrick, Mark; Virginie Ravigné (2002-03). "Speciation by Natural and Sexual Selection: Models
and Experiments". The American Naturalist 159 (3): S22–S35. doi:10.1086/338370. ISSN 00030147.
PMID 18707367. http://www.jstor.org/stable/3078919. Retrieved 2010-03-11.
11.^ CB901: No Macroevolution
12.^ CB902: Microevolution is distinct from macroevolution.
13.^ Myers 2006; NSTA 2007; IAP 2006; AAAS 2006; and Pinholster 2006; Ruling, Kitzmiller v.
Dover page 83
14.^ 29+ Evidences for Macroevolution: The Scientific Case for Common Descent , Douglas L.
Theobald, TalkOrigins Archive, Vers. 2.83, 2004, 12 Jan, 2004.
15.^ Laurence Moran (1993). "Evolution is a Fact and a Theory". TalkOrigins Archive.
http://www.talkorigins.org/faqs/evolution-fact.html. Retrieved 2008-02-07.

[edit] External links

• Introduction to macroevolution
• Macroevolution as the common descent of all life
 • Macroevolution in the 21st Century Macroevolution as an independent discipline.
• Macroevolution FAQ

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Macroevolution"

Categories: Evolutionary biology

• This page was last modified on 12 August 2010 at 17:43.

 w000

Mutation
From Wikipedia, the free encyclopedia

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For other uses, see Mutation (disambiguation).
Part of the Biology series on

Genetics
 Key components

Chromosome
DNA · RNA
Genome
Heredity
Mutation
Nucleotide
Variation

Glossary
Index
 Outline

History and topics

Introduction
History

Classical genetics
Evolution · Molecular
Mendelian inheritance
Molecular genetics

Research

DNA sequencing
Genetic engineering
Genomics · Topics
Medical genetics

Branches in genetics

Biology portal • v • d • e

Part of the Biology series on

 Evolution

Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

 Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
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Biology portal · v • d • e
 A mutation has caused this garden moss rose to produce flowers of different colors. This is a somatic
mutation that may also be passed on in the germ line.
Mutations are changes in a genomic sequence: the DNA sequence of a cell's genome or the DNA or
RNA sequence of a virus. Mutations are caused by radiation, viruses, transposons and mutagenic chemicals,
as well as errors that occur during meiosis or DNA replication.[1][2][3] They can also be induced by the
organism itself, by cellular processes such as hypermutation.
 Mutation can result in several different types of change in DNA sequences; these can either have no
effect, alter the product of a gene, or prevent the gene from functioning properly or completely. Studies in the
fly Drosophila melanogaster suggest that if a mutation changes a protein produced by a gene, this will
probably be harmful, with about 70 percent of these mutations having damaging effects, and the remainder
being either neutral or weakly beneficial.[4] Due to the damaging effects that mutations can have on cells,
organisms have evolved mechanisms such as DNA repair to remove mutations.[1] Therefore, the optimal
mutation rate for a species is a trade-off between costs of a high mutation rate, such as deleterious
mutations, and the metabolic costs of maintaining systems to reduce the mutation rate, such as DNA repair
enzymes.[5] Viruses that use RNA as their genetic material have rapid mutation rates,[6] which can be an
advantage since these viruses will evolve constantly and rapidly, and thus evade the defensive responses of
e.g. the human immune system.[7]
Contents
[hide]
• 1 Description
• 2 Causes
• 3 Classification of mutation types
• 3.1 By effect on structure
• 3.2 By effect on function
• 3.3 By effect on fitness
• 3.4 By impact on protein sequence
• 3.5 Special classes
• 3.6 Nomenclature
• 4 Harmful mutations
• 5 Beneficial mutations
• 6 Prion mutation
• 7 See also
• 8 References
• 9 External links
[edit] Description
Mutations can involve large sections of DNA becoming duplicated, usually through genetic
recombination.[8] These duplications are a major source of raw material for evolving new genes, with tens to
hundreds of genes duplicated in animal genomes every million years.[9] Most genes belong to larger families
of genes of shared ancestry.[10] Novel genes are produced by several methods, commonly through the
duplication and mutation of an ancestral gene, or by recombining parts of different genes to form new
combinations with new functions.[11][12] Here, domains act as modules, each with a particular and
independent function, that can be mixed together to produce genes encoding new proteins with novel
properties.[13] For example, the human eye uses four genes to make structures that sense light: three for
color vision and one for night vision; all four arose from a single ancestral gene.[14] Another advantage of
duplicating a gene (or even an entire genome) is that this increases redundancy; this allows one gene in the
pair to acquire a new function while the other copy performs the original function.[15][16] Other types of
mutation occasionally create new genes from previously noncoding DNA.[17][18]
Changes in chromosome number may involve even larger mutations, where segments of the DNA
within chromosomes break and then rearrange. For example, two chromosomes in the Homo genus fused to
produce human chromosome 2; this fusion did not occur in the lineage of the other apes, and they retain
these separate chromosomes.[19] In evolution, the most important role of such chromosomal
rearrangements may be to accelerate the divergence of a population into new species by making populations
less likely to interbreed, and thereby preserving genetic differences between these populations.[20]
Sequences of DNA that can move about the genome, such as transposons, make up a major fraction
of the genetic material of plants and animals, and may have been important in the evolution of genomes.[21]
For example, more than a million copies of the Alu sequence are present in the human genome, and these
sequences have now been recruited to perform functions such as regulating gene expression.[22] Another
effect of these mobile DNA sequences is that when they move within a genome, they can mutate or delete
existing genes and thereby produce genetic diversity.[2]
In multicellular organisms with dedicated reproductive cells, mutations can be subdivided into germ
line mutations, which can be passed on to descendants through their reproductive cells, and somatic
mutations (also called acquired mutations)[23]), which involve cells outside the dedicated reproductive group
and which are not usually transmitted to descendants. If the organism can reproduce asexually through
mechanisms such as cuttings or budding the distinction can become blurred.
For example, plants can sometimes transmit somatic mutations to their descendants asexually or
sexually where flower buds develop in somatically mutated parts of plants. A new mutation that was not
inherited from either parent is called a de novo mutation. The source of the mutation is unrelated to the
consequence[clarification needed], although the consequences are related to which cells were mutated.
Nonlethal mutations accumulate within the gene pool and increase the amount of genetic
variation[24]. The abundance of some genetic changes within the gene pool can be reduced by natural
selection, while other "more favorable" mutations may accumulate and result in adaptive evolutionary
changes.
For example, a butterfly may produce offspring with new mutations. The majority of these mutations
will have no effect; but one might change the color of one of the butterfly's offspring, making it harder (or
easier) for predators to see. If this color change is advantageous, the chance of this butterfly surviving and
producing its own offspring are a little better, and over time the number of butterflies with this mutation may
form a larger percentage of the population.
Neutral mutations are defined as mutations whose effects do not influence the fitness of an
individual. These can accumulate over time due to genetic drift. It is believed that the overwhelming majority
of mutations have no significant effect on an organism's fitness. Also, DNA repair mechanisms are able to
mend most changes before they become permanent mutations, and many organisms have mechanisms for
eliminating otherwise permanently mutated somatic cells.
Mutation is generally accepted by biologists as the mechanism by which natural selection acts,
generating advantageous new traits that survive and multiply in offspring as well as disadvantageous traits, in
less fit offspring, that tend to die out.

[edit] Causes
Two classes of mutations are spontaneous mutations (molecular decay) and induced mutations
caused by mutagens.
Spontaneous mutations on the molecular level can be caused by:
• Tautomerism – A base is changed by the repositioning of a hydrogen atom, altering the
hydrogen bonding pattern of that base resulting in incorrect base pairing during replication.
• Depurination – Loss of a purine base (A or G) to form an apurinic site (AP site).
• Deamination – Hydrolysis changes a normal base to an atypical base containing a keto group
in place of the original amine group. Examples include C → U and A → HX (hypoxanthine), which can
be corrected by DNA repair mechanisms; and 5MeC (5-methylcytosine) → T, which is less likely to be
detected as a mutation because thymine is a normal DNA base.
• Slipped strand mispairing - Denaturation of the new strand from the template during
replication, followed by renaturation in a different spot ("slipping"). This can lead to insertions or
deletions.
A covalent adduct between benzo[a]pyrene, the major mutagen in tobacco smoke, and DNA[25]
Induced mutations on the molecular level can be caused by:
• Chemicals
• Hydroxylamine NH2OH
• Base analogs (e.g. BrdU)
• Alkylating agents (e.g. N-ethyl-N-nitrosourea) These agents can mutate both
replicating and non-replicating DNA. In contrast, a base analog can only mutate the DNA
when the analog is incorporated in replicating the DNA. Each of these classes of chemical
mutagens has certain effects that then lead to transitions, transversions, or deletions.
• Agents that form DNA adducts (e.g. ochratoxin A metabolites)[26]
• DNA intercalating agents (e.g. ethidium bromide)
• DNA crosslinkers
• Oxidative damage
• Nitrous acid converts amine groups on A and C to diazo groups, altering their
hydrogen bonding patterns which leads to incorrect base pairing during replication.
• Radiation
• Ultraviolet radiation (nonionizing radiation). Two nucleotide bases in DNA – cytosine
and thymine – are most vulnerable to radiation that can change their properties. UV light can
induce adjacent pyrimidine bases in a DNA strand to become covalently joined as a
pyrimidine dimer. UV radiation, particularly longer-wave UVA, can also cause oxidative
damage to DNA[27].
• Ionizing radiation
 • Viral infections[28]
DNA has so-called hotspots, where mutations occur up to 100 times more frequently than the normal
mutation rate. A hotspot can be at an unusual base, e.g., 5-methylcytosine.
Mutation rates also vary across species. Evolutionary biologists have theorized that higher mutation
rates are beneficial in some situations, because they allow organisms to evolve and therefore adapt more
quickly to their environments. For example, repeated exposure of bacteria to antibiotics, and selection of
resistant mutants, can result in the selection of bacteria that have a much higher mutation rate than the
original population (mutator strains).
[edit] Classification of mutation types
 Illustrations of five types of chromosomal mutations.

Selection of disease-causing mutations, in a standard table of the genetic code of amino acids.[29]

[edit] By effect on structure

The sequence of a gene can be altered in a number of ways. Gene mutations have varying effects on
health depending on where they occur and whether they alter the function of essential proteins. Mutations in
the structure of genes can be classified as:
• Small-scale mutations, such as those affecting a small gene in one or a few nucleotides,
including:
 • Point mutations, often caused by chemicals or malfunction of DNA replication,
exchange a single nucleotide for another[30]. These changes are classified as transitions or
transversions[31]. Most common is the transition that exchanges a purine for a purine (A ↔
G) or a pyrimidine for a pyrimidine, (C ↔ T). A transition can be caused by nitrous acid, base
mis-pairing, or mutagenic base analogs such as 5-bromo-2-deoxyuridine (BrdU). Less
common is a transversion, which exchanges a purine for a pyrimidine or a pyrimidine for a
purine (C/T ↔ A/G). An example of a transversion is adenine (A) being converted into a
cytosine (C). A point mutation can be reversed by another point mutation, in which the
nucleotide is changed back to its original state (true reversion) or by second-site reversion (a
complementary mutation elsewhere that results in regained gene functionality). Point
mutations that occur within the protein coding region of a gene may be classified into three
kinds, depending upon what the erroneous codon codes for:
• Silent mutations: which code for the same amino acid.
• Missense mutations: which code for a different amino acid.
• Nonsense mutations: which code for a stop and can truncate the protein.
• Insertions add one or more extra nucleotides into the DNA. They are usually caused
by transposable elements, or errors during replication of repeating elements (e.g. AT
repeats[citation needed]). Insertions in the coding region of a gene may alter splicing of the
mRNA (splice site mutation), or cause a shift in the reading frame (frameshift), both of which
can significantly alter the gene product. Insertions can be reverted by excision of the
transposable element.
• Deletions remove one or more nucleotides from the DNA. Like insertions, these
mutations can alter the reading frame of the gene. They are generally irreversible: though
exactly the same sequence might theoretically be restored by an insertion, transposable
 elements able to revert a very short deletion (say 1–2 bases) in any location are either highly
unlikely to exist or do not exist at all. Note that a deletion is not the exact opposite of an
insertion: the former is quite random while the latter consists of a specific sequence inserting
at locations that are not entirely random or even quite narrowly defined.
• Large-scale mutations in chromosomal structure, including:
• Amplifications (or gene duplications) leading to multiple copies of all chromosomal
regions, increasing the dosage of the genes located within them.
• Deletions of large chromosomal regions, leading to loss of the genes within those
regions.
• Mutations whose effect is to juxtapose previously separate pieces of DNA, potentially
bringing together separate genes to form functionally distinct fusion genes (e.g. bcr-abl).
These include:
• Chromosomal translocations: interchange of genetic parts from
nonhomologous chromosomes.
• Interstitial deletions: an intra-chromosomal deletion that removes a segment
of DNA from a single chromosome, thereby apposing previously distant genes. For
example, cells isolated from a human astrocytoma, a type of brain tumor, were found
to have a chromosomal deletion removing sequences between the "fused in
glioblastoma" (fig) gene and the receptor tyrosine kinase "ros", producing a fusion
protein (FIG-ROS). The abnormal FIG-ROS fusion protein has constitutively active
kinase activity that causes oncogenic transformation (a transformation from normal
cells to cancer cells).
• Chromosomal inversions: reversing the orientation of a chromosomal
segment.
 • Loss of heterozygosity: loss of one allele, either by a deletion or recombination
event, in an organism that previously had two different alleles.

[edit] By effect on function

• Loss-of-function mutations are the result of gene product having less or no function. When
the allele has a complete loss of function (null allele) it is often called an amorphic mutation.
Phenotypes associated with such mutations are most often recessive. Exceptions are when the
organism is haploid, or when the reduced dosage of a normal gene product is not enough for a
normal phenotype (this is called haploinsufficiency).
• Gain-of-function mutations change the gene product such that it gains a new and abnormal
function. These mutations usually have dominant phenotypes. Often called a neomorphic mutation.
• Dominant negative mutations (also called antimorphic mutations) have an altered gene
product that acts antagonistically to the wild-type allele. These mutations usually result in an altered
molecular function (often inactive) and are characterised by a dominant or semi-dominant phenotype.
In humans, Marfan syndrome is an example of a dominant negative mutation occurring in an
autosomal dominant disease. In this condition, the defective glycoprotein product of the fibrillin gene
(FBN1) antagonizes the product of the normal allele.
• Lethal mutations are mutations that lead to the death of the organisms which carry the
mutations.
• A back mutation or reversion is a point mutation that restores the original sequence and
hence the original phenotype.[32]
 [edit] By effect on fitness
In applied genetics it is usual to speak of mutations as either harmful or beneficial.
• A harmful mutation is a mutation that decreases the fitness of the organism.
• A beneficial mutation is a mutation that increases fitness of the organism, or which promotes
traits that are desirable.
In theoretical population genetics, it is more usual to speak of such mutations as deleterious or
advantageous. In the neutral theory of molecular evolution, genetic drift is the basis for most variation at the
molecular level.
• A neutral mutation has no harmful or beneficial effect on the organism. Such mutations occur
at a steady rate, forming the basis for the molecular clock.
• A deleterious mutation has a negative effect on the phenotype, and thus decreases the
fitness of the organism.
• An advantageous mutation has a positive effect on the phenotype, and thus increases the
fitness of the organism.
• A nearly neutral mutation is a mutation that may be slightly deleterious or advantageous,
although most nearly neutral mutations are slightly deleterious.
By inheritance
• inheritable generic in pro-generic tissue or cells on path to be changed to gametes.
• non inheritable somatic (e.g., carcinogenic mutation)
• non inheritable post mortem aDNA mutation in decaying remains.
 By pattern of inheritance The human genome contains two copies of each gene – a paternal and a
maternal allele.
• A heterozygous mutation is a mutation of only one allele.
• A homozygous mutation is an identical mutation of both the paternal and maternal alleles.
• Compound heterozygous mutations or a genetic compound comprises two different
mutations in the paternal and maternal alleles.[33]
• A wildtype or homozygous non-mutated organism is one in which neither allele is mutated.
(Just not a mutation)

[edit] By impact on protein sequence

• A frameshift mutation is a mutation caused by insertion or deletion of a number of
nucleotides that is not evenly divisible by three from a DNA sequence. Due to the triplet nature of
gene expression by codons, the insertion or deletion can disrupt the reading frame, or the grouping of
the codons, resulting in a completely different translation from the original. The earlier in the
sequence the deletion or insertion occurs, the more altered the protein produced is.
• A nonsense mutation is a point mutation in a sequence of DNA that results in a premature
stop codon, or a nonsense codon in the transcribed mRNA, and possibly a truncated, and often
nonfunctional protein product.
• Missense mutations or nonsynonymous mutations are types of point mutations where a
single nucleotide is changed to cause substitution of a different amino acid. This in turn can render
the resulting protein nonfunctional. Such mutations are responsible for diseases such as
Epidermolysis bullosa, sickle-cell disease, and SOD1 mediated ALS (Boillée 2006, p. 39).
• A neutral mutation is a mutation that occurs in an amino acid codon which results in the use
of a different, but chemically similar, amino acid. The similarity between the two is enough that little
or no change is often rendered in the protein. For example, a change from AAA to AGA will encode
lysine, a chemically similar molecule to the intended arginine.
• Silent mutations are mutations that do not result in a change to the amino acid sequence of a
protein. They may occur in a region that does not code for a protein, or they may occur within a
codon in a manner that does not alter the final amino acid sequence. The phrase silent mutation is
often used interchangeably with the phrase synonymous mutation; however, synonymous mutations
are a subcategory of the former, occurring only within exons. The name silent could be a misnomer.
For example, a silent mutation in the exon/intron border may lead to alternative splicing by changing
the splice site (see Splice site mutation), thereby leading to a changed protein.

[edit] Special classes

• Conditional mutation is a mutation that has wild-type (or less severe) phenotype under
certain "permissive" environmental conditions and a mutant phenotype under certain "restrictive"
conditions. For example, a temperature-sensitive mutation can cause cell death at high temperature
(restrictive condition), but might have no deleterious consequences at a lower temperature
(permissive condition).
[edit] Nomenclature
Nomenclature of mutations specify the type of mutation and base or amino acid changes.
• Nucleotide substitution (e.g. 76A>T) - The number is the position of the nucleotide from the 5'
end, the first letter represents the wild type nucleotide, and the second letter represents the
nucleotide which replaced the wild type. In the given example, the adenine at the 76th position was
replaced by a thymine.
• If it becomes necessary to differentiate between mutations in genomic DNA,
mitochondrial DNA, and RNA, a simple convention is used. For example, if the 100th base of
a nucleotide sequence mutated from G to C, then it would be written as g.100G>C if the
mutation occurred in genomic DNA, m.100G>C if the mutation occurred in mitochondrial
DNA, or r.100g>c if the mutation occurred in RNA. Note that for mutations in RNA, the
nucleotide code is written in lower case.
• Amino acid substitution (e.g. D111E) – The first letter is the one letter code of the wild type
amino acid, the number is the position of the amino acid from the N terminus, and the second letter is
the one letter code of the amino acid present in the mutation. Nonsense mutations are represented
with an X for the second amino acid (e.g. D111X).
• Amino acid deletion (e.g. ΔF508) – The Greek letter Δ (delta) indicates a deletion. The letter
refers to the amino acid present in the wild type and the number is the position from the N terminus of
the amino acid were it to be present as in the wild type.
 [edit] Harmful mutations
Changes in DNA caused by mutation can cause errors in protein sequence, creating partially or
completely non-functional proteins. To function correctly, each cell depends on thousands of proteins to
function in the right places at the right times. When a mutation alters a protein that plays a critical role in the
body, a medical condition can result. A condition caused by mutations in one or more genes is called a
genetic disorder. Some mutations alter a gene's DNA base sequence but do not change the function of the
protein made by the gene. Studies of the fly Drosophila melanogaster suggest that if a mutation does change
a protein, this will probably be harmful, with about 70 percent of these mutations having damaging effects,
and the remainder being either neutral or weakly beneficial.[34] However, studies in yeast have shown that
only 7% of mutations that are not in genes are harmful.[35]
If a mutation is present in a germ cell, it can give rise to offspring that carries the mutation in all of its
cells. This is the case in hereditary diseases. On the other hand, a mutation may occur in a somatic cell of an
organism. Such mutations will be present in all descendants of this cell within the same organism, and
certain mutations can cause the cell to become malignant, and thus cause cancer[36].
Often, gene mutations that could cause a genetic disorder are repaired by the DNA repair system of
the cell. Each cell has a number of pathways through which enzymes recognize and repair mistakes in DNA.
Because DNA can be damaged or mutated in many ways, the process of DNA repair is an important way in
which the body protects itself from disease.

[edit] Beneficial mutations

Although most mutations that change protein sequences are neutral or harmful, some mutations
have a positive effect on an organism. In this case, the mutation may enable the mutant organism to
withstand particular environmental stresses better than wild-type organisms, or reproduce more quickly. In
these cases a mutation will tend to become more common in a population through natural selection.
For example, a specific 32 base pair deletion in human CCR5 (CCR5-Δ32) confers HIV resistance to
homozygotes and delays AIDS onset in heterozygotes.[37] The CCR5 mutation is more common in those of
European descent. One possible explanation of the etiology of the relatively high frequency of CCR5-Δ32 in
the European population is that it conferred resistance to the bubonic plague in mid-14th century Europe.
People with this mutation were more likely to survive infection; thus its frequency in the population increased.
[38] This theory could explain why this mutation is not found in southern Africa, where the bubonic plague
never reached. A newer theory suggests that the selective pressure on the CCR5 Delta 32 mutation was
caused by smallpox instead of the bubonic plague.[39]
Another example, is Sickle cell disease which is a blood disorder in which the body produces an
abnormal type of the oxygen-carrying substance hemoglobin in the red blood cells. One-third of all
indigenous inhabitants of Sub-Saharan Africa carry the gene[40], because in areas where malaria is
common, there is a survival value in carrying only a single sickle-cell gene (sickle cell trait).[41] Those with
only one of the two alleles of the sickle-cell disease are more resistant to malaria, since the infestation of the
malaria plasmodium is halted by the sickling of the cells which it infests.

[edit] Prion mutation

Prions are proteins and do not contain genetic material. However, prion replication has been shown
to be subject to mutation and natural selection just like other forms of replication.[42]
[edit] See also
• Aneuploidy
• Antioxidant
• Budgerigar colour genetics
• Homeobox
• Macromutation
• Muller's morphs
• Mutant
• Mutagenesis
• Polyploidy
• Robertsonian translocation
• Signature tagged mutagenesis
• Site-directed mutagenesis
• TILLING (molecular biology)

[edit] References
1. ^ a b Bertram J (2000). "The molecular biology of cancer". Mol. Aspects Med. 21 (6): 167–223.
doi:10.1016/S0098-2997(00)00007-8. PMID 11173079.
2. ^ a b Aminetzach YT, Macpherson JM, Petrov DA (2005). "Pesticide resistance via
transposition-mediated adaptive gene truncation in Drosophila". Science 309 (5735): 764–7.
doi:10.1126/science.1112699. PMID 16051794.
 3. ^ Burrus V, Waldor M (2004). "Shaping bacterial genomes with integrative and conjugative
elements". Res. Microbiol. 155 (5): 376–86. doi:10.1016/j.resmic.2004.01.012. PMID 15207870.
4. ^ Sawyer SA, Parsch J, Zhang Z, Hartl DL (2007). "Prevalence of positive selection among
nearly neutral amino acid replacements in Drosophila". Proc. Natl. Acad. Sci. U.S.A. 104 (16): 6504–
10. doi:10.1073/pnas.0701572104. PMID 17409186.
5. ^ Sniegowski P, Gerrish P, Johnson T, Shaver A (2000). "The evolution of mutation rates:
separating causes from consequences". Bioessays 22 (12): 1057–66. doi:10.1002/1521-
1878(200012)22:12<1057::AID-BIES3>3.0.CO;2-W. PMID 11084621.
6. ^ Drake JW, Holland JJ (1999). "Mutation rates among RNA viruses". Proc. Natl. Acad. Sci.
U.S.A. 96 (24): 13910–3. doi:10.1073/pnas.96.24.13910. PMID 10570172. PMC 24164.
http://www.pnas.org/content/96/24/13910.long.
7. ^ Holland J, Spindler K, Horodyski F, Grabau E, Nichol S, VandePol S (1982). "Rapid
evolution of RNA genomes". Science 215 (4540): 1577–85. doi:10.1126/science.7041255.
PMID 7041255.
8. ^ Hastings, P J; Lupski, JR; Rosenberg, SM; Ira, G (2009). "Mechanisms of change in gene
copy number". Nature Reviews. Genetics 10 (8): 551–564. doi:10.1038/nrg2593. PMID 19597530.
9. ^ Carroll SB, Grenier J, Weatherbee SD (2005). From DNA to Diversity: Molecular Genetics
and the Evolution of Animal Design. Second Edition . Oxford: Blackwell Publishing. ISBN 1-4051-
1950-0.
10.^ Harrison P, Gerstein M (2002). "Studying genomes through the aeons: protein families,
pseudogenes and proteome evolution". J Mol Biol 318 (5): 1155–74. doi:10.1016/S0022-
2836(02)00109-2. PMID 12083509.
 11.^ Orengo CA, Thornton JM (2005). "Protein families and their evolution-a structural
perspective". Annu. Rev. Biochem. 74: 867–900. doi:10.1146/annurev.biochem.74.082803.133029.
PMID 15954844.
12.^ Long M, Betrán E, Thornton K, Wang W (November 2003). "The origin of new genes:
glimpses from the young and old". Nat. Rev. Genet. 4 (11): 865–75. doi:10.1038/nrg1204.
PMID 14634634.
13.^ Wang M, Caetano-Anollés G (2009). "The evolutionary mechanics of domain organization
in proteomes and the rise of modularity in the protein world". Structure 17 (1): 66–78.
doi:10.1016/j.str.2008.11.008. PMID 19141283.
14.^ Bowmaker JK (1998). "Evolution of colour vision in vertebrates". Eye (London, England) 12
(Pt 3b): 541–7. PMID 9775215.
15.^ Gregory TR, Hebert PD (1999). "The modulation of DNA content: proximate causes and
ultimate consequences". Genome Res. 9 (4): 317–24. doi:10.1101/gr.9.4.317 (inactive 2009-11-14).
PMID 10207154. http://genome.cshlp.org/content/9/4/317.full.
16.^ Hurles M (July 2004). "Gene duplication: the genomic trade in spare parts". PLoS Biol. 2
(7): E206. doi:10.1371/journal.pbio.0020206. PMID 15252449.
17.^ Liu N, Okamura K, Tyler DM (2008). "The evolution and functional diversification of animal
microRNA genes". Cell Res. 18 (10): 985–96. doi:10.1038/cr.2008.278. PMID 18711447.
PMC 2712117. http://www.nature.com/cr/journal/v18/n10/full/cr2008278a.html.
18.^ Siepel A (October 2009). "Darwinian alchemy: Human genes from noncoding DNA".
Genome Res. 19 (10): 1693–5. doi:10.1101/gr.098376.109. PMID 19797681. PMC 2765273.
http://genome.cshlp.org/content/19/10/1693.full.
19.^ Zhang J, Wang X, Podlaha O (2004). "Testing the chromosomal speciation hypothesis for
humans and chimpanzees". Genome Res. 14 (5): 845–51. doi:10.1101/gr.1891104. PMID 15123584.
 20.^ Ayala FJ, Coluzzi M (2005). "Chromosome speciation: humans, Drosophila, and
mosquitoes". Proc. Natl. Acad. Sci. U.S.A. 102 (Suppl 1): 6535–42. doi:10.1073/pnas.0501847102.
PMID 15851677. PMC 1131864. http://www.pnas.org/content/102/suppl.1/6535.full.
21.^ Hurst GD, Werren JH (2001). "The role of selfish genetic elements in eukaryotic evolution".
Nat. Rev. Genet. 2 (8): 597–606. doi:10.1038/35084545. PMID 11483984.
22.^ Häsler J, Strub K (2006). "Alu elements as regulators of gene expression". Nucleic Acids
Res. 34 (19): 5491–7. doi:10.1093/nar/gkl706. PMID 17020921.
23.^ "Genome Dictionary". http://www.theodora.com/genetics/#somaticcellgeneticmutation.
Retrieved 2010-06-06. .
24.^ Eyre-Walker, A.; Keightley, P. (Aug 2007). "The distribution of fitness effects of new
mutations". Nature reviews. Genetics 8 (8): 610–618. doi:10.1038/nrg2146. ISSN 1471-0056.
PMID 17637733. edit
25.^ Created from PDB 1JDG
26.^ Pfohl-Leszkowicz A, Manderville RA (January 2007). "Ochratoxin A: An overview on toxicity
and carcinogenicity in animals and humans". Mol Nutr Food Res 51 (1): 61–99.
doi:10.1002/mnfr.200600137. PMID 17195275.
27.^ Kozmin S, Slezak G, Reynaud-Angelin A, Elie C, de Rycke Y, Boiteux S, Sage E
(September 2005). "UVA radiation is highly mutagenic in cells that are unable to repair 7,8-dihydro-8-
oxoguanine in Saccharomyces cerevisiae". Proc. Natl. Acad. Sci. U.S.A. 102 (38): 13538–43.
doi:10.1073/pnas.0504497102. PMID 16157879. PMC 1224634.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=16157879.
28.^ Pilon L, Langelier Y, Royal A (1 August 1986). "Herpes simplex virus type 2 mutagenesis:
characterization of mutants induced at the hprt locus of nonpermissive XC cells". Mol. Cell. Biol. 6
(8): 2977–83. PMID 3023954. PMC 367868. http://mcb.asm.org/cgi/pmidlookup?
view=long&pmid=3023954.
29.^ References for the image are found in Wikimedia Commons page at:
Commons:File:Notable mutations.svg#References.
30.^ Freese, Ernst (April 1959). "The Difference between Spontaneous and Base-Analogue
Induced Mutations of Phage T4". Proc. Natl. Acad. Sci. U.S.A. 45 (4): 622–33.
doi:10.1073/pnas.45.4.622. PMID 16590424.
31.^ Freese, Ernst (1959). "The Specific Mutagenic Effect of Base Analogues on Phage T4". J.
Mol. Biol. 1: 87–105. doi:10.1016/S0022-2836(59)80038-3.
32.^ Ellis NA, Ciocci S, German J (2001). "Back mutation can produce phenotype reversion in
Bloom syndrome somatic cells". Hum Genet 108 (2): 167–73. doi:10.1007/s004390000447.
PMID 11281456. http://link.springer.de/link/service/journals/00439/bibs/1108002/11080167.htm.
33.^ Medterms.com
34.^ Sawyer SA, Parsch J, Zhang Z, Hartl DL (2007). "Prevalence of positive selection among
nearly neutral amino acid replacements in Drosophila". Proc. Natl. Acad. Sci. U.S.A. 104 (16): 6504–
10. doi:10.1073/pnas.0701572104. PMID 17409186.
35.^ Doniger SW, Kim HS, Swain D, et al. (August 2008). "A catalog of neutral and deleterious
polymorphism in yeast". PLoS Genet. 4 (8): e1000183. doi:10.1371/journal.pgen.1000183.
PMID 18769710.
36.^ Ionov Y, Peinado MA, Malkhosyan S, Shibata D, Perucho M (1993). "Ubiquitous somatic
mutations in simple repeated sequences reveal a new mechanism for colonic carcinogenesis".
Nature 363 (6429): 558–61. doi:10.1038/363558a0. PMID 8505985.
37.^ Sullivan, Amy D. et al. (2001). "The coreceptor mutation CCR5Δ32 influences the dynamics
of HIV epidemics and is selected for by HIV". PNAS 95 (18): 10214–10219.
 doi:10.1073/pnas.181325198. PMID 11517319. PMC 56941.
http://www.pnas.org/content/98/18/10214.full.
38.^ "PBS:Secrets of the Dead. Case File: Mystery of the Black Death".
http://www.pbs.org/wnet/secrets/previous_seasons/case_plague/clues.html.
39.^ Galvani A, Slatkin M (2003). "Evaluating plague and smallpox as historical selective
pressures for the CCR5-Δ32 HIV-resistance allele". Proc Natl Acad Sci USA 100 (25): 15276–9.
doi:10.1073/pnas.2435085100. PMID 14645720.
40.^ Sicklecell.md
41.^ Sicklecell.md FAQ: "Why is Sickle Cell Anaemia only found in Black people?
42.^ 'Lifeless' prion proteins are 'capable of evolution'

[edit] External links

• "All About Mutations" from the Huntington's Disease Outreach Project for Education at
Stanford
• Central Locus Specific Variation Database at the Institute of Genomics and Integrative
Biology
• The mutations chapter of the WikiBooks General Biology textbook
• Examples of Beneficial Mutations
• Correcting mutation by gene therapy
• BBC Radio 4 In Our Time - GENETIC MUTATION - with Steve Jones - streaming audio
 [show]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

[show]
v•d•e
Mutation

Point Nonsense mutation · Missense mutation · Frameshift mutation · Neutral

mutation mutation · Silent mutation · substitution (Transversion, Transition)
Other Insertion · Deletion

Retrieved from "http://en.wikipedia.org/wiki/Mutation"

Categories: Mutation | Evolutionary biology | Genetics | Radiation health effects

• This page was last modified on 15 August 2010 at 18:23.

 W000

Mutation (génétique)
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir mutation.

Les mécanismes de
l'évolution biologique

Mécanismes non aléatoires :

• sélection naturelle
 • sélection
utilitaire
• sélection
sexuelle
• sélection de
parentèle
• sélection de
groupe
• sélection
stratégique
• sélection artificielle

Mécanismes aléatoires :
• mutation génétique
• recombinaison
• dérive génétique

Conséquences de l'évolution :
• spéciation
• adaptation des
espèces
• radiation évolutive
 Exemple de mutation génétique.
En génétique, une mutation est une modification irréversible de l'information génétique et héréditaire
contenue dans un génome.
Sommaire
[masquer]
• 1
Types de
mutation
• 2
Origines ou
causes de
mutations
• 3
Utilité dans
l'évolution
• 4
Agents
mutagènes
• 5
Transmission
des mutations
• 6
Conséquence
d'une mutation
• 7
Types de mutation [modifier]

Un exemple de mutation homéotique: une drosophile Antennapedia

 Chez la souris, comme chez l'Homme, les mutations germinales augmentent avec l'âge, surtout pour
les mutations aneuploïdes, XX8 et YY8 et après l'âge de deux ans (Ces données ne concernent que les cas
où les cellules (spermatozoïdes) ont acquis un chromosome supplémentaire [1]
On peut distinguer plusieurs types de mutations
• On parle de mutation germinale ou mutation de novo, quand la mutation porte sur l'ADN des
cellules souches d'un gamète . Dans ce cas, l'embryon sera porteur de la mutation, alors qu'aucun
des parents ne la possédait dans son patrimoine génétique. Ce type de mutation survient lors de la
formation ou de la vie des gamètes d'un des deux parent (ovule ou spermatozoïde).
Dans ce cas les mutations apportées par le spermatozoïde prédominent très largement ; Environ
80% des aberrations chromosomiques des chromosomes des bébés proviennent du matériel
chromosomique apporté par le spermatozoïde[2]. Ceci est pour une faible part du au fait que le
spermatozoïde possède un chromosome de plus (le X), et surtout au fait qu'il est bien plus vulnérable
 aux mutations que l'ovule, ceci d'autant plus que le père (ou plus précisément le donneur) est âgé.
Le fait que la diminution avec l'âge de la qualité génétique du sperme soit également constaté chez
la souris [2] laisse penser que l'horloge biologique pourrait être en cause, et pas seulement une
accumulation d'erreurs corrélable au temps absolu.
• Les mutations somatiques touchent des cellules particulières.
- Les mutations post-zygotiques sont les mutations qui apparaissent dans l'œuf après sa
fécondation. Elles sont plus rares.
-des mutations peuvent apparaître tout au long de la vie sur l'ADN de n'importe quelle cellule,
alors transmises à la lignée des cellules filles.
Chez les animaux pluricellulaires, les mutations de la lignée germinale peuvent être transmises à la
descendance, contrairement aux mutations somatiques.
Une mutation est dite sexuelle lorsqu'elle concerne un chromosome sexuel, par exemple X/Y chez
les mammifères ou W/Z chez les oiseaux. Une mutation est dite autosomique losqu'elle touche un autre
gène que les gènes de détermination sexuelle.
Les mutations peuvent être classées selon leurs modalités de modification du gène :
• MUTATIONS PONCTUELLES (Ne modifient qu'un nucléotide de la séquence d'ADN)
• Mutations par substitution :
• les mutations faux-sens. Cette mutation ponctuelle se traduit par le
changement d'un nucléotide par un autre. Dans certains cas, cette modification de
nucléotide entraîne une modification de l'acide aminé codé. Le changement d'un
acide aminé peut avoir ou non une répercussion en termes de fonction de la protéine
 produite par le gène, dans le cas d'un gène codant, ou d'une modification d'affinité
pour un facteur de transcription, dans le cas d'une zone promotrice de l'ADN. On
parle de mutation de transition (transition mutation) lorsqu’il y a substitution d’une
base purique par une autre base purique (ou d’une base pyrimidique par une autre
base pyrimidique). Au contraire une mutation de transversion (transversion mutation)
est une mutation causée par la substitution d’une base purique par une base
pyrimidique (ou d’une base pyrimidique par une base purique) ;
• les mutations non-sens. Le changement d'un nucléotide provoque le
remplacement d'un codon spécifiant un acide aminé par un codon stop. Cela
entraîne la production d'une protéine tronquée ;
• les mutations silencieuses. Ce sont des mutations qui ne modifient pas la
séquence d'une protéine, à cause de la redondance du code génétique (le nouveau
triplet code le même acide aminé que le triplet original), ou parce qu'elle touche une
région non codante de l'ADN, ou un intron. Cette mutation n'a aucune conséquence
sur le phénotype. Une mutation synonyme désigne une mutation silencieuse qui
touche un exon, sans changer la séquence de la protéine.
• Insertions et délétions :
• les mutations décalantes. Une addition ou une suppression de nucléotides
non multiple de 3 provoquera un changement de cadre de lecture. Au moment de la
traduction, cela générera le plus souvent une protéine tronquée par l'apparition d'un
codon-stop prématuré.
• MUTATIONS DYNAMIQUES
 Ces mutations évoluent d'une génération à l'autre, elles correspondent à des répétitions importantes
de certains triplets au niveau de l'ADN (CAG et GGG).

Origines ou causes de mutations [modifier]

Les mutations sont des « erreurs de copie » du matériel génétique. Les grandes causes connues de
mutations sont ;
• les erreurs faites lors de la préparation à la division cellulaire,
• les erreurs ou formes nouvelles dues à l'exposition à des agents mutagènes (radiations,
agents chimiques, virus).
Une très grande partie des erreurs commises au cours de la réplication du génome sont corrigées
immédiatement par des mécanismes complexes et efficaces de réparation de l'ADN, et seule une faible part
de ces erreurs deviennent des mutations transmises aux cellules-filles.

Utilité dans l'évolution [modifier]

Les mutations expliquent l'existence d'une variabilité entre les gènes.
Les mutations qui sont le moins favorables (délétères) à la survie de l'individu qui les porte, sont éliminées
par le jeu de la sélection naturelle, alors que les mutations avantageuses, beaucoup plus rares, tendent à
s'accumuler.
La plupart des mutations sont dites neutres, elles n'influencent pas la valeur sélective et peuvent se fixer ou
disparaître par le jeu de la dérive génétique.
Les mutations spontanées, généralement rares et aléatoires, constituent donc la principale source de
diversité génétique, moteur de l'évolution. Les causes des mutations spontanées sont inconnues.
Agents mutagènes [modifier]
La mutation est un phénomène spontané, dû à des erreurs dans le processus de réplication de
l'ADN. Mais, dans certaines circonstances, le taux de mutations peut être augmenté considérablement par
des facteurs physiques ou chimiques, appelés agents mutagènes ;
• Les ondes électromagnétiques (rayons X, rayons gamma, les rayons ultraviolets).
• Des substances chimiques qui interagissent avec l’ADN (ou éventuellement avec l'ARN) tels
que pesticides, dérivés de benzène, solvants, etc.
• Des substances chimiques qui interagissent avec des éléments impliqués dans la réplication
de l'ADN, comme la colchicine qui empêche la formation du fuseau achromatique d'où une altération
du nombre de chromosomes.
• une modification du système de réparation de l'ADN, qui cesse alors de corriger les erreurs
de réplications.

Transmission des mutations [modifier]

• Si une mutation affecte les cellules germinales, elle est transmise aux descendants de
l'individu mutant. Dans certains cas, cette mutation peut procurer un avantage sélectif ou au
contraire être délétère, voire létale. C'est la base du processus de l'évolution. Il est cependant admis
que la plupart des mutations interviennent entre les gènes, dans les introns, ou à des endroits où leur
effet est minime (mutations synonymes) ; la plupart des mutations sont donc probablement neutres,
et ne sont conservées (ou éliminées) que par hasard (dérive génétique).
• En revanche, comme c'est le cas pour la plupart des mutations accidentelles (provoquées
par irradiation ou substances chimiques), si elle affecte les cellules somatiques, la mutation ne se
transmet pas et n'affectera donc que le sujet l'ayant subie directement. Si les cellules se divisent
activement, il y a possibilité de création d'une tumeur pouvant évoluer en cancer. À l'opposé, s'il n'y a
pas de division l'effet est négligeable.

Conséquence d'une mutation [modifier]

Un exemple de mutation génétique : le leucistisme du lion blanc.

Les mutations peuvent être classées selon leurs conséquences phénotypiques :
• la plupart des mutations ont de plus ou moins importantes conséquences phénotypiques
(certaines d'entre elles peuvent avoir des conséquences graves comme le cancer ou des maladies
 génétiques, car la modification d'un seul acide aminé dans la chaîne constituant une protéine peut
modifier complètement sa structure spatiale, qui conditionne son fonctionnement) ;
• les mutations neutres ne modifient pas le fonctionnement de la protéine et n'ont pas de
conséquence phénotypique macroscopique ;
• Les mutations conditionnelles", ne s'expriment que dans certaines conditions particulières
(élévation de la température, niveau d'hydratation etc)
• les mutations silencieuses ou muettes n'entraînent aucun changement dans la séquence
d'acides aminés, ce qui est dû aux nombreuses redondances dans le code génétique. En effet, la
troisième base d'un codon n'est en général pas codante (de fait, plusieurs codons différents codent
le même acide aminé). Cette propriété est appelée redondance (ou dégénérescence) du codage.

Notes et références [modifier]

1. ↑ LLNL d'après X. Lowe, et al., "Aneuploidies and Micronuclei in the Germ Cells of Male
Mice of Advanced Age", Mutation Research ; 1995) D'après X. Lowe, et al., "Aneuploidies and
Micronuclei in the Germ Cells of Male Mice of Advanced Age", Mutation Research ; 1995
2. ↑ a et b The Genetic Contribution of Sperm:Healthy Baby or Not? [archive] ; Science &
Technology Review November/December 1995, LLNL, USA

Voir aussi [modifier]

Sur les autres projets Wikimédia :
• Mutation (génétique) sur Wikimedia Commons (ressources multimédia)
• Mutation (génétique) sur le Wiktionnaire (dictionnaire universel)
Articles connexes [modifier]
• Horloge moléculaire
• Mutation récessive
• Mutation dans la fiction,

Liens externes [modifier]

• (en) Moroccan Human Mutation Database
• (fr) Cours de génétique sur le site de l'Université Paris-Sud 11

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Natural selection
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For other uses, see Natural Selection (disambiguation).
Part of the Biology series on

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Biology portal · v • d • e
Natural selection is a natural law by which genetically heritable traits become more or less common
in a population over successive generations. This selection in interaction with the production of variation, the
possible genetic fixation process and possibly, in several cases, with little epigenetic process determine the
evolution of the species.
 The natural genetic variation within a population of organisms means that some individuals will
survive and reproduce more successfully than others in their current environment. For example, the
peppered moth exists in both light and dark colors in the United Kingdom, but during the industrial revolution
many of the trees on which the moths rested became blackened by soot, giving the dark-colored moths an
advantage in hiding from predators. This gave dark-colored moths a better chance of surviving to produce
dark-colored offspring, and in just a few generations the majority of the moths were dark. Factors which affect
reproductive success are also important, an issue which Charles Darwin developed in his ideas on sexual
selection.
Natural selection acts on the phenotype, or the observable characteristics of an organism, but the
genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common
in a population (see allele frequency). Over time, this process can result in adaptations that specialize
populations for particular ecological niches and may eventually result in the emergence of new species. In
other words, natural selection is an important process (though not the only process) by which evolution takes
place within a population of organisms. As opposed to artificial selection, in which humans favor specific
traits, in natural selection the environment acts as a sieve through which only certain variations can pass.
Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin
in his influential 1859 book On the Origin of Species,[1] in which natural selection was described as
analogous to artificial selection, a process by which animals and plants with traits considered desirable by
human breeders are systematically favored for reproduction. The concept of natural selection was originally
developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of
modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and
molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary
explanation for adaptive evolution.
Contents
[hide]
• 1 General principles
• 1.1 Nomenclature and usage
• 1.2 Fitness
• 1.3 Types of selection
• 1.4 Sexual selection
• 2 Examples of natural selection
• 3 Evolution by means of natural selection
• 3.1 Speciation
• 4 Historical development
• 4.1 Pre-Darwinian theories
• 4.2 Darwin's theory
• 4.3 Modern evolutionary synthesis
• 5 Impact of the idea
• 5.1 Cell and molecular biology
• 5.2 Social and psychological theory
• 5.3 Information and systems theory
• 6 Genetic basis of natural selection
• 6.1 Genotype and phenotype
• 6.2 Directionality of selection
[edit] General principles

Darwin's illustrations of beak variation in the finches of the Galápagos Islands, which hold 13 closely
related species that differ most markedly in the shape of their beaks. The beak of each species is suited to its
preferred food, suggesting that beak shapes evolved by natural selection.
Natural variation occurs among the individuals of any population of organisms. Many of these
differences do not affect survival (such as differences in eye color in humans), but some differences may
improve the chances of survival of a particular individual. A rabbit that runs faster than others may be more
likely to escape from predators, and algae that are more efficient at extracting energy from sunlight will grow
faster. Individuals that have better odds for survival also have better odds for reproduction.
If the traits that give these individuals a reproductive advantage are also heritable, that is, passed
from parent to child, then there will be a slightly higher proportion of fast rabbits or efficient algae in the next
generation. This is known as differential reproduction. Even if the reproductive advantage is very slight, over
many generations any heritable advantage will become dominant in the population, due to exponential
growth. In this way the natural environment of an organism "selects" for traits that confer a reproductive
advantage, causing gradual changes or evolution of life. This effect was first described and named by
Charles Darwin.
The concept of natural selection predates the understanding of genetics, which is the study of
heredity. In modern times, it is understood that selection acts on an organism's phenotype, or observable
characteristics, but it is the organism's genetic make-up or genotype that is inherited. The phenotype is the
result of the genotype and the environment in which the organism lives (see Genotype-phenotype
distinction).
This is the link between natural selection and genetics, as described in the modern evolutionary
synthesis. Although a complete theory of evolution also requires an account of how genetic variation arises in
the first place (such as by mutation and sexual reproduction) and includes other evolutionary mechanisms
(such as gene flow), natural selection is still understood as a fundamental mechanism for evolution.

[edit] Nomenclature and usage

The term natural selection has slightly different definitions in different contexts. It is most often
defined to operate on heritable traits, because these are the traits that directly participate in evolution.
However, natural selection is "blind" in the sense that changes in phenotype (physical and behavioral
characteristics) can give a reproductive advantage regardless of whether or not the trait is heritable (non
heritable traits can be the result of environmental factors or the life experience of the organism).
Following Darwin's primary usage[1] the term is often used to refer to both the evolutionary
consequence of blind selection and to its mechanisms.[2][3] It is sometimes helpful to explicitly distinguish
between selection's mechanisms and its effects; when this distinction is important, scientists define "natural
selection" specifically as "those mechanisms that contribute to the selection of individuals that reproduce",
without regard to whether the basis of the selection is heritable. This is sometimes referred to as "phenotypic
natural selection".[4]
Traits that cause greater reproductive success of an organism are said to be selected for, whereas
those that reduce success are selected against. Selection for a trait may also result in the selection of other
correlated traits that do not themselves directly influence reproductive advantage. This may occur as a result
of pleiotropy or gene linkage.[5]

[edit] Fitness
Main article: Fitness (biology)
The concept of fitness is central to natural selection. Broadly, individuals which are more "fit" have
better potential for survival, as in the well-known phrase "survival of the fittest". However, as with natural
selection above, the precise meaning of the term is much more subtle, and Richard Dawkins manages in his
later books to avoid it entirely. (He devotes a chapter of his book, The Extended Phenotype, to discussing the
various senses in which the term is used). Modern evolutionary theory defines fitness not by how long an
organism lives, but by how successful it is at reproducing. If an organism lives half as long as others of its
species, but has twice as many offspring surviving to adulthood, its genes will become more common in the
adult population of the next generation.
Though natural selection acts on individuals, the effects of chance mean that fitness can only really
be defined "on average" for the individuals within a population. The fitness of a particular genotype
corresponds to the average effect on all individuals with that genotype. Very low-fitness genotypes cause
their bearers to have few or no offspring on average; examples include many human genetic disorders like
cystic fibrosis.
Since fitness is an averaged quantity, it is also possible that a favorable mutation arises in an
individual that does not survive to adulthood for unrelated reasons. Fitness also depends crucially upon the
environment. Conditions like sickle-cell anemia may have low fitness in the general human population, but
because the sickle-cell trait confers immunity from malaria, it has high fitness value in populations which
have high malaria infection rates.

[edit] Types of selection

Natural selection can act on any phenotypic trait, and selective pressure can be produced by any
aspect of the environment, including sexual selection and competition with members of the same species.
However, this does not imply that natural selection is always directional and results in adaptive evolution;
natural selection often results in the maintenance of the status quo by eliminating less fit variants.
The unit of selection can be the individual or it can be another level within the hierarchy of biological
organisation, such as genes, cells, and kin groups. There is still debate about whether natural selection acts
at the level of groups or species to produce adaptations that benefit a larger, non-kin group. Selection at a
different level such as the gene can result in an increase in fitness for that gene, while at the same time
reducing the fitness of the individuals carrying that gene, in a process called intragenomic conflict. Overall,
the combined effect of all selection pressures at various levels determines the overall fitness of an individual,
and hence the outcome of natural selection.

The life cycle of a sexually reproducing organism. Various components of natural selection are
indicated for each life stage.[6]
Natural selection occurs at every life stage of an individual. An individual organism must survive until
adulthood before it can reproduce, and selection of those that reach this stage is called viability selection. In
many species, adults must compete with each other for mates via sexual selection, and success in this
competition determines who will parent the next generation. When individuals can reproduce more than
once, a longer survival in the reproductive phase increases the number of offspring, called survival selection.
The fecundity of both females and males (for example, giant sperm in certain species of Drosophila)
[7] can be limited via "fecundity selection". The viability of produced gametes can differ, while intragenomic
conflicts such as meiotic drive between the haploid gametes can result in gametic or "genic selection".
Finally, the union of some combinations of eggs and sperm might be more compatible than others; this is
termed compatibility selection.

[edit] Sexual selection

Main article: Sexual selection
It is useful to distinguish between "ecological selection" and "sexual selection". Ecological selection
covers any mechanism of selection as a result of the environment (including relatives, e.g. kin selection,
competition, and infanticide), while "sexual selection" refers specifically to competition for mates.[8]
Sexual selection can be intrasexual, as in cases of competition among individuals of the same sex in
a population, or intersexual, as in cases where one sex controls reproductive access by choosing among a
population of available mates. Most commonly, intrasexual selection involves male–male competition and
intersexual selection involves female choice of suitable males, due to the generally greater investment of
resources for a female than a male in a single offspring. However, some species exhibit sex-role reversed
behavior in which it is males that are most selective in mate choice; the best-known examples of this pattern
occur in some fishes of the family Syngnathidae, though likely examples have also been found in amphibian
and bird species.[9]
 Some features that are confined to one sex only of a particular species can be explained by selection
exercised by the other sex in the choice of a mate, for example, the extravagant plumage of some male birds.
Similarly, aggression between members of the same sex is sometimes associated with very distinctive
features, such as the antlers of stags, which are used in combat with other stags. More generally, intrasexual
selection is often associated with sexual dimorphism, including differences in body size between males and
females of a species.[10]
[edit] Examples of natural selection
 Resistance to antibiotics is increased though the survival of individuals which are immune to the
effects of the antibiotic, whose offspring then inherit the resistance, creating a new population of resistant
bacteria.
A well-known example of natural selection in action is the development of antibiotic resistance in
microorganisms. Since the discovery of penicillin in 1928 by Alexander Fleming, antibiotics have been used
to fight bacterial diseases. Natural populations of bacteria contain, among their vast numbers of individual
members, considerable variation in their genetic material, primarily as the result of mutations. When exposed
to antibiotics, most bacteria die quickly, but some may have mutations that make them slightly less
susceptible. If the exposure to antibiotics is short, these individuals will survive the treatment. This selective
elimination of maladapted individuals from a population is natural selection.
These surviving bacteria will then reproduce again, producing the next generation. Due to the
elimination of the maladapted individuals in the past generation, this population contains more bacteria that
have some resistance against the antibiotic. At the same time, new mutations occur, contributing new genetic
variation to the existing genetic variation. Spontaneous mutations are very rare, and advantageous mutations
are even rarer. However, populations of bacteria are large enough that a few individuals will have beneficial
mutations. If a new mutation reduces their susceptibility to an antibiotic, these individuals are more likely to
survive when next confronted with that antibiotic.
Given enough time, and repeated exposure to the antibiotic, a population of antibiotic-resistant
bacteria will emerge. This new changed population of antibiotic-resistant bacteria is optimally adapted to the
context it evolved in. At the same time, it is not necessarily optimally adapted any more to the old antibiotic
free environment. The end result of natural selection is two populations that are both optimally adapted to
their specific environment, while both perform substandard in the other environment.
 The widespread use and misuse of antibiotics has resulted in increased microbial resistance to
antibiotics in clinical use, to the point that the methicillin-resistant Staphylococcus aureus (MRSA) has been
described as a "superbug" because of the threat it poses to health and its relative invulnerability to existing
drugs.[11] Response strategies typically include the use of different, stronger antibiotics; however, new
strains of MRSA have recently emerged that are resistant even to these drugs.[12]
This is an example of what is known as an evolutionary arms race, in which bacteria continue to
develop strains that are less susceptible to antibiotics, while medical researchers continue to develop new
antibiotics that can kill them. A similar situation occurs with pesticide resistance in plants and insects. Arms
races are not necessarily induced by man; a well-documented example involves the elaboration of the RNA
interference pathway in plants as means of innate immunity against viruses.[13]

[edit] Evolution by means of natural selection

Main articles: Evolution and Darwinism
A prerequisite for natural selection to result in adaptive evolution, novel traits and speciation, is the
presence of heritable genetic variation that results in fitness differences. Genetic variation is the result of
mutations, recombinations and alterations in the karyotype (the number, shape, size and internal
arrangement of the chromosomes). Any of these changes might have an effect that is highly advantageous or
highly disadvantageous, but large effects are very rare. In the past, most changes in the genetic material
were considered neutral or close to neutral because they occurred in noncoding DNA or resulted in a
synonymous substitution. However, recent research suggests that many mutations in non-coding DNA do
have slight deleterious effects.[14][15] Although both mutation rates and average fitness effects of mutations
are dependent on the organism, estimates from data in humans have found that a majority of mutations are
slightly deleterious.[16]
 The exuberant tail of the peacock is thought to be the result of sexual selection by females. This
peacock is an albino; selection against albinos in nature is intense because they are easily spotted by
predators or are unsuccessful in competition for mates.
By the definition of fitness, individuals with greater fitness are more likely to contribute offspring to the
next generation, while individuals with lesser fitness are more likely to die early or fail to reproduce. As a
result, alleles which on average result in greater fitness become more abundant in the next generation, while
alleles which generally reduce fitness become rarer. If the selection forces remain the same for many
generations, beneficial alleles become more and more abundant, until they dominate the population, while
alleles with a lesser fitness disappear. In every generation, new mutations and re-combinations arise
spontaneously, producing a new spectrum of phenotypes. Therefore, each new generation will be enriched
by the increasing abundance of alleles that contribute to those traits that were favored by selection,
enhancing these traits over successive generations.
 Some mutations occur in so-called regulatory genes. Changes in these can have large effects on the
phenotype of the individual because they regulate the function of many other genes. Most, but not all,
mutations in regulatory genes result in non-viable zygotes. Examples of nonlethal regulatory mutations occur
in HOX genes in humans, which can result in a cervical rib[17] or polydactyly, an increase in the number of
fingers or toes.[18] When such mutations result in a higher fitness, natural selection will favor these
phenotypes and the novel trait will spread in the population.

X-ray of the left hand of a ten year old boy with polydactyly.
Established traits are not immutable; traits that have high fitness in one environmental context may
be much less fit if environmental conditions change. In the absence of natural selection to preserve such a
trait, it will become more variable and deteriorate over time, possibly resulting in a vestigial manifestation of
the trait, also called evolutionary baggage. In many circumstances, the apparently vestigial structure may
retain a limited functionality, or may be co-opted for other advantageous traits in a phenomenon known as
preadaptation. A famous example of a vestigial structure, the eye of the blind mole rat, is believed to retain
function in photoperiod perception.[19]

[edit] Speciation
Main article: Speciation
Speciation requires selective mating, which result in a reduced gene flow. Selective mating can be
the result of 1. Geographic isolation, 2. Behavioral isolation, or 3. Temporal isolation. For example, a change
in the physical environment (geographic isolation by an extrinsic barrier) would follow number 1, a change in
camouflage for number 2 or a shift in mating times (i.e., one species of deer shifts location and therefore
changes its "rut") for number 3.[citation needed] Over time, these subgroups might diverge radically to
become different species, either because of differences in selection pressures on the different subgroups, or
because different mutations arise spontaneously in the different populations, or because of founder effects –
some potentially beneficial alleles may, by chance, be present in only one or other of two subgroups when
they first become separated. A lesser-known mechanism of speciation occurs via hybridization, well-
documented in plants and occasionally observed in species-rich groups of animals such as cichlid fishes.[20]
Such mechanisms of rapid speciation can reflect a mechanism of evolutionary change known as punctuated
equilibrium, which suggests that evolutionary change and particularly speciation typically happens quickly
after interrupting long periods of stasis.
Genetic changes within groups result in increasing incompatibility between the genomes of the two
subgroups, thus reducing gene flow between the groups. Gene flow will effectively cease when the distinctive
mutations characterizing each subgroup become fixed. As few as two mutations can result in speciation: if
each mutation has a neutral or positive effect on fitness when they occur separately, but a negative effect
when they occur together, then fixation of these genes in the respective subgroups will lead to two
reproductively isolated populations. According to the biological species concept, these will be two different
species.

[edit] Historical development

Main articles: History of evolutionary thought, Inception of Darwin's theory, and Development of
Darwin's theory
 The modern theory of natural selection derives from the work of Charles Darwin in the nineteenth
century.
[edit] Pre-Darwinian theories
Several ancient philosophers expressed the idea that nature produces a huge variety of creatures,
apparently randomly, and that only those creatures survive that manage to provide for themselves and
reproduce successfully; well-known examples include Empedocles[21] and his intellectual successor,
Lucretius,[22] while related ideas were later refined by Aristotle.[23] The struggle for existence was later
described by Al-Jahiz, who argued that environmental factors influence animals to develop new
characteristics to ensure survival.[24][25][26] Abu Rayhan Biruni described the idea of artificial selection and
argued that nature works in much the same way.[27] Similar ideas were later expressed by Nasir al-Din
Tusi[28] and Ibn Khaldun.[29][30] Such classical arguments were reintroduced in the 18th century by Pierre
Louis Maupertuis[31] and others, including Charles Darwin's grandfather Erasmus Darwin. While these
forerunners had an influence on Darwinism, they later had little influence on the trajectory of evolutionary
thought after Charles Darwin.
Until the early 19th century, the prevailing view in Western societies was that differences between
individuals of a species were uninteresting departures from their Platonic idealism (or typus) of created kinds.
However, the theory of uniformitarianism in geology promoted the idea that simple, weak forces could act
continuously over long periods of time to produce radical changes in the Earth's landscape. The success of
this theory raised awareness of the vast scale of geological time and made plausible the idea that tiny,
virtually imperceptible changes in successive generations could produce consequences on the scale of
differences between species. Early 19th century evolutionists such as Jean Baptiste Lamarck suggested the
inheritance of acquired characteristics as a mechanism for evolutionary change; adaptive traits acquired by
an organism during its lifetime could be inherited by that organism's progeny, eventually causing
transmutation of species.[32] This theory has come to be known as Lamarckism and was an influence on the
anti-genetic ideas of the Stalinist Soviet biologist Trofim Lysenko.[33]
[edit] Darwin's theory
In 1859, Charles Darwin set out his theory of evolution by natural selection as an explanation for
adaptation and speciation. He defined natural selection as the "principle by which each slight variation [of a
trait], if useful, is preserved".[34] The concept was simple but powerful: individuals best adapted to their
environments are more likely to survive and reproduce. As long as there is some variation between them,
there will be an inevitable selection of individuals with the most advantageous variations. If the variations are
inherited, then differential reproductive success will lead to a progressive evolution of particular populations
of a species, and populations that evolve to be sufficiently different eventually become different species.[35]
Darwin's ideas were inspired by the observations that he had made on the Beagle voyage, and by
the work of two political economists. The first was the Reverend Thomas Malthus, who in An Essay on the
Principle of Population, noted that population (if unchecked) increases exponentially whereas the food supply
grows only arithmetically; thus inevitable limitations of resources would have demographic implications,
leading to a "struggle for existence" as a divinely ordained law "in order to rouse man into action, and form
his mind to reason" for the greater good despite the "partial evil" limiting population.[36] The second was
Adam Smith who, in The Wealth of Nations, identified a regulating mechanism in free markets, which he
referred to as the "invisible hand", which suggests that prices self-adjust according to supplies and demand.
[37] When Darwin read Malthus in 1838 he was already primed by his work as a naturalist to appreciate the
"struggle for existence" in nature and it struck him that as population outgrew resources, "favourable
variations would tend to be preserved, and unfavourable ones to be destroyed. The result of this would be
the formation of new species."[38]
Here is Darwin's own summary of the idea, which can be found in the fourth chapter of the Origin:
 If during the long course of ages and under varying conditions of life, organic beings vary at all
in the several parts of their organisation, and I think this cannot be disputed; if there be, owing to the
high geometrical powers of increase of each species, at some age, season, or year, a severe struggle
for life, and this certainly cannot be disputed; then, considering the infinite complexity of the relations of
all organic beings to each other and to their conditions of existence, causing an infinite diversity in
structure, constitution, and habits, to be advantageous to them, I think it would be a most extraordinary
fact if no variation ever had occurred useful to each being's own welfare, in the same way as so many
variations have occurred useful to man. But if variations useful to any organic being do occur,
assuredly individuals thus characterised will have the best chance of being preserved in the struggle
for life; and from the strong principle of inheritance they will tend to produce offspring similarly
characterised. This principle of preservation, I have called, for the sake of brevity, Natural Selection.
Once he had his theory "by which to work", Darwin was meticulous about gathering and refining
evidence as his "prime hobby" before making his idea public. He was in the process of writing his "big book"
to present his researches when the naturalist Alfred Russel Wallace independently conceived of the principle
and described it in an essay he sent to Darwin to forward to Charles Lyell. Lyell and Joseph Dalton Hooker
decided (without Wallace's knowledge) to present his essay together with unpublished writings which Darwin
had sent to fellow naturalists, and On the Tendency of Species to form Varieties; and on the Perpetuation of
Varieties and Species by Natural Means of Selection was read to the Linnean Society announcing co-
discovery of the principle in July 1858.[39] Darwin published a detailed account of his evidence and
conclusions in On the Origin of Species in 1859. In the 3rd edition of 1861 Darwin acknowledged that others
— notably William Charles Wells in 1813, and Patrick Matthew in 1831 — had proposed similar ideas, but had
neither developed them nor presented them in notable scientific publications.[40]
 Darwin thought of natural selection by analogy to how farmers select crops or livestock for breeding,
which he called "artificial selection"; in his early manuscripts he referred to a 'Nature' which would do the
selection. At the time, other mechanisms of evolution such as evolution by genetic drift were not yet explicitly
formulated, and Darwin believed that selection was likely only part of the story: "I am convinced that [it] has
been the main, but not exclusive means of modification."[41] In a letter to Charles Lyell in September 1860,
Darwin regretted the use of the term "Natural Selection", preferring the term "Natural Preservation".[42] For
Darwin and his contemporaries, natural selection was essentially synonymous with evolution by natural
selection. After the publication of On the Origin of Species, educated people generally accepted that
evolution had occurred in some form. However, natural selection remained controversial as a mechanism,
partly because it was perceived to be too weak to explain the range of observed characteristics of living
organisms, and partly because even supporters of evolution balked at its "unguided" and non-progressive
nature,[43] a response that has been characterized as the single most significant impediment to the idea's
acceptance.[44] However, some thinkers enthusiastically embraced natural selection; after reading Darwin,
Herbert Spencer introduced the term survival of the fittest, which became a popular summary of the theory.
[45] The fifth edition of On the Origin of Species published in 1869 included Spencer's phrase as an
alternative to natural selection, with credit given: "But the expression often used by Mr. Herbert Spencer, of
the Survival of the Fittest, is more accurate, and is sometimes equally convenient."[46] Although the phrase
is still often used by non-biologists, modern biologists avoid it because it is tautological if "fittest" is read to
mean "functionally superior" and is applied to individuals rather than considered as an averaged quantity
over populations.[47]

[edit] Modern evolutionary synthesis

Main article: Modern evolutionary synthesis
 Natural selection relies crucially on the idea of heredity, but it was developed long before the basic
concepts of genetics. Although the Austrian monk Gregor Mendel, the father of modern genetics, was a
contemporary of Darwin's, his work would lie in obscurity until the early 20th century. Only after the
integration of Darwin's theory of evolution with a complex statistical appreciation of Gregor Mendel's 're-
discovered' laws of inheritance did natural selection become generally accepted by scientists. The work of
Ronald Fisher (who developed the required mathematical language and The Genetical Theory of Natural
Selection),[2] J.B.S. Haldane (who introduced the concept of the "cost" of natural selection),[48] Sewall
Wright (who elucidated the nature of selection and adaptation),[49] Theodosius Dobzhansky (who
established the idea that mutation, by creating genetic diversity, supplied the raw material for natural
selection: see Genetics and the Origin of Species),[50] William Hamilton (who conceived of kin selection),
Ernst Mayr (who recognised the key importance of reproductive isolation for speciation: see Systematics and
the Origin of Species)[51] and many others formed the modern evolutionary synthesis. This synthesis
cemented natural selection as the foundation of evolutionary theory, where it remains today.

[edit] Impact of the idea

Darwin's ideas, along with those of Adam Smith and Karl Marx, had a profound influence on 19th
century thought. Perhaps the most radical claim of the theory of evolution through natural selection is that
"elaborately constructed forms, so different from each other, and dependent on each other in so complex a
manner" evolved from the simplest forms of life by a few simple principles. This claim inspired some of
Darwin's most ardent supporters—and provoked the most profound opposition. The radicalism of natural
selection, according to Stephen Jay Gould,[52] lay in its power to "dethrone some of the deepest and most
traditional comforts of Western thought". In particular, it challenged long-standing beliefs in such concepts as
a special and exalted place for humans in the natural world and a benevolent creator whose intentions were
reflected in nature's order and design.
[edit] Cell and molecular biology
In the 19th century, Wilhelm Roux, a founder of modern embryology, wrote a book entitled « Der
Kampf der Teile im Organismus » (The struggle of parts in the organism) in which he suggested that the
development of an organism results from a Darwinian competition between the parts of the embryo,
occurring at all levels, from molecules to organs. In recent years, a modern version of this theory has been
proposed by Jean-Jacques Kupiec. According to this cellular Darwinism, stochasticity at the molecular level
generates diversity in cell types whereas cell interactions impose a characteristic order on the developing
embryo.

[edit] Social and psychological theory

The social implications of the theory of evolution by natural selection also became the source of
continuing controversy. Friedrich Engels, a German political philosopher and co-originator of the ideology of
communism, wrote in 1872 that "Darwin did not know what a bitter satire he wrote on mankind when he
showed that free competition, the struggle for existence, which the economists celebrate as the highest
historical achievement, is the normal state of the animal kingdom".[53] Interpretation of natural selection as
necessarily 'progressive', leading to increasing 'advances' in intelligence and civilisation, was used as a
justification for colonialism and policies of eugenics, as well as broader sociopolitical positions now described
as Social Darwinism. Konrad Lorenz won the Nobel Prize in Physiology or Medicine in 1973 for his analysis
of animal behavior in terms of the role of natural selection (particularly group selection). However, in
Germany in 1940, in writings that he subsequently disowned, he used the theory as a justification for policies
of the Nazi state. He wrote "... selection for toughness, heroism, and social utility...must be accomplished by
some human institution, if mankind, in default of selective factors, is not to be ruined by domestication-
induced degeneracy. The racial idea as the basis of our state has already accomplished much in this
respect."[54] Others have developed ideas that human societies and culture evolve by mechanisms that are
analogous to those that apply to evolution of species.[55]
More recently, work among anthropologists and psychologists has led to the development of
sociobiology and later evolutionary psychology, a field that attempts to explain features of human psychology
in terms of adaptation to the ancestral environment. The most prominent such example, notably advanced in
the early work of Noam Chomsky and later by Steven Pinker, is the hypothesis that the human brain is
adapted to acquire the grammatical rules of natural language.[56] Other aspects of human behavior and
social structures, from specific cultural norms such as incest avoidance to broader patterns such as gender
roles, have been hypothesized to have similar origins as adaptations to the early environment in which
modern humans evolved. By analogy to the action of natural selection on genes, the concept of memes –
"units of cultural transmission", or culture's equivalents of genes undergoing selection and recombination –
has arisen, first described in this form by Richard Dawkins[57] and subsequently expanded upon by
philosophers such as Daniel Dennett as explanations for complex cultural activities, including human
consciousness.[58] Extensions of the theory of natural selection to such a wide range of cultural phenomena
have been distinctly controversial and are not widely accepted.[59]

[edit] Information and systems theory

In 1922, Alfred Lotka proposed that natural selection might be understood as a physical principle
which could be described in terms of the use of energy by a system,[60] a concept that was later developed
by Howard Odum as the maximum power principle whereby evolutionary systems with selective advantage
maximise the rate of useful energy transformation. Such concepts are sometimes relevant in the study of
applied thermodynamics.
 The principles of natural selection have inspired a variety of computational techniques, such as "soft"
artificial life, that simulate selective processes and can be highly efficient in 'adapting' entities to an
environment defined by a specified fitness function.[61] For example, a class of heuristic optimization
algorithms known as genetic algorithms, pioneered by John Holland in the 1970s and expanded upon by
David E. Goldberg,[62] identify optimal solutions by simulated reproduction and mutation of a population of
solutions defined by an initial probability distribution.[63] Such algorithms are particularly useful when applied
to problems whose solution landscape is very rough or has many local minima.

[edit] Genetic basis of natural selection

The idea of natural selection predates the understanding of genetics. We now have a much better
idea of the biology underlying heritability, which is the basis of natural selection.

[edit] Genotype and phenotype

See also: Genotype-phenotype distinction.
Natural selection acts on an organism's phenotype, or physical characteristics. Phenotype is
determined by an organism's genetic make-up (genotype) and the environment in which the organism lives.
Often, natural selection acts on specific traits of an individual, and the terms phenotype and genotype are
used narrowly to indicate these specific traits.
When different organisms in a population possess different versions of a gene for a certain trait, each
of these versions is known as an allele. It is this genetic variation that underlies phenotypic traits. A typical
example is that certain combinations of genes for eye color in humans which, for instance, give rise to the
phenotype of blue eyes. (On the other hand, when all the organisms in a population share the same allele for
a particular trait, and this state is stable over time, the allele is said to be fixed in that population.)
Some traits are governed by only a single gene, but most traits are influenced by the interactions of
many genes. A variation in one of the many genes that contributes to a trait may have only a small effect on
the phenotype; together, these genes can produce a continuum of possible phenotypic values.[64]

[edit] Directionality of selection

When some component of a trait is heritable, selection will alter the frequencies of the different
alleles, or variants of the gene that produces the variants of the trait. Selection can be divided into three
classes, on the basis of its effect on allele frequencies.[65]
Directional selection occurs when a certain allele has a greater fitness than others, resulting in an
increase of its frequency. This process can continue until the allele is fixed and the entire population shares
the fitter phenotype. It is directional selection that is illustrated in the antibiotic resistance example above.
Far more common is stabilizing selection (which is commonly confused with purifying selection[66]
[67]), which lowers the frequency of alleles that have a deleterious effect on the phenotype – that is, produce
organisms of lower fitness. This process can continue until the allele is eliminated from the population.
Purifying selection results in functional genetic features, such as protein-coding genes or regulatory
sequences, being conserved over time due to selective pressure against deleterious variants.
Finally, a number of forms of balancing selection exist, which do not result in fixation, but maintain an
allele at intermediate frequencies in a population. This can occur in diploid species (that is, those that have
two pairs of chromosomes) when heterozygote individuals, who have different alleles on each chromosome
at a single genetic locus, have a higher fitness than homozygote individuals that have two of the same
alleles. This is called heterozygote advantage or overdominance, of which the best-known example is the
malarial resistance observed in heterozygous humans who carry only one copy of the gene for sickle cell
anemia. Maintenance of allelic variation can also occur through disruptive or diversifying selection, which
favors genotypes that depart from the average in either direction (that is, the opposite of overdominance),
and can result in a bimodal distribution of trait values. Finally, balancing selection can occur through
frequency-dependent selection, where the fitness of one particular phenotype depends on the distribution of
other phenotypes in the population. The principles of game theory have been applied to understand the
fitness distributions in these situations, particularly in the study of kin selection and the evolution of reciprocal
altruism.[68][69]

[edit] Selection and genetic variation

A portion of all genetic variation is functionally neutral in that it produces no phenotypic effect or
significant difference in fitness; the hypothesis that this variation accounts for a large fraction of observed
genetic diversity is known as the neutral theory of molecular evolution and was originated by Motoo Kimura.
When genetic variation does not result in differences in fitness, selection cannot directly affect the frequency
of such variation. As a result, the genetic variation at those sites will be higher than at sites where variation
does influence fitness.[65] However, after a period with no new mutation, the genetic variation at these sites
will be eliminated due to genetic drift.

[edit] Mutation selection balance

Natural selection results in the reduction of genetic variation through the elimination of maladapted
individuals and consequently of the mutations that caused the maladaptation. At the same time, new
mutations occur, resulting in a mutation-selection balance. The exact outcome of the two processes depends
both on the rate at which new mutations occur and on the strength of the natural selection, which is a function
of how unfavorable the mutation proves to be. Consequently, changes in the mutation rate or the selection
pressure will result in a different mutation-selection balance.

[edit] Genetic linkage

Genetic linkage occurs when the loci of two alleles are linked, or in close proximity to each other on
the chromosome. During the formation of gametes, recombination of the genetic material results in
reshuffling of the alleles. However, the chance that such a reshuffle occurs between two alleles depends on
the distance between those alleles; the closer the alleles are to each other, the less likely it is that such a
reshuffle will occur. Consequently, when selection targets one allele, this automatically results in selection of
the other allele as well; through this mechanism, selection can have a strong influence on patterns of
variation in the genome.
Selective sweeps occur when an allele becomes more common in a population as a result of positive
selection. As the prevalence of one allele increases, linked alleles can also become more common, whether
they are neutral or even slightly deleterious. This is called genetic hitchhiking. A strong selective sweep
results in a region of the genome where the positively selected haplotype (the allele and its neighbours) are
essentially the only ones that exist in the population.
Whether a selective sweep has occurred or not can be investigated by measuring linkage
disequilibrium, or whether a given haplotype is overrepresented in the population. Normally, genetic
recombination results in a reshuffling of the different alleles within a haplotype, and none of the haplotypes
will dominate the population. However, during a selective sweep, selection for a specific allele will also result
in selection of neighbouring alleles. Therefore, the presence of a block of strong linkage disequilibrium might
indicate that there has been a 'recent' selective sweep near the center of the block, and this can be used to
identify sites recently under selection.
Background selection is the opposite of a selective sweep. If a specific site experiences strong and
persistent purifying selection, linked variation will tend to be weeded out along with it, producing a region in
the genome of low overall variability. Because background selection is a result of deleterious new mutations,
which can occur randomly in any haplotype, it does not produce clear blocks of linkage disequilibrium,
although with low recombination it can still lead to slightly negative linkage disequilibrium overall.[70]

[edit] See also

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Artificial selection
• Co-evolution
• Gene-centered view of evolution
• Negative selection
• Unit of selection
[edit] References
1. ^ a b Darwin C (1859) On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life John Murray, London; modern reprint
Charles Darwin, Julian Huxley (2003). On The Origin of Species. Signet Classics. ISBN 0-451-
52906-5. Published online at The complete work of Charles Darwin online: On the origin of species
by means of natural selection, or the preservation of favoured races in the struggle for life.
2. ^ a b Fisher RA (1930) The Genetical Theory of Natural Selection Clarendon Press, Oxford
3. ^ Works employing or describing this usage:
Endler JA (1986). Natural Selection in the Wild. Princeton, New Jersey: Princeton University Press.
ISBN 0-691-00057-3.
Williams GC (1966). Adaptation and Natural Selection. Oxford University Press.
4. ^ Works employing or describing this usage:
Lande R & Arnold SJ (1983) The measurement of selection on correlated characters. Evolution
37:1210-26
Futuyma DJ (2005) Evolution. Sinauer Associates, Inc., Sunderland, Massachusetts. ISBN 0-87893-
187-2
Haldane, J.B.S. 1953. The measurement of natural selection. Proceedings of the 9th International
Congress of Genetics. 1: 480-487
5. ^ Sober E (1984; 1993) The Nature of Selection: Evolutionary Theory in Philosophical Focus
University of Chicago Press ISBN 0-226-76748-5
6. ^ Modified from Christiansen FB (1984) The definition and measurement of fitness. In:
Evolutionary ecology (ed. Shorrocks B) pp65–79. Blackwell Scientific, Oxford by adding survival
selection in the reproductive phase
 7. ^ Pitnick S & Markow TA (1994) Large-male advantage associated with the costs of sperm
production in Drosophila hydei, a species with giant sperm. Proc Natl Acad Sci USA 91:9277-81;
Pitnick S (1996) Investment in testes and the cost of making long sperm in Drosophila. Am Nat
148:57-80
8. ^ Andersson, M (1995). Sexual Selection. Princeton, New Jersey: Princeton University
Press. ISBN 0-691-00057-3.
9. ^ Eens M, Pinxten R. (2000). Sex-role reversal in vertebrates: behavioural and
endocrinological accounts. Behav Processes 51(1-3):135-147. PMID 11074317
10.^ Barlow GW. (2005). How Do We Decide that a Species is Sex-Role Reversed? The
Quarterly Review of Biology 80(1):28–35. PMID 15884733
11.^ "MRSA Superbug News". http://www.inboxrobot.com/news/MRSASuperbug. Retrieved
2006-05-06.
12.^ Schito GC (2006). "The importance of the development of antibiotic resistance in
Staphylococcus aureus". Clin Microbiol Infect 12 Suppl 1: 3–8. doi:10.1111/j.1469-
0691.2006.01343.x. PMID 16445718. [1]
13.^ Lucy A, Guo H, Li W, Ding S (2000). "Suppression of post-transcriptional gene silencing by
a plant viral protein localized in the nucleus". EMBO J 19 (7): 1672–80. PMID 10747034.
14.^ Kryukov GV, Schmidt S & Sunyaev S (2005) Small fitness effect of mutations in highly
conserved non-coding regions. Human Molecular Genetics 14:2221-9
15.^ Bejerano G, Pheasant M, Makunin I, Stephen S, Kent WJ, Mattick JS & Haussler D (2004)
Ultraconserved elements in the human genome. Science 304:1321-5
16.^ Eyre-Walker A, Woolfit M, Phelps T. (2006). The distribution of fitness effects of new
deleterious amino acid mutations in humans. Genetics 173(2):891-900. PMID 16547091
 17.^ Galis F (1999) Why do almost all mammals have seven cervical vertebrae? developmental
constraints, Hox genes, and cancer. J Exp Zool 285:19-26
18.^ Zakany J, FromentalRamain C, Warot X & Duboule D (1997) Regulation of number and
size of digits by posterior Hox genes: a dose-dependent mechanism with potential evolutionary
implications. Proc Natl Acad Sci USA 94:13695-700
19.^ Sanyal S, Jansen HG, de Grip WJ, Nevo E, de Jong WW. (1990). The eye of the blind mole
rat, Spalax ehrenbergi. Rudiment with hidden function? Invest Ophthalmol Vis Sci. 1990 31(7):1398-
404. PMID 2142147
20.^ Salzburger W, Baric S, Sturmbauer C. (2002). Speciation via introgressive hybridization in
East African cichlids? Mol Ecol 11(3): 619–625. PMID 11918795
21.^ Empedocles. On Nature. Book II
22.^ Lucretius. De rerum natura. Book V
23.^ Aristotle. Physics. Book II, Chapters 4 and 8
24.^ Conway Zirkle (1941). Natural Selection before the "Origin of Species", Proceedings of the
American Philosophical Society 84 (1), p. 71-123.
25.^ Mehmet Bayrakdar (Third Quarter, 1983). "Al-Jahiz And the Rise of Biological
Evolutionism", The Islamic Quarterly. London.
26.^ Paul S. Agutter & Denys N. Wheatley (2008). Thinking about Life: The History and
Philosophy of Biology and Other Sciences. Springer. p. 43. ISBN 1402088655
27.^ Jan Z. Wilczynski (December 1959). "On the Presumed Darwinism of Alberuni Eight
Hundred Years before Darwin". Isis 50 (4): 459–466. doi:10.1086/348801
28.^ Farid Alakbarov (Summer 2001). A 13th-Century Darwin? Tusi's Views on Evolution,
Azerbaijan International 9 (2).
29.^ Franz Rosenthal and Ibn Khaldun, Muqaddimah, Chapter 6, Part 5
 30.^ Franz Rosenthal and Ibn Khaldun, Muqaddimah, Chapter 6, Part 29
31.^ Maupertuis, Pierre Louis (1748). "Derivation of the laws of motion and equilibrium from a
metaphysical principle (Original French text)". Histoire de l'academie des sciences et belle lettres de
Berlin 1746: 267–294.
32.^ Chevalier de Lamarck J-B, de Monet PA (1809) Philosophie Zoologique
33.^ Joravsky D. (1959). Soviet Marxism and Biology before Lysenko. Journal of the History of
Ideas 20(1):85-104.
34.^ Darwin 1859, p. 61
35.^ Darwin 1859, p. 5
36.^ T. Robert Malthus (1798). "An Essay on the Principle of Population". Rogers State
University. http://www.faculty.rsu.edu/~felwell/Theorists/Malthus/Essay.htm#112. Retrieved 2008-11-
03.
37.^ Orrell, David (2007) Apollo's Arrow Toronto: HarperCollins Publishers Ltd. [2]
38.^ Charles Darwin; ed. Nora Barlow (1958). "The autobiography of Charles Darwin 1809-
1882". London: Collins. pp. 120. http://darwin-online.org.uk/content/frameset?
viewtype=text&itemID=F1497&pageseq=124. Retrieved 2008-11-03.
39.^ Wallace, Alfred Russel (1870) Contributions to the Theory of Natural Selection New York:
Macmillan & Co. [3]
40.^ Darwin 1861, p. xiii
41.^ Darwin 1859, p. 6
42.^ "Darwin Correspondence Online Database: Darwin, C. R. to Lyell, Charles, 28 September
1860". http://www.darwinproject.ac.uk/darwinletters/calendar/entry-2931.html. Retrieved 2006-05-
10.
 43.^ Eisley L. (1958). Darwin's Century: Evolution and the Men Who Discovered It. Doubleday &
Co: New York, USA.
44.^ Kuhn TS. [1962] (1996). The Structure of Scientific Revolution 3rd ed. University of
Chicago Press: Chicago, Illinois, USA. ISBN 0-226-45808-3
45.^ "Letter 5145 — Darwin, C. R. to Wallace, A. R., 5 July (1866)". Darwin Correspondence
Project. http://www.darwinproject.ac.uk/entry-5145#mark-5145.f3. Retrieved 2010-01-12.
Maurice E. Stucke. "Better Competition Advocacy". http://works.bepress.com/cgi/viewcontent.cgi?
article=1000&context=maurice_stucke. Retrieved 2007-08-29. "Herbert Spencer in his Principles of
Biology of 1864, vol. 1, p. 444, wrote “This survival of the fittest, which I have here sought to express
in mechanical terms, is that which Mr. Darwin has called ‘natural selection’, or the preservation of
favoured races in the struggle for life.”"
46.^ Darwin 1872, p. 49.
47.^ Mills SK, Beatty JH. [1979] (1994). The Propensity Interpretation of Fitness. Originally in
Philosophy of Science (1979) 46: 263-286; republished in Conceptual Issues in Evolutionary Biology
2nd ed. Elliott Sober, ed. MIT Press: Cambridge, Massachusetts, USA. pp3-23. ISBN 0-262-69162-0.
48.^ Haldane JBS (1932) The Causes of Evolution; Haldane JBS (1957) The cost of natural
selection. J Genet 55:511-24([4].
49.^ Wright S (1932) The roles of mutation, inbreeding, crossbreeding and selection in evolution
Proc 6th Int Cong Genet 1:356–66
50.^ Dobzhansky Th (1937) Genetics and the Origin of Species Columbia University Press, New
York. (2nd ed., 1941; 3rd edn., 1951)
51.^ Mayr E (1942) Systematics and the Origin of Species Columbia University Press, New
York. ISBN 0-674-86250-3
52.^ The New York Review of Books: Darwinian Fundamentalism (accessed May 6, 2006)
 53.^ Engels F (1873-86) Dialectics of Nature 3d ed. Moscow: Progress, 1964 [5]
54.^ Quoted in translation in Eisenberg L (2005) Which image for Lorenz? Am J Psychiatry
162:1760 [6]
55.^ e.g. Wilson, DS (2002) Darwin's Cathedral: Evolution, Religion, and the Nature of Society .
University of Chicago Press, ISBN 0-226-90134-3
56.^ Pinker S. [1994] (1995). The Language Instinct: How the Mind Creates Language.
HarperCollins: New York, NY, USA. ISBN 0-06-097651-9
57.^ Dawkins R. [1976] (1989). The Selfish Gene. Oxford University Press: New York, NY, USA,
p.192. ISBN 0-19-286092-5
58.^ Dennett DC. (1991). Consciousness Explained. Little, Brown, and Co: New York, NY, USA.
ISBN 0-316-18066-1
59.^ For example, see Rose H, Rose SPR, Jencks C. (2000). Alas, Poor Darwin: Arguments
Against Evolutionary Psychology. Harmony Books. ISBN 0-609-60513-5
60.^ Lotka AJ (1922a) Contribution to the energetics of evolution [PDF] Proc Natl Acad Sci USA
8:147–51
Lotka AJ (1922b) Natural selection as a physical principle [PDF] Proc Natl Acad Sci USA 8:151–4
61.^ Kauffman SA (1993) The Origin of order. Self-organization and selection in evolution. New
York: Oxford University Press ISBN 0-19-507951-5
62.^ Goldberg DE. (1989). Genetic Algorithms in Search, Optimization and Machine Learning.
Addison-Wesley: Boston, MA, USA
63.^ Mitchell, Melanie, (1996), An Introduction to Genetic Algorithms, MIT Press, Cambridge,
MA.
64.^ Falconer DS & Mackay TFC (1996) Introduction to Quantitative Genetics Addison Wesley
Longman, Harlow, Essex, UK ISBN 0-582-24302-5
 65.^ a b Rice SH. (2004). Evolutionary Theory: Mathematical and Conceptual Foundations.
Sinauer Associates: Sunderland, Massachusetts, USA. ISBN 0-87893-702-1 See esp. ch. 5 and 6 for
a quantitative treatment.
66.^ Lemey, Philippe; Marco Salemi, Anne-Mieke Vandamme (2009). The Phylogenetic
Handbook. Cambridge University Press. ISBN 978-0-521-73071.
67.^ http://www.nature.com/scitable/topicpage/Negative-Selection-1136
68.^ Hamilton WD. (1964). The genetical evolution of social behaviour I and II. Journal of
Theoretical Biology 7: 1-16 and 17-52. PMID 5875341 PMID 5875340
69.^ Trivers RL. (1971). The evolution of reciprocal altruism. Q Rev Biol 46: 35-57.
70.^ Keightley PD. and Otto SP (2006). "Interference among deleterious mutations favours sex
and recombination in finite populations". Nature 443: 89-92.

[edit] Further reading

• For technical audiences
• Gould, Stephen Jay (2002). The Structure of Evolutionary Theory. Harvard University
Press. ISBN 0-674-00613-5.
• Maynard Smith, John (1993). The Theory of Evolution: Canto Edition. Cambridge
University Press. ISBN 0-521-45128-0.
• Popper, Karl (1978) Natural selection and the emergence of mind. Dialectica 32:339-
55. See [7]
• Sober, Elliott (1984) The Nature of Selection: Evolutionary Theory in Philosophical
Focus. University of Chicago Press.
 • Williams, George C. (1966) Adaptation and Natural Selection: A Critique of Some
Current Evolutionary Thought. Oxford University Press.
• Williams George C. (1992) Natural Selection: Domains, Levels and Challenges.
Oxford University Press.
• For general audiences
• Dawkins, Richard (1996) Climbing Mount Improbable. Penguin Books, ISBN 0-670-
85018-7.
• Dennett, Daniel (1995) Darwin's Dangerous Idea: Evolution and the Meanings of
Life. Simon & Schuster ISBN 0-684-82471-X.
• Gould, Stephen Jay (1997) Ever Since Darwin: Reflections in Natural History.
Norton, ISBN 0-393-06425-5.
• Jones, Steve (2001) Darwin's Ghost: The Origin of Species Updated. Ballantine
Books ISBN 0-345-42277-5. Also published in Britain under the title Almost like a whale: the
origin of species updated. Doubleday. ISBN 1-86230-025-9.
• Lewontin, Richard (1978) Adaptation. Scientific American 239:212-30
• Weiner, Jonathan (1994) The Beak of the Finch: A Story of Evolution in Our Time.
Vintage Books, ISBN 0-679-73337-X.
• Historical
• Zirkle C (1941). Natural Selection before the "Origin of Species", Proceedings of the
American Philosophical Society 84 (1), p. 71-123.
 • Kohm M (2004) A Reason for Everything: Natural Selection and the English
Imagination. London: Faber and Faber. ISBN 0-571-22392-3. For review, see [8] van Wyhe J
(2005) Human Nature Review 5:1-4

[edit] External links

• On the Origin of Species by Charles Darwin – Chapter 4,Natural Selection
• Natural Selection- Modeling for Understanding in Science Education, University of Wisconsin
• Natural Selection from University of Berkeley education website
• T. Ryan Gregory: Understanding Natural Selection: Essential Concepts and Common
Misconceptions Evolution: Education and Outreach

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic Genetic drift · Gene flow · Mutation · Natural selection

mechanisms

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)
 Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

[hide]
v•d•e
Topics in population genetics

Key concepts Hardy-Weinberg law · Genetic linkage · Linkage disequilibrium · Fisher's

fundamental theorem · Neutral theory · Price equation

Selection Natural · Sexual · Artificial · Ecological

Effects of
selection Genetic hitchhiking · Background selection
on genomic variation

Genetic drift Small population size · Population bottleneck · Founder effect ·

Coalescence · Balding–Nichols model
Founders R. A. Fisher · J. B. S. Haldane · Sewall Wright

Related topics Evolution · Microevolution · Evolutionary game theory · Fitness landscape ·

Genetic genealogy

List of evolutionary biology topics

Retrieved from "http://en.wikipedia.org/wiki/Natural_selection"

Categories: Selection | Evolutionary biology | Ecological processes

• This page was last modified on 25 August 2010 at 22:57.

 W000

Sélection naturelle
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Les mécanismes de
l'évolution biologique

Mécanismes non aléatoires :

• sélection naturelle
• sélection
 utilitaire
• sélection
sexuelle
• sélection de
parentèle
• sélection de
groupe
• sélection
stratégique
• sélection artificielle

Mécanismes aléatoires :
• mutation génétique
• recombinaison
• dérive génétique

Conséquences de l'évolution :
• spéciation
• adaptation des
espèces
• radiation évolutive
 En biologie, la sélection naturelle est l'un des mécanismes qui guident l'évolution des espèces. Ce
mécanisme est particulièrement important du fait qu'il explique l'adaptation des espèces aux milieux. La
théorie de la sélection naturelle permet d'expliquer et de comprendre comment l'environnement influe sur
l'évolution des espèces et des populations en sélectionnant les individus les plus adaptés et constitue donc
un aspect fondamental de la théorie de l'évolution.
De façon sommaire, la sélection naturelle désigne le fait que les traits qui favorisent la survie et la
reproduction, voient leur fréquence s'accroître d'une génération à l'autre. Cela découle logiquement du fait
que les porteurs de ces traits ont plus de descendants, et aussi que ces derniers portent ces traits (puisqu'ils
sont héréditaires).
Sommaire
[masquer]
• 1 Historique
• 2 Principes de la sélection naturelle
• 2.1 Principe 1 : Les individus diffèrent les uns des
autres
• 2.2 Principe 2 : Les individus les plus adaptés au
milieu survivent et se reproduisent davantage
• 2.3 Principe 3 : Les caractéristiques avantageuses
doivent être héréditaires
• 2.4 Une histoire imaginée par Richard Dawkins
• 3 La sélection naturelle explique l'adaptation des espèces à
leur milieu
• 3.1 Adaptations convergentes
• 4 Origine des variations héréditaires dans une population
• 4.1 L'information génétique portée par l'ADN est
relativement instable
• 4.2 La sélection naturelle agit après les
modifications de l'information génétique
• 4.3 L'hypothèse des caractères acquis
• 5 La sélection naturelle en génétique des populations
• 6 Cas de sélection naturelle scientifiquement démontrés
Historique [modifier]

Charles Darwin
Alors que plusieurs théories évolutives existaient déjà sous le nom de transformisme, Charles
Darwin (1809-1882) propose ce mécanisme que l'on désigne sous le terme de darwinisme ou sélection
darwinienne. Le terme "sélection naturelle" a été imaginé par Darwin par analogie avec la sélection artificielle
pratiquée par les humains depuis des millénaires : les agriculteurs ou éleveurs choisissent à chaque
génération les individus présentant les meilleurs caractéristiques pour les faire se reproduire. Le mécanisme
de sélection darwinienne permet donc d'expliquer de façon naturaliste la complexité adaptative des êtres
vivants, sans avoir recours au finalisme ni à une intervention surnaturelle, d'origine divine, par exemple.

Principes de la sélection naturelle [modifier]

La théorie de la sélection naturelle telle qu'elle a été initialement décrite par Charles Darwin, repose
sur trois principes[1]:
 1. le principe de variation
2. le principe d'adaptation
3. le principe d'hérédité

Principe 1 : Les individus diffèrent les uns des autres [modifier]

En général, dans une population d'individus d'une même espèce, il existe des différences plus ou
moins importantes entre ces individus. En biologie, on appelle caractère, tout ce qui est visible et peut varier
d'un individu à l'autre. On dit qu'il existe plusieurs traits pour un même caractère. Par exemple, chez l'être
humain, la couleur de la peau, la couleur des yeux sont des caractères pour lesquels il existe de multiples
variations ou traits. La variation d'un caractère chez un individu donné constitue son phénotype. C'est là, la
première condition pour qu'il y ait sélection naturelle : au sein d'une population, certains caractères doivent
présenter des variations, c'est le principe de variation.
Principe 2 : Les individus les plus adaptés au milieu survivent et se
reproduisent davantage [modifier]

Certains individus portent des variations qui leur permettent de se reproduire davantage que les
autres, dans un environnement précis. On dit qu'ils disposent d'un avantage sélectif sur leurs congénères:
• La première possibilité est, par exemple, qu'en échappant mieux aux prédateurs, en étant
moins malades, en accédant plus facilement à la nourriture, ces individus atteignent plus facilement
l'âge adulte, pour être apte à la reproduction. Ceux qui ont une meilleure capacité de survie pourront
donc se reproduire davantage.
• Dans le cas particulier de la reproduction sexuée, les individus ayant survécu peuvent être
porteurs d'un caractère particulièrement attirant pour les partenaires de sexe opposé. Ceux-là seront
capables d'engendrer une plus grande descendance en copulant davantage.
 Dans les deux cas, l'augmentation de la capacité à survivre et à se reproduire se traduit par une
augmentation du taux de reproduction et donc par une descendance plus nombreuse, pour les individus
porteurs de ces caractéristiques. On dit alors que ce trait de caractère donné offre un avantage sélectif, par
rapport à d'autres. C'est dans ce principe d'adaptation uniquement, qu'intervient le milieu de vie.

Principe 3 : Les caractéristiques avantageuses doivent être héréditaires

[modifier]

La troisième condition pour qu'il y ait sélection naturelle est que les caractéristiques des individus
doivent être héréditaires, c'est-à-dire qu'elles puissent être transmises à leur descendance. En effet certains
caractères, comme le bronzage ou la culture, ne dépendent pas du génotype, c'est-à-dire l'ensemble des
gènes de l'individu. Lors de la reproduction, ce sont donc les gènes qui, transmis aux descendants,
entraîneront le passage de certains caractères d'une génération à l'autre. C'est le principe d'hérédité.
Ces trois premiers principes entraînent donc que les variations héréditaires qui confèrent un
avantage sélectif seront davantage transmises à la génération suivante que les variations moins
avantageuses. En effet les individus qui portent les variations avantageuses se reproduisent plus. Au fil des
générations, on verra donc la fréquence des gènes désavantageux diminuer jusqu'à éventuellement
disparaître, tandis que les variations avantageuses se répandront dans la population, jusqu'à éventuellement
être partagées par tous les membres de la population ou de l'espèce. Par exemple, dans la population
humaine, la bipédie est un caractère commun à tous les êtres humains modernes.

Une histoire imaginée par Richard Dawkins [modifier]

Cette histoire amusante n'a d'autre but que de bien fixer un point important de la théorie
darwinienne.
Deux brontosaures voient un T-Rex avancer dans leur direction et se mettent à courir aussi vite qu'ils
le peuvent. Puis l'un des deux dit à l'autre :
« Pourquoi nous fatiguons-nous au juste ? Nous n'avons de toute façon pas la moindre chance d'arriver à
courir plus vite qu'un T-Rex ! »
Et l'autre lui répond cyniquement :
« Je ne cherche pas à courir plus vite que le T-Rex. Je cherche juste à courir plus vite que toi ! »
L'idée est de rappeler que le processus concerne moins une compétition entre espèces, qu'une
compétition à l'intérieur de chaque espèce. C'est à partir de ce constat et de la découverte du conflit sexuel
que Thierry Lodé[2] développe l'hypothèse que le conflit au sens large (conflit sexuel, conflit de reproduction,
coévolution) serait un puissant vecteur d'évolution, né de multiples interactions antagonistes. En fait, le rôle
des interactions et des mécanismes coévolutifs est encore probablement sous-estimé.
La sélection naturelle explique l'adaptation des espèces à leur milieu
[modifier]
Lorsqu'on observe des espèces dans leur milieu de vie, elles semblent toutes être profondément
adaptées à chacun de leur milieu : le long cou et les longues pattes de la girafe sont en effet bien adaptés
pour attraper des feuilles hautes des acacias des savanes africaines. On pourrait tout aussi bien dire que ce
sont les organismes non adaptés qui n'ont pas survécu dans ce milieu.

L'ours polaire
En outre, certaines variations avantageuses dans un environnement donné peuvent devenir néfastes
sous d'autres conditions. Par exemple, dans un milieu enneigé, une fourrure blanche permet de ne pas être
vu par ses futures proies ou ses prédateurs, mais si le milieu devient forestier et plus sombre, il n'y aura plus
de camouflage et les individus porteurs de fourrure blanche perdront leur avantage sélectif. La conséquence
de ce phénomène est donc qu'au fil des générations, par la sélection naturelle, les caractères observés dans
une population seront plus ou moins adaptés aux évolutions de son écosystème.
 Autres exemples, chez les humains la couleur de la peau est une adaptation due à la sélection
naturelle, et non à un bronzage qui se serait « fixé » à tout jamais dans certaines populations. En zones
ensoleillées les individus à la peau claire ont plus de risque de développer un cancer de la peau à cause des
rayons UV, ils sont donc désavantagés car leur espérance de vie est moindre. En zones moins ensoleillées
ces individus seraient avantagés car la lumière du soleil permet au corps de produire de la vitamine D, et de
plus le corps économise de l'énergie et des nutriments en fabriquant moins de mélanine, le pigment de la
peau.
Répartition de l'intensité de la couleur de la
Répartition de l'énergie solaire reçue.
peau humaine R. Biasutti avant 1940.

Les facteurs de l'environnement qui peuvent donc entraîner une sélection naturelle peuvent être:
 • Des facteurs physico-chimiques (le biotope) : climat, milieu occupé (terrestre, aquatique,
cavernicole…)
• D'autres êtres vivants (la biocénose) : présence de prédateurs, de parasites, de microbes, de
compétiteurs, etc.

Adaptations convergentes [modifier]

Article détaillé : convergence évolutive.
L'adaptation des espèces à leur niche écologique peut parfois conduire deux espèces qui occupent
un milieu similaire, à acquérir des ressemblances qui ne sont alors pas dues à leur éventuelle parenté. On
parle dans ce cas d'évolution convergente. Ce phénomène s'interprète comme le fait que les mêmes
contraintes du milieu mènent aux mêmes « solutions adaptatives ».
Les yeux des vertébrés et des céphalopodes constituent l'un des exemples les plus frappants de
convergence alors même que l'ancêtre commun de ces deux taxons ne possédait pas d'yeux complexes. ces
deux lignées ont évolué vers des systèmes optiques qui présentent une très forte similarité. La différence
résidant surtout dans l'orientation des cellules sensorielles dans la rétine
De tels cas de convergence évolutive sont souvent mis en avant pour argumenter en faveur d'une
conception adaptationniste de l'évolution par sélection naturelle selon laquelle l'essentiel des
caractéristiques observées dans les espèces vivantes ne sont pas dues au hasard mais sont le résultat de
diverses pressions de sélection au cours de l'histoire évolutive des espèces.
Origine des variations héréditaires dans une population [modifier]
Pour qu'il y ait sélection, encore faut-il que plusieurs variations d'un même caractère soient
présentes dans la population afin que l'individu le plus adapté l'« emporte » sur les autres. En effet, dans
l'exemple ci-dessus, si la totalité des individus sont identiques et porteurs de la variation phénotypique
« fourrure blanche », en cas de réchauffement climatique aucun individu ne pourra survivre, et l'espèce
s'éteindra. En cas de modification de l'environnement, pour qu'une espèce survive, il faut qu'elle s'adapte par
la sélection naturelle. Il est donc indispensable, qu'avant le changement du milieu elle présente en son sein
une diversité génétique importante.
L'information génétique portée par l'ADN est relativement instable [modifier]

Les mutations précèdent la sélection naturelle

La diversité génétique dans une population d'individus a pour origine des modifications de
l'information génétique dans l'ADN des cellules. Il s'agit:
1. de mutations ponctuelles dans la séquence d'ADN des gènes, ou de mutations plus
importantes comme des réarrangements chromosomiques ;
 2. de recombinaisons génétiques qui se produisent lors des transferts d'ADN des bactéries et
lors de la reproduction sexuée des eucaryotes.
On peut ajouter aux modifications du génome citées plus haut, une autre source de diversité
génétique, soient les migrations par lesquelles le stock génétique dans une population donnée se voit
renouvelé par l'arrivée d'autres membres de l'espèce porteuse d'un pool génétique différent.
Les modifications génétiques sont aléatoires : ce n'est pas l'environnement qui « dicte » quel gène
doit muter, mais bien le hasard. C'est pour cela qu'on observe dans les populations beaucoup de variations
inadaptées au milieu de vie (par exemple, les maladies génétiques rares). Une erreur courante consiste à
croire que les modifications génétiques sont une conséquence de la sélection naturelle. En revanche la
sélection naturelle a bien le pouvoir de cumuler les innovations adaptées, ce qui aboutit à des adaptations
complexes.

La sélection naturelle agit après les modifications de l'information génétique

[modifier]
C'est donc parmi la grande diversité génétique des individus, que vont ensuite être sélectionnés les
phénotypes et les allèles les plus adaptés à l'environnement. Pour en faire la démonstration on pourrait
simplement montrer que les phénotypes nouveaux étaient présents avant le changement du milieu. Ceci
n'est pas toujours évident et quand bien même ce serait le cas, un mécanisme d'induction d'une mutation
donnée par l'environnement n'est pas à exclure. C'est grâce à une astuce mathématique que Luria et
Delbrück montrent que ce sont bien les mutations préexistantes dans une population de bactéries qui sont
sélectionnées quand on ajoute un virus.
Article détaillé : Expérience de Luria et Delbrück.
L'hypothèse des caractères acquis [modifier]
Article détaillé : transmission des caractères acquis.
La génétique moléculaire n'existant pas à son époque, Darwin ne pouvait prendre en compte les
mécanismes moléculaires à l'origine des nouveaux caractères. Sa théorie de la sélection naturelle incluait
donc l'hypothèse de la transmission des caractères acquis. Dans son ouvrage de 1868, La variation des
animaux et des plantes sous l'effet de la domestication , il alla même jusqu'à proposer une théorie pour cette
transmission des caractères acquis[3]..
Dans la théorie initiale de Darwin telle qu'il l'expose dans L'Origine des espèces, ces variations entre
les individus trouvent leur origine dans le fait que des individus acquièrent des caractéristiques différentes au
cours de leur vie. Ces caractères acquis seraient alors transmis à leur descendance et cela expliquerait les
variations observées et l'évolution des caractéristiques de l'espèce. Toute autre source de variation reste
pourtant acceptable, comme par exemple le hasard; en revanche, le problème qui se poserait alors serait de
savoir comment empêcher la dilution, puis l'éventuelle disparition de ces caractères s'ils ne sont pas
"entretenus".
La théorie de l'hérédité des caractères acquis a été considérée comme invalidée par August
Weismann à la fin du XIXe siècle. En réponse aux néo-lamarckiens qui soutenaient le contraire, il montra que
des mutilations n'étaient pas transmises. On en déduisit abusivement qu'aucun caractère acquis ne pouvait
se transmettre, alors qu'une mutilation ne peut être assimilée à une acquisition par l'organisme de fonctions
nouvelles comme le voulait Lamarck. On ne peut prouver avec certitude l’impossibilité d'hérédité de
caractères acquis (une inexistence ne peut être prouvée qu'en mathématiques, par l'absurde). On peut à
défaut en chercher s'il existe quelque exemple réel réfutant cette impossibilité. Plusieurs recherches ont été
menées en ce sens au début du XXe siècle, notamment par Paul Kammerer. Cela qui fait aussi l'objet de
recherches dans le domaine de l'immunologie.
Les caractères innés sont bel et bien transmis au cours de la reproduction mais avec des variations
qui suivent les lois de l'hérédité mendélienne, du nom de leur découvreur, Gregor Mendel (1822-1884) dont
les travaux sur les lignées de pois ne furent redécouverts qu'au début du XXe siècle et étaient
malheureusement ignorés de Darwin. Mendel apporte la réponse au problème de la dilution : un caractère ne
s'affaiblit pas; il est simplement transmis en tant que dominant, transmis en tant que récessif, ou éliminé;
mais chez les individus qui le portent, il reste totalement présent, ce qui assure sa pérennité s'il est favorable
à son porteur (ou, pour être plus précis, à la descendance de son porteur).

La sélection naturelle en génétique des populations [modifier]

La convergence entre la théorie darwinienne et la théorie de l'hérédité donnera alors naissance au
cours des années 1930 à la génétique des populations, en particulier grâce aux travaux théoriques de
Ronald Fisher. À la même période, grâce aux expériences de Thomas Morgan et Theodosius Dobzhansky
sur les mouches drosophiles, les mécanismes moléculaires responsables des phénomènes d'hérédité
génétique commenceront à être identifiés. L'une des découvertes majeures de la biologie sera alors de
montrer que la diversité génétique qui garantit la variété des phénotypes est due à des modifications
aléatoires du génotype (mutations, recombinaisons génétiques, …) en particulier lors de sa transmission
d'une génération à l'autre, au moment de la reproduction.
Même si ce n'est pas le mécanisme qui avait été envisagé par Darwin dans sa théorie de la sélection
naturelle, il n'en reste pas moins que ces processus permettent de rendre parfaitement compte de la
sélection naturelle dans le cadre de ce qui est considéré comme la théorie centrale de la biologie moderne,
la théorie synthétique de l'évolution ou synthèse néo-darwinienne qui fait le lien entre les mécanismes au
niveau de la génétique moléculaire et les phénomènes d'évolution à l'échelle des populations.
Ainsi la sélection naturelle peut se "mesurer" grâce à des calculs statistiques.
Articles détaillés : génétique des populations et valeur sélective.

Cas de sélection naturelle scientifiquement démontrés [modifier]

Il apparaît aujourd'hui évident que tout organe ayant une fonction définie, par exemple la nageoire du
poisson, est une adaptation à un milieu et le résultat d'une sélection naturelle. Cependant la démonstration
scientifique doit, elle, passer par la mise en évidence d'une corrélation chiffrée entre les variations d'un
caractère héréditaire et celles d'un paramètre précis de l'environnement. Parmi les exemples les plus
célèbres, on peut citer :

Les deux morphes du phalène du

bouleau :
le clair (à gauche) et le sombre (à droite).
 • Les pinsons des Galapagos : les épisodes de sécheresse furent suivis par une raréfaction
des graines molles et donc par une augmentation de la taille du bec des pinsons leur permettant de
briser la coquille des graines restantes, plus dures.
• La sélection naturelle chez les bactéries de résistances aux virus bactériophages a été mise
en évidence par l'expérience de Luria et Delbrück. Ils ont en effet démontré pour la première fois que
les mutations préexistent à la sélection et qu'elles sont bien aléatoires.
• Le mélanisme industriel de la phalène du bouleau en Angleterre: dans cette espèce de
papillons, la proportion d'individus clairs aurait diminué du fait de la pollution qui détruisait les lichens
(blanc) qui se développaient sur les surfaces des troncs d'arbre sur lesquels ils se posaient. En effet,
Les individus clairs (initialement présents en majorité), étant désormais plus visibles que les autres
sur les troncs noir, étaient plus sujets à la prédation que les autres. Les phalènes sombres qui
existaient avant la pollution seraient alors devenues majoritaires. Puis, le phénomène se serait
inversé lorsque les industries polluantes ont progressivement disparu permettant le retour du
développement du lichen sur les troncs. En fait, cette belle histoire est discutable car apparemment
les phalènes ne se posent pas sur les troncs mais sous les feuilles des bouleaux. Les modifications
des fréquences des morphes sombres et clairs pourraient être comme des réponses à la toxicité des
rejets industriels plutôt qu'envers la modification de la couleur du revêtement des troncs. Mais
pourquoi les phalènes y seraient-elles seules sensibles ?

Sélection naturelle dans l'espèce humaine [modifier]

La sélection naturelle produit aussi ses effets dans l'espèce humaine :
• La capacité chez l'adulte à digérer le lactose du lait a été sélectionnée il y a environ 9000
ans dans les populations humaines originaires d'Europe du nord ou d'Asie centrale où on élevait du
 bétail pour son lait, mais pas dans d'autres populations où cet aliment est absent[4]. Récemment, il a
été découvert que cette capacité était présente également chez certaines ethnies d'Afrique de l'Est,
les Tutsis et Fulanis du Soudan, de Tanzanie et du Kenya, mais à partir de trois autres mutations,
sélectionnées indépendamment l'une de l'autre[5]. Ces dernières seraient apparues il y a 7000 à
3000 ans. Il s'agit là d'un exemple de convergence évolutive.
• La persistance d'une maladie génétique comme la drépanocytose est due au fait que l'allèle
responsable de la maladie confère aussi un avantage sélectif en augmentant aussi la résistance au
parasite responsable du paludisme.

Relation entre sélection naturelle et les activités humaines [modifier]

L'homme peut aussi exercer involontairement une pression de sélection sur certains organismes
dont l'évolution, en retour, peut être néfaste pour l'économie ou la santé humaine :
• La résistance aux antibiotiques : Par exemple, depuis 1961 les bactéries staphylocoques
dorés résistantes à la méticilline se sont multipliées et rendent inefficaces le traitement par cet
antibiotique dans un grand nombre de cas. Ces résistances sont à l'origine de nombreuses maladies
nosocomiales, en milieu hospitalier.
• La résistance des criquets pèlerins aux insecticides : Comme dans l'exemple précédent,
l'utilisation massive d'insecticide pour éradiquer les criquets s'est traduite par une plus grande
résistance. En effet, les individus survivant au traitement insecticide peuvent engendrer plusieurs
milliers de descendants en quelques générations, soit en quelques années, qui eux-mêmes héritent
de cette capacité de résistance. L'élimination de cette nouvelle population exige alors un traitement
encore plus agressif pour l'écosystème.
 Sélection naturelle, coévolution et compétition [modifier]
Dans les exemples précédents, il s'agit d'espèces évoluant au gré des améliorations techniques
humaines mais ce phénomène peut aussi s'observer dans les interactions biologiques entre deux espèces.
Un exemple très étudié d'une telle coévolution est le parasitisme de ponte chez les coucous[6]. Dans ces
espèces, le parent pond ses œufs dans le nid d'autres oiseaux. Dès sa naissance, le jeune coucou expulse
les œufs présents afin d'être le seul à bénéficier des soins prodigués par les parents de l'espèce hôte ainsi
trompés. Parmi les espèces parasitées, certaines ont évolué des stratégies antiparasitiques, en l'occurrence
une aptitude à distinguer les œufs de coucou de leurs propres œufs. Cela a créé une pression de sélection
pour certaines espèces de coucou qui ont évolué une forme de mimétisme des œufs de telle sorte que ceux-
ci ressemblent étonnamment aux œufs de leurs hôtes. À leur tour, les espèces parasitées (comme la pie-
grièche écorcheur) ont développé des capacités accrues de discrimination de leurs propres œufs des œufs
mimétiques du coucou, capacité qui est absente chez des espèces non ou moins parasitées.
De tels phénomènes de coévolution sont fréquemment observés dans les cas de parasitisme mais
parfois aussi dans certains cas de compétition interspécifique ou dans le cadre compétition intraspécifique
de la sélection sexuelle[7]. La coévolution inter-sexes s'observent souvent en réponse à la sélection sexuelle
post-copulatoire : par exemple, dans certaines espèces d'oiseaux[8] et d'invertébrés[9], les pénis des mâles
ont une forme hélicoïdale qui fonctionne comme un goupillon éliminant la semence des autres mâles et leur
permettant de déposer leurs propres gamètes au plus profond du vagin des femelles avec lesquelles ils
s'accouplent afin de s'assurer la paternité de la progéniture ; dans ces mêmes espèces, on observe une
augmentation proportionnelle de la taille du vagin avec une morphologie tout en sinuosités qui contrecarre
les stratégies reproductives des mâles.
La sélection artificielle [modifier]

Rose cultivée et Rose sauvage Rosa canina.

L'adjectif naturelle s'oppose chez Darwin au concept de sélection artificielle connue et pratiquée
depuis quelques milliers d'années par les éleveurs. En effet les animaux d'élevage domestiques ou les
espèces de plantes cultivées (vaches, chiens, roses…) constituent autant de variations « monstrueuses »
absentes dans la nature. Elles sont le fruit de la lente sélection d'individus intéressants (pour les rendements,
ou du point de vue esthétique) par les éleveurs et les agriculteurs (voir élevage sélectif des animaux). C'est
cette observation qui permit à Darwin d'émettre l'hypothèse d'une sélection opérée par la nature sur les
espèces sauvages.
Par exemple :
• La rose cultivée est une mutation d'une rose sauvage. Cette mutation a été sélectionnée par
les horticulteurs, elle est répandue dans les jardins: c'est la sélection artificielle.
• Dans la nature des roses cultivées ne peuvent se reproduire, c'est la sélection naturelle.
 La sélection artificielle, malgré son intense pression (élimination de tout géniteur qui ne répond pas
aux critères du choix), ne parvient pas, après des pratiques millénaires, à faire naître de nouvelles espèces.
Les races ne s'isolent pas et peuvent s'hybrider sans perte ou baisse de fécondité. La domestication et la
culture révèlent les limites, assez étroites entre lesquelles l'espèce varie sans courir de péril, mais elles
n'impriment pas un mouvement évolutif aux espèces qu'elles concernent.

La sélection utilitaire et la sélection sexuelle [modifier]

Articles détaillés : Sélection utilitaire et Sélection sexuelle.
La sélection darwinienne s'appuie sur de deux mécanismes conjoints. Le premier est la sélection
utilitaire (ou sélection de survie ou sélection écologique) le second est la sélection sexuelle. Dans le grand
public, ce dernier aspect de sélection sexuelle est souvent ignoré et on identifie la sélection naturelle avec la
sélection de survie. Or c'est une erreur car ces deux mécanismes sont bien à l'œuvre dans le monde vivant.
La sélection utilitaire correspond à un processus de tri entre individus en vertu de leur capacité à
survivre et/ou à être féconds [10]. Ce terme désigne plus spécifiquement le mécanisme qui fait évoluer les
espèces sous la pression "externe" de l'environnement ou "interne" de la compétition intraspécifique. En
effet, elle repose sur l'idée que pour pouvoir se reproduire, il faut d'abord survivre. Ce type de sélection
favorise donc les individus capables d'échapper ou de se protéger des prédateurs mais aussi de résister aux
parasites : c'est la compétition interspécifique. Il existe aussi une compétition intraspécifique : les individus
d'une même espèce étant en compétition entre eux pour trouver des ressources dans l'environnement, qu'il
s'agisse de proies ou d'autres ressources non-nutritives comme des abris (terrier, nid, …). Enfin, il y a des
facteurs dits abiotiques qui sélectionnent les individus les mieux capables de résister à l'environnement
biotopique, aux conditions climatiques, etc.
 La sélection sexuelle est un phénomène qui a lieu à une étape différente de la vie de l'individu. Elle
désigne le fait qu'il y a aussi une compétition au sein de chaque espèce pour accéder aux partenaires
sexuels dans le cadre de la reproduction sexuée. Cet aspect de la théorie fut pleinement développé par
Darwin dans son ouvrage intitulé La Descendance de l'Homme. Dans le cadre de la sélection sexuelle, il va
donc se produire une compétition intra-sexe, entre les individus d'un même sexe, mais aussi inter-sexe,
entre les sexes (les individus d'un sexe devant choisir avec quel individu de l'autre sexe ils vont s'accoupler).
La sélection sexuelle permet donc d'expliquer des caractères ou des comportements qui pénalisent la survie
quand ils sont analysés en dehors du contexte reproductif, comme la queue du paon, les bois des
mégacéros.Très importante chez les oiseaux (parades nuptiales) elle pourrait,être à l'origine des plumes
dont elle explique la forme aplatie pour exhiber les couleurs.

Pression de sélection [modifier]

Rétrospectivement, les modifications successives au cours des générations des populations peuvent
sembler orientées, comme si ces modifications étaient "tirées" ou "poussées" dans une certaine direction.
Par exemple, en suivant les observations de Darwin sur les pinsons des Galapagos, on peut observer que
certaines espèces semblent suivre une tendance vers un élargissement du bec qui devient de plus en plus
massif alors que chez d'autres espèces de pinsons, la tendance est plutôt vers un affinement du bec.
Ce phénomène qui se manifeste comme une tendance apparente dans l'évolution d'une ou plusieurs
espèces a reçu le nom de pression de sélection. Ces pressions de sélection sont 'orientées' par les
pressions dites intérieures à l'espèce (sélection sexuelle, compétition intraspécifique) et les pressions dites
extérieures à l'espèce (limitation des ressources, modifications de l'environnement, prédateurs, parasites…),
bref, tout ce qui influence la survie et la reproduction des individus.
 Les pressions de sélection s'exercent différemment d'une espèce à l'autre ou d'un milieu écologique
à un autre, voire d'une sous-population d'individus à une autre. Ainsi il peut se produire au sein d'une même
espèce une divergence si deux sous-populations sont soumises à des pressions de sélection légèrement
différentes. Ces deux populations évolueront vers des formes différentes qu'on appelle morphes et si le
phénomène se poursuit dans le temps on peut aboutir à la formation de deux espèces distinctes, c'est la
spéciation sympatrique. Les deux espèces occupent alors des niches écologiques suffisamment distinctes
pour qu'elles n'entrent plus directement en compétition l'une avec l'autre et suivent alors des "trajectoires"
évolutives différentes en réponse aux pressions de sélection spécifiques auxquelles elles sont soumises.
Article détaillé : pression de sélection.

La sélection naturelle permet l'apparition d'adaptations biologiques

complexes [modifier]

L'œil, une structure très complexe

 La myoglobine. Cette protéine, indispensable pour l'oxygénation des muscles, doit sa fonction à la
complexité de sa structure.
Les systèmes vivants apparaissent comme très complexes et sont des adaptations tellement
poussées à un milieu que les humains y trouvent une source d'innovations techniques et industrielles (par
exemple, les attaches scratch ou velcro, les industries pharmaceutique, et chimique). voir:bionique.
Cette complexité n'a pu voir le jour sans le pouvoir qu'a la sélection naturelle d'accumuler les
« bonnes » innovations génétiques :
Chaque innovation évolutive apparaît de manière aléatoire. La sélection naturelle favorise ensuite
chacun de ces petits "sauts" évolutifs (a, puis ab, puis abc… puis abcde). Elle permet ainsi l'apparition
d'adaptations de plus en plus poussées (abcde). En effet, si le caractère (a) n'avait pas été sélectionné, le
caractère (abcde) ne serait jamais apparu. Car (c) dépend de (a). Un caractère complexe, comme une
enzyme, résulte d'une accumulation d'innovations sélectionnées successivement, et non de simples
apparitions indépendantes, au hasard des innovations génétiques (même si certains "sauts" évolutifs
peuvent être plus ou moins importants ou graduels, voir équilibre ponctué). Donc la sélection naturelle ne fait
pas que favoriser les adaptations les plus complexes ; elle permet aussi leur apparition.
Cela n'est valable que si la sélection naturelle s'opère de manière continue, ce qui est envisageable
par une compétition intra-spécifique, que Malthus et Darwin estiment inévitable dans une population. En effet
les êtres vivants ont une tendance naturelle et universelle à se reproduire en plus grand nombre qu'à la
génération précédente.
Les simples innovations dues au hasard sur quelques générations ne suffisent pas à rendre compte
de la complexité des êtres vivants et de leur adaptation à leur milieu. Il faut la sélection naturelle pour
accumuler les petites innovations et pour en arriver à un organe aussi complexe que l'œil de mammifère, par
exemple.
Cela peut être une réponse aux critiques de certains néo-créationnistes, qui affirment que les
systèmes vivants (enzymes…) sont trop complexes et harmonieux pour que leur apparition ne soit due qu'à
des mutations aléatoires, et que donc selon eux il n'aurait pas eu d'évolution.

Limites de la sélection naturelle pour expliquer l'évolution de la lignée

humaine [modifier]
Comme toutes les autres espèces animales, l'espèce humaine est le produit de l'évolution et de la
sélection naturelle. Par exemple, on peut penser que la bipédie a offert dans le passé un tel avantage sélectif
aux individus capables de se déplacer debout que les gènes associés à ce mode de locomotion se sont
répandus dans toute la population humaine.
 Il reste néanmoins difficile de reconstruire le détail de l'histoire évolutive de notre espèce et en
particulier d'identifier quels facteurs exacts et quels processus précis ont pu intervenir dans l'évolution
humaine (e.g., sélection sexuelle, sélection de groupe, sélection culturelle, dérive génétique …) Dans certains
cas, il est néanmoins possible d'identifier les pressions de sélection et les adaptations résultant de la
sélection naturelle dans les populations humaines. L'un des exemples les plus documentés est la capacité
de résistance au paludisme. Les individus porteurs d'un gène entraînant une anomalie de leurs cellules
sanguines résistent mieux au parasite qui cause cette maladie. Par contre, leurs enfants courent le risque de
souffrir de drépanocytose s'ils héritent de ce gène des deux parents. L'avantage sélectif fournit par la
résistance au paludisme permet donc d'expliquer pourquoi dans les populations humaines où ce parasite est
endémique (Afrique) se maintient l'allèle d'un gène pouvant entraîner une maladie relativement grave.
L'une des tentatives les plus décriées d'appliquer la théorie darwinienne à l'espèce humaine fut
l'utilisation qu'il fut faite de la sociobiologie développée par E.O. Wilson pour expliquer le comportement des
espèces ultra-sociales (fourmis, termites, abeilles, …). En effet, même s'il est vrai que l'une des
caractéristiques de l'être humain (et aussi d'autres primates) est son mode de vie très social, la théorie de
Wilson nécessite aussi un mode de reproduction et d'organisation sociale très particulier dite eusocialité, très
différents de ce qu'on observe chez les humains. Comme s'en défend Wilson lui-même, il est donc absurde
de vouloir appliquer directement les conclusions des travaux menés sur ces espèces à l'espèce humaine.
Toutefois certains outils théoriques développés initialement dans le cadre de la sociobiologie peuvent se
révéler parfaitement pertinents pour l'étude de l'homme.
Le défi majeur de la paléoanthropologie reste néanmoins de parvenir à un cadre théorique pour
expliquer l'évolution humaine au moyen de mécanismes plus riches que la seule sélection naturelle. Or
l'importance de phénomènes comme la culture, la sélection sexuelle, la dérive génétique reste difficile à
évaluer :
 « Ainsi, même dans le cadre des théories modernes de l'évolution, qu'on appelle
néodarwinisme ou théorie synthétique de l'évolution, les évolutionnistes s'efforcent de réserver
une place à part à l'homme, étant entendu que si son corps a évolué, il reste que ce qui fait
l'humain échappe aux lois de l'évolution[11]. »

Un exemple récent d'une telle difficulté concerne le rôle de la sélection de groupe. Alors que ce
mécanisme qui "favorise la survie du groupe au détriment de la survie de l'individu" a été très critiqué dans
l'évolution animale. Il semble que l'une des particularités de l'espèce humaine est que justement, des
phénomènes de sélection multi-niveaux ont pu jouer un rôle important au cours de son évolution[12] et en
particulier dans l'évolution de sa psychologie.

Sélection naturelle, comportement et culture [modifier]

Tout comportement a une composante génétique et héréditaire. Il a été démontré que
l'environnement pouvait agir sur l'évolution d'un comportement héréditaire et inné, chez certains
animaux[13].
Quant à la culture qui ne se transmet pas par l'ADN, mais par l'apprentissage, elle peut également
être sujette à une sélection. Par exemple, si j'ai un comportement qui m'apporte de la satisfaction, comme
m'habiller à la mode, je le reproduirai et je l'enseignerai ou on m'imitera. Et inversement, ce comportement
ne sera pas transmis, si cela ne donne pas satisfaction. La théorie de la mémétique émise par Richard
Dawkins, désigne ces entités, qu'on appelle mèmes, comme éléments de base de la sélection que subit la
culture, au même titre que le sont les gènes pour l'évolution du vivant.
 Relations entre les évolutions culturelle, environnementale et génétique
Article détaillé : mémétique.
En retour la culture peut entraîner de nouvelles conditions de vie, et donc modifier la sélection
naturelle[14]. Par exemple l'utilisation de lait de vache dans l'alimentation a favorisé les génotypes tolérants
à la digestion du lactose (sucre présent dans le lait).
Ainsi, l'évolution de la culture serait le produit d'une interaction entre une sélection naturelle et une
sélection culturelle [réf. nécessaire].
Ainsi on peut envisager que ce modèle d'évolution faisant jouer des rétroactions puisse suivre un
cercle vicieux ou vertueux, ce qui entraîne une évolution perpétuelle et continue. À condition que les
rétroactions soient positives. Ce modèle est corroboré par les observations: la culture humaine suit bien une
évolution continue [réf. nécessaire], et les structures anatomiques qui ont permis cette évolution (volume
crânien, structure du cortex) ont aussi évolué de manière continue depuis 2 millions d'années, au moins.
La culture humaine, qui fait toute la singularité de notre espèce, pourrait donc être le résultat d'un tel
modèle d'évolution, avec une certaine indépendance vis-à-vis de l'environnement, si on ne tient pas compte
des modifications engendrées par les humains eux-mêmes.
E.O. Wilson parle de co-évolution[15] des gènes et de la culture. Mais cette approche est
critiquée[réf. nécessaire].
Cela est expliqué par certains scientifiques dans la théorie de la construction de niche[16]: le
comportement ou une autre activité peut influencer sur l'environnement immédiat (la niche écologique) et, en
retour, modifier la pression de sélection naturelle.

Les autres mécanismes de l'évolution [modifier]

Lorsque les fréquences de certaines variations héréditaires changent uniquement à cause du
hasard, on parle de dérive, voire (si le groupe est très réduit) d'effet fondateur. Ces caractères doivent être
relativement neutres pour la sélection naturelle (il n'y a ni avantage, ni désavantage sélectif). Si une même
évolution se produit de manière répétée dans un même milieu au cours des générations, ce n'est pas la
dérive, mais on peut parler de sélection.

Dimension universelle de la sélection naturelle [modifier]

Dans le cadre de cette théorie, tout système dans lequel s'observeraient ces trois premiers principes
donnerait lieu à un phénomène d'évolution par sélection naturelle. Dans le monde vivant, la transmission
héréditaire de l'information génétique, qui obéit à ces trois principes, résulte donc dans une évolution des
espèces par sélection naturelle. Cependant, d'un point de vue théorique, l'évolution par sélection naturelle ne
dépend pas de la nature précise des mécanismes qui permettent l'apparition de variations, la transmission
héréditaire et la traduction de l'information héréditaire en caractères phénotypiques. Le fait que Darwin lui-
même ignorait jusqu'à l'existence des gènes illustre bien le distinguo qu'il convient de faire entre le cadre
théorique de la sélection darwinienne et ses manifestations observables dans les écosystèmes terrestres.
Par conséquent, les phénomènes évolutionnaires observés dans le monde vivant pourraient tout à
fait se manifester dans d'autres systèmes qui mettraient en œuvre les trois principes fondamentaux de la
sélection darwinienne. C'est par exemple, l'hypothèse faite en exobiologie selon laquelle des formes de vie
extraterrestres pourraient être apparues sur la base de mécanismes fondamentaux différents de ceux que
l'on connait de la biologie terrestre. Au sein de telles formes de vie évoluant par sélection naturelle, on
devrait donc observer des phénomènes similaires à ceux que l'on connait sur Terre : adaptation, coévolution,
reproduction sexuée…
Les plus adaptationnistes des biologistes de l'évolution (comme Simon Conway Morris) ont ainsi proposé
que les contraintes environnementales sont suffisamment fortes et similaires pour que les formes de vie
extraterrestres devraient présenter d'importantes convergences évolutives avec la vie terrestre ; parmi
lesquelles la présence d'yeux, l'eusocialité, ou des capacités cognitives complexes[17].
Plus près de nous, l'application des principes de la sélection naturelle à la sphère culturelle humaine
a donné lieu à la mémétique qui cherche à expliquer les variations, la transmission, et la stabilisation des
phénomènes culturels par analogie avec les espèces vivantes. Dans ce cadre théorique, les mèmes sont les
unités de sélection, ou réplicateurs, des phénomènes culturels. La sélection intervient sur les mèmes en
fonction de leur capacité à "survivre" c'est-à-dire à persister dans l'esprit des individus et de leur capacité à
"se reproduire", c'est-à-dire à passer d'un individu à un autre par l'imitation, la communication,
l'enseignement, etc. À chaque reproduction, un mème peut donc "muter" : à force d'être racontée, une même
histoire sera par exemple déformée, c'est le principe du téléphone arabe. Ainsi, malgré les différences
notables entre la nature des réplicateurs biologiques et culturels, certaines analogies peuvent être
envisagées : coévolution (y compris entre gène et mème[18]) ou formation de complexes de mèmes. La
pertinence de l'application de la théorie darwinienne aux phénomènes culturels reste néanmoins très
débattue[19].
Enfin on peut aussi citer l'exemple des algorithmes évolutionnaires utilisés dans le domaine de
l'optimisation en ingénierie. Ceux-ci permettent de rechercher une solution à un problème donné en mettant
en compétition une population de solutions potentielles dont seules les meilleures sont conservées pour être
recombinées et donner naissance à une nouvelle génération de solutions.
Étant donné que dans cette méthode la sélection est le résultat d'une intervention humaine, ce cas relève
plutôt de la sélection artificielle.

Notes et références [modifier]

1. ↑ Cf. Ernst Mayr, The Growth of Biological Thought: Diversity, Evolution and Inheritance ,
Cambridge, Harvard University Press, 1982 — traduit en français sous le titre de Histoire de la
biologie. Diversité, évolution et hérédité, Fayard (1989) : 894 p. (ISBN 2213018944).
2. ↑ Thierry Lodé La guerre des sexes chez les animaux, une histoire naturelle de la sexualité.
2006, Eds Odile Jacob, Paris,ISBN 2-7381-1901-8
3. ↑ Cf. André Pichot, Histoire de la notion de vie, éd. Gallimard, coll. TEL, 1993.
4. ↑ (en) Genetic Signatures of Strong Recent Positive Selection at the Lactase
Gene [archive]. Todd Bersaglieri, Pardis C. Sabeti, Nick Patterson, Trisha Vanderploeg, Steve F.
Schaffner, Jared A. Drake, Matthew Rhodes, David E. Reich, and Joel N. Hirschhorn. The American
Journal of Human Genetics, volume 74 (2004), pages 1111–1120.
5. ↑ (en)[pdf] Convergent adaptation of human lactase persistence in Africa and
Europe [archive]. Sarah A. Tishkoff, Floyd A. Reed, Alessia Ranciaro, Benjamin F. Voight, Courtney
C. Babbitt, Jesse S. Silverman, Kweli Powell, Holly M. Mortensen, Jibril B. Hirbo, Maha Osman,
Muntaser Ibrahim, Sabah A. Omar, Godfrey Lema, Thomas B. Nyambo, Jilur Ghori, Suzannah
Bumpstead, Jonathan K. Pritchard, Gregory A. Wray & Panos Deloukas. Nature Genetics, volume 39
(January 2007), pages 31-40.
6. ↑ (en)Egg mimicry by cuckoos Cuculus canorus in relation to discrimination by hosts. M.
de L. Brooke & N. B. Davies. [archive]Nature 335, 630-632 . sur www.nature.com. Consulté le 31
mai 2010.
7. ↑ (en)Arms Races between and within Species. [archive] R Dawkins, JR Krebs.
Proceedings of the Royal Society of London. Series B (1979) 205, 1161. sur
www.journals.royalsoc.ac.uk. Consulté le 31 mai 2010.
8. ↑ Brennan PL, Prum RO, McCracken KG, Sorenson MD, Wilson RE, et al. (2007)
Coevolution of Male and Female Genital Morphology in Waterfowl. PLoS ONE 2(5): e418.
doi:10.1371/journal.pone.0000418 [archive]
9. ↑ Arnqvist G, Rowe L. (2002) Antagonistic coevolution between the sexes in a group of
insects. Nature 415: 787–789.
10.↑ E. Danchin, L.-A. Giraldeau & F. Cézilly (2005). Écologie comportementale. Dunod
11.↑ PICQ P L'humain à l'aube de l'humanité p. 41 dans Qu'est-ce que l'humain ?
12.↑ (en) Wilson, D.S. & Sober, E. (1994). Reintroducing group selection to the human
behavioral sciences. Behavioral and Brain Sciences 17 (4): 585-654. [archive] sur
www.bbsonline.org. Consulté le 31 mai 2010.
 13.↑ Boyd et Silk, L'aventure humaine, de la molécule à la culture, De Boeck 2004, partie 1,
chapitre 3, pages 68-72 ; ISBN 2 8041 4333 3
14.↑ Laland et Coolen, la culture autre moteur de l'évolution, dans "la Recherche" n°377 juillet-
août 2004
15.↑ (en)Vidéo d'une conférence sur la relation entre biologie et sciences humaines [archive]
16.↑ (en)Site des auteurs de la théorie de la construction de niche [archive]
17.↑ (en) Life's Solution Inevitable Humans in a Lonely Universe, Simon Conway Morris,
Cambridge University Press, New York, 2003. 486 pp. $30, $18.95. ISBN 0-521-82704-3.
18.↑ [pdf]On meme--gene coevolution. Bull L, Holland O, Blackmore S. Artif Life. 2000
Summer;6(3):227-35. pdf [archive]
19.↑ (en)Atran, S. (2001) The Trouble with Memes : Inference versus Imitation in Cultural
Creation. Revue scientifique. Human Nature 12(4):351-381. [archive]

Voir aussi [modifier]

Articles connexes [modifier]
• Anthropologie
• Élevage sélectif des animaux
• Historique des critiques des théories de l'évolution
• Sélection (biologie), radiation évolutive
 Bibliographie [modifier]
• Serre M, Picq P, Vincent J-D (2003) Qu’est-ce que l’Humain ?, ed. Le Pommier et le Collège
de la Cité des sciences, (ISBN 2746501309)
• Chayeux J-P (2003) Gènes et cultures - Symposium annuel, ed. Odile Jacob, 304 pages
(ISBN 2-7381-1310-9)

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive
 Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie
l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation
 • Portail de l’origine et de l’évolution du vivant
Ce document provient de « http://fr.wikipedia.org/wiki/S%C3%A9lection_naturelle ».

Catégorie : Mécanisme de l'évolution | [+]

 W000

Genetic drift
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Part of the Biology series on

 Evolution

Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

 Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
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Molecular evolution
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Biology portal · v • d • e
Genetic drift or allelic drift is the change in the relative frequency in which a gene variant (allele)
occurs in a population due to random sampling and chance: The alleles in offspring are a sample of those in
the parents, and chance has a role in determining whether a given individual survives and reproduces. A
population's allele frequency is the fraction of the gene copies that share a particular form.[1]
Genetic drift is an important evolutionary process, which leads to changes in allele frequencies over
time. It may cause gene variants to disappear completely, and thereby reduce genetic variability. In contrast
to natural selection, which makes gene variants more common or less common depending on their
reproductive success,[2] the changes due to genetic drift are not driven by environmental or adaptive
pressures, and may be beneficial, neutral, or detrimental to reproductive success.
The effect of genetic drift is larger in small populations, and smaller in large populations. Vigorous
debates wage among scientists over the relative importance of genetic drift compared with natural selection.
Ronald Fisher held the view that genetic drift plays at the most a minor role in evolution, and this remained
the dominant view for several decades. In 1968 Motoo Kimura rekindled the debate with his neutral theory of
molecular evolution which claims that most of the changes in the genetic material are caused by genetic drift.
[3]
Contents
[hide]
• 1 Analogy with marbles in a jar
• 2 Probability and allele frequency
• 2.1 Wright-Fisher model
• 2.2 Other causes than sampling error
• 3 Drift and fixation
• 3.1 Time to fixation or loss
• 4 Genetic drift versus natural selection
• 5 Population bottleneck
• 5.1 Founder effect
• 6 History of the concept
• 7 See also
• 8 Notes
• 9 External links

[edit] Analogy with marbles in a jar

The process of genetic drift can be illustrated using 20 marbles in a jar to represent 20 organisms in
a population.[4] Half of them are red and half blue, and both colors correspond to two different gene alleles in
the population. The offspring they reproduce for the next generation are represented in another jar. In each
new generation the organisms reproduce at random. To represent this reproduction, randomly select a
marble from the original jar and deposit a new marble with the same color as its "parent" in the second jar.
(The selected marble remains in the original jar.) Repeat this process until there are 20 new marbles in the
second jar. The second jar will then contain a second generation of "offspring", 20 marbles of various colors.
Unless the second jar contains exactly 10 red and 10 blue marbles, there will have been a purely random
shift in the allele frequencies.
Repeat this process a number of times, randomly reproducing each generation of marbles to form
the next. The numbers of red and blue marbles picked each generation will fluctuate: sometimes more red,
sometimes more blue. This fluctuation is genetic drift – a change in the population's allele frequency resulting
from a random variation in the distribution of alleles from one generation to the next.
It is even possible that in any one generation no marbles of a particular color will be chosen, meaning
they have no offspring. In this example, if no red marbles are selected the jar representing the new
generation will contain only blue offspring. If this happens, the red allele has been lost permanently in the
population, while the remaining blue allele has become fixed: all future generations will be entirely blue. In
small populations, fixation to a single surviving allele can occur in just a few generations. Given enough time,
this outcome is nearly inevitable for populations of any size.
 In this simulation, there is fixation in the blue "allele" within five generations.

[edit] Probability and allele frequency

Genetic drift is a change in the allele frequency within the population due to random chance: a
variation from generation to generation, resulting from random sampling. Chance events can change the
allele frequencies in a population because any individual's reproductive success can be determined by
factors other than adaptive pressures. Genetic drift occurs when these allele frequencies change as a
consequence of sampling error. In probability theory, the law of large numbers predicts little change taking
place over time when the population is large. When the reproductive population is small, however, the effects
of sampling error can alter the allele frequencies significantly. Genetic drift is therefore considered to be a
consequential mechanism of evolutionary change primarily within small, isolated breeding populations.[5]
 This effect can be illustrated using a simplified example. Consider a very large colony of bacteria
which are isolated within a drop of solution. The bacteria are genetically identical except for a single gene for
which there are two different alleles. The alleles are labeled A and B. Half of the bacteria have allele A and
the other half have allele B. Thus both A and B have allele frequency 1/2.
A and B are neutral alleles, meaning they do not affect the bacteria's ability to survive and reproduce.
This being the case, all bacteria in this colony are equally likely to survive and reproduce. The drop of
solution then shrinks until it has only enough food to sustain four bacteria. All the others die without
reproducing. Among the four who survive, there are sixteen possible combinations for the A and B alleles:
(A-A-A-A), (B-A-A-A), (A-B-A-A), (B-B-A-A),
(A-A-B-A), (B-A-B-A), (A-B-B-A), (B-B-B-A),
(A-A-A-B), (B-A-A-B), (A-B-A-B), (B-B-A-B),
(A-A-B-B), (B-A-B-B), (A-B-B-B), (B-B-B-B).
If each of the combinations with the same number of A and B respectively are counted, we get the
following table. The probabilities are calculated with the slightly faulty premise that the peak population size
was infinite.
A B Combinations Probability

4 0 1 1/16

3 1 4 4/16

2 2 6 6/16
 1 3 4 4/16

0 4 1 1/16
The probability of any one possible combination is

where 1/2 (the probability of the A or B allele for each surviving bacterium) is multiplied four times
(the total sample size, which in this example is the total number of surviving bacteria).
As seen in the table, the total number of possible combinations to have an equal (conserved) number
of A and B alleles is six, and its probability is 6/16. The total number of possible alternative combinations is
ten, and the probability of unequal number of A and B alleles is 10/16.
The total number of possible combinations can be represented as binomial coefficients and they can
be derived from Pascal's triangle. The probability for any one of the possible combinations can be calculated
with the formula
 where N is the number of bacteria and k is the number of A (or B) alleles in the combination. The
function '()' signifies the binomial coefficient and can be expressed as "N choose k". Using the formula to
calculate the probability that between them the surviving four bacteria have two A alleles and two B alleles.[6]

Genetic drift occurs when a population's allele frequencies change due to random events. In this
example the population contracted to just four random survivors, a phenomenon known as population
bottleneck. The original colony began with an equal distribution of A and B alleles but chances are that the
remaining population of four members has an unequal distribution. There is a higher probability that this
surviving population will undergo drift (10/16) than the probability it will remain the same (6/16).

[edit] Wright-Fisher model

The example given uses bacteria to illustrate the effect of sampling and probability in genetic drift.
Each bacterium has a single allele, A or B. In diploid populations consisting of N individuals there are 2N
copies of each gene. An individual can have two copies of the same allele or two different alleles. We can
call the frequency of one allele p and the frequency of the other q. One formula to calculate the probability of
obtaining k copies of the allele with frequency p in the next generation is[7]
 where the symbol "!" signifies the factorial function. This expression can also be formulated using the
binomial coefficient,

[edit] Other causes than sampling error

Random changes in allele frequencies can also be caused by other effects than sampling error, for
example random changes in selection pressure.[8]
One important alternative source of stochasticity, perhaps more important than genetic drift, is
genetic draft.[9] Genetic draft has very different mathematical properties; unlike genetic drift, the direction of
the random error is strongly autocorrelated over time.

[edit] Drift and fixation

The Hardy–Weinberg principle states that within sufficiently large, randomly mating populations the
allele frequencies will tend to remain constant from one generation to the next unless the equilibrium is
disturbed by migration, genetic mutation, or selection.[10] However, there is no residual influence on this
probability from the frequency distribution of alleles in the grandparent, or any earlier, population—only that of
the parent population. The predicted distribution of alleles among the offspring is a memory-less probability
as described in the Markov property. This means that the mathematical probabilities associated with the
distribution of alleles in any generation are only derived from the distribution of alleles in the generation
immediately prior. Thus when random fluctuations result in a change of the allele frequency from the parent
generation to the offspring generation, that deviation establishes new expected values for the allele
distributions in the next generation to follow.[11]
Populations do not gain new alleles from the random sampling of alleles passed to the next
generation, but the sampling can cause an existing allele to disappear. Because random sampling can
remove but not replace an allele, and because random declines or increases in allele frequency will influence
the expected allele distributions for the next following generation, genetic drift drives a population towards
genetic uniformity over time. When an allele reaches a frequency of 1 (100%) it is said to be "fixed" in the
population and when an allele reaches a frequency of 0 (0%) it is lost. Once an allele becomes fixed, genetic
drift comes to a halt, and the allele frequency cannot change unless a new allele is introduced in the
population via mutation or gene flow. Thus even while genetic drift is a random, directionless process, it acts
to eliminate genetic variation over time.[12]
[edit] Time to fixation or loss
 Ten simulations of random genetic drift of a single given allele with an initial frequency distribution
0.5 measured over the course of 50 generations, repeated in three reproductively synchronous populations
of different sizes. In general, alleles drift to loss or fixation (frequency of 0.0 or 1.0) significantly faster in
smaller populations.
Assuming genetic drift is the only evolutionary force acting upon an allele, at any given point in time
the probability that an allele will eventually become fixed in the population is simply its frequency in the
population at that given point in time.[13] For example, if the frequency p for allele A is 75% and the
frequency q for allele B is 25%, then given unlimited time the probability A will ultimately become fixed in the
population is 75% and the probability that B becomes fixed is 25%.
The time needed, which is measured by the number of generations, for fixation to occur can be
estimated with probability, and is proportional to the population size, such that fixation is predicted to occur
much more rapidly in smaller populations.[14] Normally the effective population size, which is smaller than
the total population, is used to determine these probabilities. The effective population size ( Ne) takes into
account factors such as the level of inbreeding, the number of organisms that are too old or young to breed,
and the lower probability that two organisms that live far apart will reproduce.[15]
One forward-looking formula used for approximating the time needed for a neutral allele to become
fixed through genetic drift is
 where T is the number of generations, Ne is the effective population size, and p is the initial
frequency for the given allele. The result is the number of generations expected to pass before fixation
occurs for a given allele in a population with given size ( Ne) and allele frequency (p).[16]

The expected time for the neutral allele to be lost through genetic drift can be calculated as[17]

These calculations are also used to understand the impact genetic drift will have on a new allele
introduced through genetic mutation. In estimating the drift of a neutral mutation in a large population, (which
can be assumed to begin as a single occurrence of the allele and thus its initial frequency will be negligible),
the formulas can be simplified to[18]

for average number of generations needed for fixation of a neutral mutation, and

for the average number of generations needed for the loss of a neutral mutation.[19] In cases where
Ne and N are assumed to be equal, the ratio of time-to-fixation over time-to-loss is[16]
[edit] Genetic drift versus natural selection
Although both processes drive evolution, genetic drift operates randomly while natural selection
functions non-randomly. This is because natural selection emblematizes the ecological interaction of a
population, whereas drift is regarded as a sampling procedure across successive generations without regard
to fitness pressures imposed by the environment. While natural selection is directioned, guiding evolution by
impelling heritable adaptations to the environment, genetic drift has no direction and is guided only by the
mathematics of chance.[20]
As a result, drift acts upon the genotypic frequencies within a population without regard their
relationship to the phenotype. Changes to the genotype caused by genetic drift may or may not result in
changes to the phenotype. In drift each allele in a population is randomly and independently affected, yet the
fluctuations in their allele frequencies are all driven in a quantitatively similar manner. Drift is blind with
respect to any advantage or disadvantage the allele may bring. Alternatively, natural selection acts directly
on the phenotype and indirectly on its underlying genotype. Selection responds specifically to the adaptive
advantage or disadvantage presented by a phenotypic trait, and thus affects genes differentially. Selection
indirectly rewards the alleles that develop adaptively advantageous phenotypes; with an increase in
reproductive success for the phenotype comes an increase in allele frequency. By the same token, selection
lowers the frequencies for alleles that cause unfavorable traits, and ignores those which are neutral.[21]
 In natural populations, genetic drift and natural selection do not act in isolation; both forces are
always at play. However, the degree to which alleles are affected by drift or selection varies according to
population size. The statistical effect of sampling error during the reproduction of alleles is much greater in
small populations than in large ones. When populations are very small, drift will predominate, and may
preserve unfavorable alleles and eliminate favorable ones (this means purifying selection has a stronger
effect in species with a larger effective population[22]). Weak selective effects may not be seen at all, as the
small changes in frequency they would produce are overshadowed by drift.[23]
In a large population, the probability of sampling error is small and little change to the allele
frequencies is expected, even over many generations. Even weak selection forces acting upon an allele will
push its frequency upwards or downwards (depending on whether the allele's influence is beneficial or
harmful). However, in cases where the allele frequency is very small, drift can also overpower selection—even
in large populations. For example, while disadvantageous mutations are usually eliminated quickly in large
populations, new advantageous mutations are almost as vulnerable to loss through genetic drift as are
neutral mutations. It is not until the allele frequency for the advantageous mutation reaches a certain
threshold that genetic drift will have little effect.[21]
Most mutations have a clear negative selective effect and cause the gametes that they occur in to
disappear after a few generations. It is possible to calculate how many percent of each generation will be
removed by such mutations. The size of the remaining population, is said to be a factor f 0, the equilibrium
frequency of non-deleterious alleles, times the total population (f 0 is between zero and one). When a neutral
mutation spreads by drift in a population, some of the occurrencies will be removed because they are linked
to such negative mutations. That is, they are located in chromosomes that are removed because of selection
against a mutation in another part of the same chromosome. As a consequence, the effective population size
is reduced by the factor f0. This means that mutation and selection in combination, causes the drift to have
more effect. Because strength of genetic linkage varies along the chromosome, effective population size, and
thereby genetic drift, also varies. With a higher recombination rate, linkage decreases and with it this local
effect on drift.[24][25] This effect is visible in molecular data as a correlation between local recombination rate
and genetic diversity,[26] and negative correlation between gene density and diversity at noncoding sites.[27]

[edit] Population bottleneck

Main article: Population bottleneck
 Changes in a population's allele frequency following a population bottleneck: the rapid and radical
decline in population size has reduced the population's genetic variation.
A population bottleneck is when a population contracts to a significantly smaller size over a short
period of time due to some random environmental event.[28] In a true population bottleneck, the odds for
survival of any member of the population are purely random, and are not improved by any particular inherent
genetic advantage. The bottleneck can result in radical changes in allele frequencies, completely
independent of selection. And its impact can be sustained, even when the bottleneck is caused by a one-time
event such as a natural catastrophe. Even when the allele frequency of the original population is carried
forward in the surviving population, a radical reduction in population size increases the likelihood of further
allele fluctuation from drift in generations to come.
A population's genetic variation can be greatly reduced by a bottleneck, and even beneficial
adaptations may be permanently eliminated.[29] The loss of variation leaves the surviving population
vulnerable to any new selection pressures such as disease, climate change or shift in the available food
source, because adapting in response to environmental changes requires sufficient genetic variation in the
population for natural selection to take place.[30][31]
There have been many known cases of population bottleneck in the recent past. Prior to the arrival of
Europeans, North American prairies were habitat for millions of greater prairie chickens. In Illinois alone, their
numbers plummeted from about 100 million birds in 1900 to about 50 birds in the 1990s. The declines in
population resulted from hunting and habitat destruction, but the random consequence has been a loss of
most of the species' genetic diversity. DNA analysis comparing birds from the mid century to birds in the
1990s documents a steep decline in the genetic variation in just in the latter few decades. Currently the
greater prairie chicken is experiencing low reproductive success.[32]
 Over-hunting also caused a severe population bottleneck in the northern elephant seal in the 19th
century. Their resulting decline in genetic variation can be deduced by comparing it to that of the southern
elephant seal which were not so aggressively hunted.[33]

[edit] Founder effect

Main article: Founder effect
 When very few members of a population migrate to form a separate new population, the founder
effect occurs. For a period after the foundation, the small population experiences intensive drift. In the figure
this results in fixation of the red allele.
The founder effect is a special case of genetic drift, occurring when a small group in a population
splinters off from the original population and forms a new one. The random sample of alleles in the just
formed new colony is expected to grossly misrepresent the original population in at least some respects.[34]
It is even possible that the number of alleles for some genes in the original population is larger than the
number of gene copies in the founders, making complete representation impossible. When a newly formed
colony is small, its founders can strongly affect the population's genetic make-up far into the future.
A well documented example is found in the Amish migration to Pennsylvania in 1744. Two members
of the new colony shared the recessive allele for Ellis–van Creveld syndrome. Members of the colony and
their descendants tend to be religious isolates and remain relatively insular. As a result of many generations
of inbreeding, Ellis-van Creveld syndrome is now much more prevalent among the Amish than in the general
population.[21][35]
The difference in gene frequencies between the original population and colony may also trigger the
two groups to diverge significantly over the course of many generations. As the difference, or genetic
distance, increases, the two separated populations may become distinct, both genetically and phenetically,
although not only genetic drift but also natural selection, gene flow and mutation will all contribute to this
divergence. This potential for relatively rapid changes in the colony's gene frequency led most scientists to
consider the founder effect (and by extension, genetic drift) a significant driving force in the evolution of new
species. Sewall Wright was the first to attach this significance to random drift and small, newly isolated
populations with his shifting balance theory of speciation.[36] Following after Wright, Ernst Mayr created
many persuasive models to show that the decline in genetic variation and small population size following the
founder effect were critically important for new species to develop.[37] However, there is much less support
for this view today since the hypothesis has been tested repeatedly through experimental research and the
results have been equivocal at best.[38]

[edit] History of the concept

The concept for genetic drift was first introduced by one of the founders in the field of population
genetics, Sewall Wright. His first use of the term "drift" was in 1929,[39] though at the time he was using it in
the sense of a directed process of change, or natural selection. Later that year he used it to refer to a purely
random process, or change due to the effects of sampling error. It came to be known as the "Sewall-Wright
effect", though he was never entirely comfortable to see his name given to it. He preferred "drifting at
random", and "drift" came to be adopted as a technical term in the stochastic sense exclusively.[40]
In the early days of the modern evolutionary synthesis, scientists were just beginning to blend the
new science of population genetics with Charles Darwin's theory of natural selection. Working within this new
framework, Wright focused on the effects of inbreeding on small relatively isolated populations. He
introduced the concept of an adaptive landscape in which phenomena such as cross breeding and genetic
drift in small populations could push them away from adaptive peaks, which would in turn allow natural
selection to push them towards new adaptive peaks.[41] Wright thought smaller populations were more
suited for natural selection because "inbreeding was sufficiently intense to create new interaction systems
through random drift but not intense enough to cause random nonadaptive fixation of genes."[40]
Wright's views on the role of genetic drift in the evolutionary scheme were controversial almost from
the very beginning. One of the most vociferous and influential critics was colleague Ronald Fisher. Fisher
conceded genetic drift played some role in evolution, but an insignificant one. Fisher has been accused of
misunderstanding Wright's views because in his criticisms Fisher seemed to argue Wright had rejected
selection almost entirely. To Fisher, viewing the process of evolution as a long, steady, adaptive progression
was the only way to explain the ever increasing complexity from simpler forms. But the debates have
continued between the "gradualists" and those who lean more toward the Wright model of evolution where
selection and drift together play an important role.[2]
In 1968,[42] population geneticist Motoo Kimura rekindled the debate with his neutral theory of
molecular evolution, which claims that most of the genetic changes are caused by genetic drift acting on
neutral mutations.[3]

[edit] See also

Evolutionary biology portal

• Allopatric speciation
• Antigenic drift
• Gene pool
• Small population size

[edit] Notes
1. ^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. Glossary. ISBN 0-
87893-189-9.
2. ^ a b Avers, Charlotte (1989). Process and Pattern in Evolution. Oxford University Press.
 3. ^ a b Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. 320. ISBN 0-
87893-189-9.
4. ^ "Evolution 101:Sampling Error and Evolution". University of California Berkeley.
http://evolution.berkeley.edu/evosite/evo101/IIID1Samplingerror.shtml. Retrieved 2009-11-01.
5. ^ Zimmer, Carl (2002). Evolution : The Triumph of an Idea. New York, NY: Perennial.
pp. 364. ISBN 0-06-095850-2.
6. ^ Walker J. "Introduction to Probability and Statistics". The RetroPsychoKinesis Project.
Fourmilab. http://www.fourmilab.ch/rpkp/experiments/statistics.html. Retrieved 2009-11-17.
7. ^ Hartl, Daniel (2007). Principles of Population Genetics. Sinauer Associates. p. 102.
ISBN 978-0-87893-308-2.
8. ^ Li, Wen-Hsiung; Dan Graur (1991). Fundamentals of Molecular Evolution. Sinauer
Associates. p. 28. ISBN 0-87893-452-9.
9. ^ Gillespie, John H. (2001). "Is the population size of a species relevant to its evolution?".
Evolution 55 (11): 2161–2169.
10.^ Hartwell el al, Leland (2004). Genetics: From Genes to Genomes; 2nd edition. McGraw Hill.
p. 680. ISBN 0-07-246248-5.
11.^ Hartl, Daniel; Andrew Clark (2007). Principles of Population Genetics. Sinauer Associates.
p. 102. ISBN 978-0-87893-308-2.
12.^ Li, Wen-Hsiung; Dan Graur (1991). Fundamentals of Molecular Evolution. Sinauer
Associates. p. 29. ISBN 0-87893-452-9.
13.^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. 300. ISBN 0-
87893-189-9.
 14.^ Otto S, Whitlock M (1 June 1997). "The probability of fixation in populations of changing
size". Genetics 146 (2): 723–33. PMID 9178020. PMC 1208011.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=9178020.
15.^ Charlesworth B (March 2009). "Fundamental concepts in genetics: Effective population
size and patterns of molecular evolution and variation". Nat. Rev. Genet. 10 (3): 195–205.
doi:10.1038/nrg2526. PMID 19204717.
16.^ a b Hedrick, Philip W. (2004). Genetics of Populations. Jones and Bartlett Publishers.
pp. 737. ISBN 0763747726.
17.^ Daniel Hartl, Andrew Clark (2007). Principles of Population Genetics, 4th edition . Sinauer
Associates. p. 112. ISBN 978-0-87893-308-2.
18.^ Wen-Hsiung Li, Dan Graur (1991). Fundamentals of Molecular Evolution. Sinauer
Associates. p. 33. ISBN 978-0-87893-452-9.
19.^ Kimura, Motoo; Ohta, Tomoko (2001). Theoretical Aspects of Population Genetics.
Princeton University Press. pp. 232. ISBN 0691080984.
20.^ "Natural Selection: How Evolution Works (An interview with Douglas Futuyma, see answer
to question Is natural selection the only mechanism of evolution?)". ActionBioscience.org.
http://www.actionbioscience.org/evolution/futuyma.html. Retrieved 2009-11-24.
21.^ a b c Cavalli-Sforza, L. L.; Menozzi, Paolo; Piazza, Alberto (1996). The history and
geography of human genes. Princeton, N.J.: Princeton University Press. pp. 413. ISBN 0-691-02905-
9.
22.^ Small KS, Brudno M, Hill MM, Sidow A (March 2007). "Extreme genomic variation in a
natural population". Proc. Natl. Acad. Sci. U.S.A. 104 (13): 5698–703. doi:10.1073/pnas.0700890104.
PMID 17372217. PMC 1838466. http://www.pnas.org/content/104/13/5698.long.
 23.^ Simpson, George Gaylord (1967). The Meaning of Evolution (Second ed.). Yale University
Press.
24.^ Golding B (1994). Non-neutral evolution: theories and molecular data . Springer. pp. 46.
ISBN 978-0-412-05391-7.
25.^ Charlesworth B, Morgan MT, Charlesworth D (August 1993). "The effect of deleterious
mutations on neutral molecular variation". Genetics 134 (4): 1289–303. PMID 8375663.
PMC 1205596. http://www.genetics.org/cgi/reprint/134/4/1289.
26.^ Presgraves DC (September 2005). "Recombination enhances protein adaptation in
Drosophila melanogaster". Curr. Biol. 15 (18): 1651–6. doi:10.1016/j.cub.2005.07.065.
PMID 16169487.
27.^ Nordborg M, Hu TT, Ishino Y, et al. (July 2005). "The pattern of polymorphism in
Arabidopsis thaliana". PLoS Biol. 3 (7): e196. doi:10.1371/journal.pbio.0030196. PMID 15907155.
28.^ Population Bottleneck | Macmillan Genetics
29.^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. pp. 303–304. ISBN 0-
87893-189-9.
30.^ O'Corry-Crowe G (2008). "Climate change and the molecular ecology of arctic marine
mammals". Ecological Applications 18 (2 Suppl): S56–S76. doi:10.1890/06-0795.1. PMID 18494363.
http://www.esajournals.org/doi/full/10.1890/06-0795.1.
31.^ Cornuet JM, Luikart G (1996). "Description and power analysis of two tests for detecting
recent population bottlenecks from allele frequency data". Genetics 144 (4): 2001–14.
PMID 8978083.
32.^ Hillis, David M.; Sadava, David E.; Heller, Craig H.; Orians, Gordon H.; Purves, William K.
(2006). "Chs. 1, 21–33, 52–57". Life: The Science of Biology. II: Evolution, Diversity and Ecology. San
Francisco: W. H. Freeman. pp. 1251. ISBN 9780716798569.
 33.^ "Evolution 101: Bottlenecks and Founder Effects". University of California, Berkeley.
http://evolution.berkeley.edu/evosite/evo101/IIID3Bottlenecks.shtml. Retrieved 2009-04-07.
34.^ Neill, Campbell (1996). Biology; Fourth edition. The Benjamin/Cummings Publishing
Company. p. 423. ISBN 0-8053-1940-9.
35.^ "Genetic Drift and the Founder Effect". Evolution. Public Broadcast System.
http://www.pbs.org/wgbh/evolution/library/06/3/l_063_03.html. Retrieved 2009-04-07.
36.^ Wade, Michael S.; Wolf, Jason; Brodie, Edmund D. (2000). Epistasis and the evolutionary
process. Oxford [Oxfordshire]: Oxford University Press. p. 330. ISBN 0-19-512806-0.
37.^ Mayr, Ernst, Jody Hey, Walter M. Fitch, Francisco José Ayala (2005). Systematics and the
Origin of Species: on Ernst Mayr's 100th anniversary (Illustrated ed.). National Academies Press.
p. 367. ISBN 9780309095365.
38.^ Howard, Daniel J.; Berlocher, Steward H. (1998). Endless Forms (Illustrated ed.). United
States: Oxford University Press. p. 470. ISBN 9780195109016.
39.^ Wright S (1929). "The evolution of dominance". The American Naturalist 63 (689): 556–61.
doi:10.1086/280290.
40.^ a b Stevenson, Joan C. (1991). Dictionary of Concepts in Physical Anthropology. Westport,
Conn: Greenwood Press. ISBN 0-313-24756-0.
41.^ Larson, Edward J. (2004). Evolution: The Remarkable History of a Scientific Theory.
Modern Library. ISBN 978-0679642886.
42.^ Kimura M (1968). "Evolutionary rate at the molecular level". Nature 217 (5129): 624–26.
doi:10.1038/217624a0. PMID 5637732.
 [edit] External links
• The TalkOrigins Archive
• Genetic drift illustrations in Barton et al.

[show]
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Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts
 Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
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Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

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History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

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 Key concepts Hardy-Weinberg law · Genetic linkage · Linkage disequilibrium · Fisher's
fundamental theorem · Neutral theory · Price equation

Selection Natural · Sexual · Artificial · Ecological

Effects of
selection Genetic hitchhiking · Background selection
on genomic variation

Genetic drift Small population size · Population bottleneck · Founder effect ·

Coalescence · Balding–Nichols model

Founders R. A. Fisher · J. B. S. Haldane · Sewall Wright

Related topics Evolution · Microevolution · Evolutionary game theory · Fitness landscape ·

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List of evolutionary biology topics

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Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
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Les mécanismes de
l'évolution biologique

Mécanismes non aléatoires :

• sélection naturelle
 • sélection
utilitaire
• sélection
sexuelle
• sélection de
parentèle
• sélection de
groupe
• sélection
stratégique
• sélection artificielle

Mécanismes aléatoires :
• mutation génétique
• recombinaison
• dérive génétique

Conséquences de l'évolution :
• spéciation
• adaptation des
espèces
• radiation évolutive
 La dérive génétique est l'évolution d'une population ou d'une espèce causée par des phénomènes
aléatoires, impossible à prévoir. Du point de vue génétique, c'est la modification de la fréquence d'un allèle,
ou d'un génotype, au sein d'une population, indépendamment des mutations, de la sélection naturelle et des
migrations. La dérive génétique est causée par des phénomènes aléatoires et imprévisibles, comme par
exemple, le hasard des rencontres des spermatozoïdes et des ovules, dans le cas d'une reproduction
sexuée.
Les effets de la dérive génétique sont d'autant plus importants que la population est petite, car les
écarts observés par rapport aux fréquences alléliques y seront d'autant plus perceptibles. Elle concerne
surtout les allèles neutres c’est-à-dire qui ne confèrent ni avantage ni désavantage sélectif, puisque on peut
espérer pouvoir ne pas y imputer les écarts observés. La dérive génétique est un des mécanismes majeurs
de l'évolution.
Sommaire
[masquer]
• 1 Principe
• 2 Effet fondateur
• 3 Effet fondateur et spéciation
• 4 Dérive génétique et érosion de la biodiversité
• 5 Notes et références
• 6 Voir aussi
• 6.1 Articles connexes
• 6.2 Liens externes

Principe [modifier]
Concrètement, dans le cadre d'une reproduction sexuée, un individu qui ne se reproduit qu'une seule
fois, ne va transmettre à son descendant que la moitié de ses allèles. C'est au cours du brassage génétique
aléatoire, lors de la méiose que vont être transmis certains allèles et pas d'autres. Pour qu'un individu puisse
transmettre à coup sûr la totalité de ses allèles, il faudrait que le nombre de descendants tendent vers l'infini.
En conséquence, dans toute population, il est statistiquement inévitable que certains allèles (chacune des
variantes d'un même gène) ne soient transmis par aucun adulte à leurs descendance. De plus, certains
individus n'ont pas de descendance du tout. Le nombre des allèles (la variabilité génétique) se réduit donc.
Parmi les allèles « survivants », certains vont voir leur fréquence originelle diminuer ou au contraire
augmenter. C'est pour cette raison que le principe de Hardy-Weinberg, repose, entre autres, sur l'hypothèse
d'une taille infinie de population. Dans une population de taille infinie, les fréquences génotypiques
observées seront égales à leur espérance, puisque l'écart potentiel des tirages à aux fréquences alléliques
(variance), est d'autant plus grand que la population est petite.
Du point de vue d'un gène, la dérive génétique conduit à l'augmentation ou la diminution de la
fréquence dans la population, de l'une de ses versions ( = allèle), deux exemples concrets, en sont un effet
de fondation et une diminution de la diversité génétique.

Effet fondateur [modifier]

Illustration de l'effet fondateur: les populations pionnières ne sont pas le reflet exact de la population
de départ.
 La population de la région québécoise du Saguenay-Lac-Saint-Jean a une particularité: elle présente
certaines maladies génétiques avec des fréquences beaucoup plus élevées qu'ailleurs. Ces maladies sont,
par exemple, l'ataxie spastique ou le Syndrome d'Andermann. Cela s'explique, entre autres, par un effet
fondateur:
Au XVIIe siècle quelques milliers d'individus colonisent la région, qui aujourd'hui compte 300 000
habitants. Il faut comprendre que suite à ce début de colonisation, peu de colons sont venus s'ajouter au
groupe fondateur qui s'est alors reproduit durant un certain nombre d'années sans apport extérieur (encore
aujourd'hui, peu de gens viennent s'installer dans cette région par manque d'emploi et elle subit donc un
exode partiel de sa population plus jeune). L'étude des généalogies ont prouvé que chacune des maladies
n'a été introduite que par un seul pionnier[1]. C'est le hasard qui a fait qu'un pionnier porteur de l'allèle
malade se trouvât dans la population fondatrice.
Lorsqu'un nombre réduit d'individus se sépare d'une population plus vaste, pour aller coloniser une
île ou un nouveau milieu, ces individus ne vont "emporter" qu'un échantillon d'allèles du pool d'allèles de la
population mère, et ce, de manière que l'on suppose aléatoire. La nouvelle population peut donc présenter
des fréquences génotypiques fort différentes de la population initiale. Cet écart peut changer radicalement le
profil (allélique, génotypique et phénotypique) de la population fondatrice, par rapport à la population initiale.

Effet fondateur et spéciation [modifier]

L'effet fondateur, peut s'il conduit à un isolement reproducteur, c'est-à-dire la cessation d'échange de
versions géniques entre l'ensemble des individus de l'espèce, aboutir à une spéciation. L'effet fondateur est
donc un puissant moteur de la spéciation allopatrique.
Dérive génétique et érosion de la biodiversité [modifier]

20 simulations par ordinateur de l'évolution de la fréquence d'un allèle, fixée au départ à 0,5.
On voit que l'effet de la dérive génétique est plus important dans la petite population.
l'Orang outan, une espèce en danger de disparition
Dans les simulations numériques ci-contre, avec une population de 10 individus, sur les 20 essais :
• 13 essais aboutissent à une disparition de l'allèle (la fréquence atteint 0),
• 5 essais mènent à une « fixation » de l'allèle qui remplace les autres allèles (la fréquence
atteint la valeur maximum 1).
• Dans les 3 autres essais tous les allèles sont conservés.
 Dans la majorité des cas (18 sur 20 cas), la dérive génétique aboutit donc à une baisse de la
diversité génétique ce qui n'est pas favorable à l'adaptation des espèces à un changement du milieu.
Dans une population plus grande (100 individus), un allèle ne se fixe que dans 2 cas sur 20
seulement. La simulation peut être reproduite grâce à ce site. On peut en conclure que, plus une population
est petite, et plus les effets de la dérive génétique sont importants, et plus la diversité génétique dans la
population sera menacée.
La dérive génétique et la perte de diversité génétique sont des phénomènes naturels, mais ils
peuvent être amplifiés par des pratiques artificielles, aboutissant à la réduction des effectifs, par la prédation
(chasse) ou une fragmentation du paysage (déforestation, utilisation agricole) ; ou encore la dépression de
consanguinité, notamment par effet Allee qui ont pour conséquence de réduire l'effectif de certaines
espèces : orang-outang[2], tigre, Centaurea corymbosa[3]…
En effet l'arrivée d'un nouveau parasite par exemple peut être « supportée » par une population si
elle est assez vaste, car il y aura une sélection naturelle des génotypes résistants. Si la population est trop
petite, la probabilité qu'il existe un allèle adapté au nouveau facteur de l'environnement est faible. Le risque
d'extinction de l'espèce sera important. C'est pourquoi il est nécessaire de maintenir les espèces à l'état
sauvage avec des effectifs significatifs, afin d'empêcher les effets délétères de la dérive génétique.

Notes et références [modifier]

1. ↑ Science et Vie, Hors-série, mars 2005 n°230
2. ↑ Genetic Evidence that Humans Have Pushed Orang-utans to the Brink of Extinction, Gross
LPLoS Biology Vol. 4, No. 2, e57
 3. ↑ Centaurea corymbosa, a cliff-dwelling species tottering on the brink of extinction: A
demographic and genetic study, Coals et al., PNAS, 1997, Vol. 94, pp. 3471-3476

Voir aussi [modifier]

Articles connexes [modifier]
• Horloge moléculaire
• Théorie neutraliste de l'évolution
• évolution (biologie)
• Théories de l'évolution
• Arbre phylogénétique
• Génétique des populations

Liens externes [modifier]

• Effectuer vos simulations, en ligne, en faisant varier les effectifs et les fréquences allèliques

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes
 Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et
Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Mammifères Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire
 évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de la biologie

• Portail de l’origine et de l’évolution du vivant

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Catégories : Génétique des populations | Mécanisme de l'évolution | [+]

 W000

Gene flow
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article needs additional citations for verification.
Please help improve this article by adding reliable references. Unsourced material may be
challenged and removed. (March 2010)

Part of the Biology series on

 Evolution

Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

 Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics

Biology portal · v • d • e
In population genetics, gene flow (also known as gene migration) is the transfer of alleles of genes
from one population to another.
 Migration into or out of a population may be responsible for a marked change in allele frequencies
(the proportion of members carrying a particular variant of a gene). Immigration may also result in the
addition of new genetic variants to the established gene pool of a particular species or population.
There are a number of factors that affect the rate of gene flow between different populations. One of
the most significant factors is mobility, as greater mobility of an individual tends to give it greater migratory
potential. Animals tend to be more mobile than plants, although pollen and seeds may be carried great
distances by animals or wind.
Maintained gene flow between two populations can also lead to a combination of the two gene pools,
reducing the genetic variation between the two groups. It is for this reason that gene flow strongly acts
against speciation, by recombining the gene pools of the groups, and thus, repairing the developing
differences in genetic variation that would have led to full speciation and creation of daughter species.
For example, if a species of grass grows on both sides of a highway, pollen is likely to be transported
from one side to the other and vice versa. If this pollen is able to fertilize the plant where it ends up and
produce viable offspring, then the alleles in the pollen have effectively been able to move from the population
on one side of the highway to the other.
Contents
[hide]
• 1 Barriers to gene flow
• 2 Gene flow in humans
• 3 Gene flow between species
• 3.1 Genetic pollution
• 4 Models of gene flow
• 5 Gene flow mitigation
• 6 See also
• 7 External links
• 8 References

[edit] Barriers to gene flow

Physical barriers to gene flow are usually, but not always, natural. They may include impassable
mountain ranges, oceans, or vast deserts. In some cases, they can be artificial, man-made barriers, such as
the Great Wall of China, which has hindered the gene flow of native plant populations[1]. Samples of the
same species which grow on either side have been shown to have developed genetic differences, because
there is little to no gene flow to provide recombination of the gene pools.
 Barriers to gene flow need not always be physical. Species can live in the same environment, yet
show very limited gene flow due to limited hybridization or hybridization yielding unfit hybrids.

[edit] Gene flow in humans

See also: African admixture in Europe
Gene flow has been observed in humans. For example, in the United States, gene flow was
observed between a white European population and a black West African population, which were recently
brought together. In West Africa, where malaria is prevalent, the Duffy antigen provides some resistance to
the disease, and this allele is thus present in nearly all of the West African population. In contrast, Europeans
have either the allele Fya or Fyb, because malaria is almost non-existent. By measuring the frequencies of
the West African and European groups, scientists found that the allele frequencies became mixed in each
population because of movement of individuals. It was also found that this gene flow between European and
West African groups is much greater in the Northern U.S. than in the South.[ citation needed]

[edit] Gene flow between species

Gene flow can occur between species, either through hybridization or gene transfer from bacteria or
virus to new hosts.
Gene transfer, defined as the movement of genetic material across species boundaries, which
includes horizontal gene transfer, antigenic shift, and reassortment is sometimes an important source of
genetic variation. Viruses can transfer genes between species [2]. Bacteria can incorporate genes from other
dead bacteria, exchange genes with living bacteria, and can exchange plasmids across species boundaries
[3]. "Sequence comparisons suggest recent horizontal transfer of many genes among diverse species
including across the boundaries of phylogenetic "domains". Thus determining the phylogenetic history of a
species can not be done conclusively by determining evolutionary trees for single genes." [4]
Biologist Gogarten suggests "the original metaphor of a tree no longer fits the data from recent
genome research". Biologists [should] instead use the metaphor of a mosaic to describe the different
histories combined in individual genomes and use the metaphor of an intertwined net to visualize the rich
exchange and cooperative effects of horizontal gene transfer. [5]
"Using single genes as phylogenetic markers, it is difficult to trace organismal phylogeny in the
presence of HGT [horizontal gene transfer]. Combining the simple coalescence model of cladogenesis with
rare HGT [horizontal gene transfer] events suggest there was no single last common ancestor that contained
all of the genes ancestral to those shared among the three domains of life. Each contemporary molecule has
its own history and traces back to an individual molecule cenancestor. However, these molecular ancestors
were likely to be present in different organisms at different times." [6]

[edit] Genetic pollution

Main article: Genetic pollution
Purebred, naturally-evolved, region-specific, wild species can be threatened with extinction in a big
way[2] through the process of genetic pollution, potentially causing uncontrolled hybridization, introgression
and genetic swamping. These processes can lead to homogenization or replacement of local genotypes as a
result of either a numerical and/or fitness advantage of introduced plant or animal[3]. Nonnative species can
bring about a form of extinction of native plants and animals by hybridization and introgression either through
purposeful introduction by humans or through habitat modification, bringing previously isolated species into
contact. These phenomena can be especially detrimental for rare species coming into contact with more
abundant ones. Interbreeding between the species can cause a 'swamping' of the rarer species' gene pool,
creating hybrids that drive the originally purebred native stock to complete extinction. The extent of this facet
of gene flow is not always apparent from morphological (outward appearance) observations alone. Some
degree of gene flow may be due to normal, evolutionarily constructive processes, and all constellations of
genes and genotypes cannot be preserved. That being said, hybridization with or without introgression may
threaten a rare species' existence nonetheless.[4][5].

[edit] Models of gene flow

Models of gene flow can be derived from population genetics, e.g. Sewall Wright's neighborhood
model, Wright's island model and the stepping stone model.

This section requires expansion.

[edit] Gene flow mitigation

This section does not cite any references or sources.
Please help improve this article by adding citations to reliable sources. Unsourced material
may be challenged and removed. (March 2010)
When cultivating genetically modified (GM) plants or livestock, it becomes necessary to prevent
"genetic pollution" i.e. their genetic modification from reaching other conventionally hybridized or wild native
plant and animal populations by using gene flow mitigation usually through unintentional cross pollination and
crossbreeding. Reasons to limit gene flow may include biosafety or agricultural co-existence, in which GM
and non-GM cropping systems work side by side.
Scientists in several large research programmes are investigating methods of limiting gene flow in
plants. Among these programmes are Transcontainer, which investigates methods for biocontainment,
SIGMEA, which focuses on the biosafety of genetically modified plants, and Co-Extra, which studies the co-
existence of GM and non-GM product chains.
Generally, there are three approaches to gene flow mitigation: keeping the genetic modification out of
the pollen, preventing the formation of pollen, and keeping the pollen inside the flower.
• The first approach requires transplastomic plants. In transplastomic plants, the modified DNA
is not situated in the cell's nucleus but is present in plastids, which are cellular compartments outside
the nucleus. An example for plastids are chloroplasts, in which photosynthesis occurs. In some
plants, the pollen does not contain plastids and, consequently, any modification located in plastids
cannot be transmitted by the pollen.
• The second approach relies on male sterile plants. Male sterile plants are unable to produce
functioning flowers and therefore cannot release viable pollen. Cytoplasmic male sterile plants are
known to produce higher yields. Therefore, researchers are trying to introduce this trait to genetically
modified crops.
• The third approach works by preventing the flowers from opening. This trait is called
cleistogamy and occurs naturally in some plants. Cleistogamous plants produce flowers which either
open only partly or not at all. However, it remains unclear how reliable cleistogamy is for gene flow
mitigation: a Co-Extra research project on rapeseed investigating the matter has published
preliminary results which cast doubt on the attainment of a high degree of reliability.
[edit] See also
• Biological dispersal
• Genetic erosion
• Genetic admixture

[edit] External links

• Co-Extra research on gene flow mitigation
• Transcontainer research on biocontainment
• SIGMEA research on the biosafety of GMOs: http://sigmea.dyndns.org

[edit] References
1. ^ Su, H et al. (2003) "The Great Wall of China: a physical barrier to gene flow?." Heredity,
Volume 9 Pages 212-219
2. ^ Hybridization and Introgression; Extinctions; from "The evolutionary impact of invasive
species; by H. A. Mooney and E. E. Cleland" Proc Natl Acad Sci U S A. 2001 May 8; 98(10): 5446–
5451. doi: 10.1073/pnas.091093398. Proc Natl Acad Sci U S A, v.98(10); May 8, 2001, The National
Academy of Sciences
3. ^ Glossary: definitions from the following publication: Aubry, C., R. Shoal and V. Erickson.
2005. Grass cultivars: their origins, development, and use on national forests and grasslands in the
Pacific Northwest. USDA Forest Service. 44 pages, plus appendices.; Native Seed Network (NSN),
Institute for Applied Ecology, 563 SW Jefferson Ave, Corvallis, OR 97333, USA
 4. ^ EXTINCTION BY HYBRIDIZATION AND INTROGRESSION; by Judith M. Rhymer ,
Department of Wildlife Ecology, University of Maine, Orono, Maine 04469, USA; and Daniel
Simberloff, Department of Biological Science, Florida State University, Tallahassee, Florida 32306,
USA; Annual Review of Ecology and Systematics, November 1996, Vol. 27, Pages 83-109 (doi:
10.1146/annurev.ecolsys.27.1.83), [1]
5. ^ Genetic Pollution from Farm Forestry using eucalypt species and hybrids; A report for the
RIRDC/L&WA/FWPRDC; Joint Venture Agroforestry Program; by Brad M. Potts, Robert C. Barbour,
Andrew B. Hingston; September 2001; RIRDC Publication No 01/114; RIRDC Project No CPF - 3A;
ISBN 0 642 58336 6; ISSN 1440-6845; Australian Government, Rural Industrial Research and
Development Corporation
• Su, H et al. (2003) "The Great Wall of China: a physical barrier to gene flow?. " Heredity,
Volume 9 Pages 212-219

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation
 Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)
Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Gene_flow"

Categories: Population genetics

• This page was last modified on 28 July 2010 at 12:59.

 W000

Flux de gènes
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Cet article est une ébauche concernant la biologie et l'origine et évolution de la

vie.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
En génétique des populations, le flux génétique, aussi nommé flux de gènes ou migration des gènes
désigne l'échange de plusieurs gènes ou de leurs allèles entre différentes populations en général
apparentées.
 C'est un mécanisme d'évolution non sexué qui agit à l'encontre du phénomène de spéciation. On le
retrouve chez les plantes, les bactéries, divers micro-organismes, les champignons[1], les moustiques[2].
En botanique par exemple, certains gènes peuvent être transférés d’une population de plantes à une
autre par pollinisation. Ainsi en agriculture, ce mécanisme explique la contamination de plantations non-
OGM par des plantations d'OGM voisines.
En microbiologie, il est possible que des bactéries s'échange du matériel génétique (par
transformation, transduction et conjugaison), ce phénomène leur permet ainsi de développer une certaine
résistance aux antibiotiques.

Sommaire
[masquer]
• 1 Notes et références
• 2 Voir aussi
• 2.1 Articles connexes
• 2.2 Liens externes

Notes et références [modifier]

1. ↑ Exemple chez les champignons : Flux de gènes au sein d'un paysage agricole chez le
champignon Mycosphaerella fijiensis [archive]
 2. ↑ Exemple chez les moustiques : Variabilité Génétique et Flux de Gènes chez les
Moustiques Anopheles arabiensis et Anopheles gambiae, Vecteurs du Paludisme en Afrique. Apport
des Loci Microsatellites. [archive]

Voir aussi [modifier]

Articles connexes [modifier]
• Théories de l'évolution
• Recombinaison génétique
• Cléistogamie

Liens externes [modifier]

• Portail de la biologie

• Portail de la biologie cellulaire et moléculaire

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Flux_de_g%C3%A8nes ».

Catégories : Génétique | Mécanisme de l'évolution | Génétique des populations | [+]

 W000

Speciation
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics

Biology portal · v • d • e
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F.
Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,'
as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages.[1][2] Whether genetic drift is a
minor or major contributor to speciation is the subject of much ongoing discussion. There are four geographic
modes of speciation in nature, based on the extent to which speciating populations are geographically
isolated from one another: allopatric, peripatric, parapatric, and sympatric. Speciation may also be induced
artificially, through animal husbandry or laboratory experiments. Observed examples of each kind of
speciation are provided throughout.[3]
Contents
[hide]
• 1 Natural speciation
• 1.1 Allopatric
• 1.2 Peripatric
• 1.3 Parapatric
• 1.4 Sympatric
• 1.4.1 Speciation via polyploidization
• 1.5 Hawthorn fly
• 1.5.1 Speciation via hybrid formation
• 1.6 Reinforcement (Wallace effect)
• 2 Artificial speciation
• 3 Genetics
• 3.1 Hybrid speciation
• 3.2 Gene transposition as a cause
• 3.3 Interspersed repeats
• 3.4 Human speciation
• 4 See also
• 5 References
• 6 Further reading
• 7 External links
[edit] Natural speciation
All forms of natural speciation have taken place over the course of evolution; however it still remains
a subject of debate as to the relative importance of each mechanism in driving biodiversity.[4]

The three-spined stickleback (Gasterosteus aculeatus)

One example of natural speciation is the diversity of the three-spined stickleback, a marine fish that,
after the last ice age, has undergone speciation into new freshwater colonies in isolated lakes and streams.
Over an estimated 10,000 generations, the sticklebacks show structural differences that are greater than
those seen between different genera of fish including variations in fins, changes in the number or size of their
bony plates, variable jaw structure, and color differences.[5]
There is debate as to the rate at which speciation events occur over geologic time. While some
evolutionary biologists claim that speciation events have remained relatively constant over time, some
palaeontologists such as Niles Eldredge and Stephen Jay Gould have argued that species usually remain
unchanged over long stretches of time, and that speciation occurs only over relatively brief intervals, a view
known as punctuated equilibrium.

[edit] Allopatric
 Comparison of allopatric, peripatric, parapatric and sympatric speciation.
Main article: allopatric speciation
During allopatric speciation, a population splits into two geographically isolated allopatric populations
(for example, by habitat fragmentation due to geographical change such as mountain building or social
change such as emigration). The isolated populations then undergo genotypic and/or phenotypic divergence
as: (a) they become subjected to dissimilar selective pressures; (b) they independently undergo genetic drift;
(c) different mutations arise in the two populations. When the populations come back into contact, they have
evolved such that they are reproductively isolated and are no longer capable of exchanging genes.
Observed instances
Island genetics, the tendency of small, isolated genetic pools to produce unusual traits, has been
observed in many circumstances, including insular dwarfism and the radical changes among certain famous
island chains, for example on Komodo. The Galápagos islands are particularly famous for their influence on
Charles Darwin. During his five weeks there he heard that Galápagos tortoises could be identified by island,
and noticed that Mockingbirds differed from one island to another, but it was only nine months later that he
reflected that such facts could show that species were changeable. When he returned to England, his
speculation on evolution deepened after experts informed him that these were separate species, not just
varieties, and famously that other differing Galápagos birds were all species of finches. Though the finches
were less important for Darwin, more recent research has shown the birds now known as Darwin's finches to
be a classic case of adaptive evolutionary radiation.[6]

[edit] Peripatric
Main article: Peripatric speciation
 In peripatric speciation, new species are formed in isolated, small peripheral populations that are
prevented from exchanging genes with the main population. It is related to the concept of a founder effect,
since small populations often undergo bottlenecks. Genetic drift is often proposed to play a significant role in
peripatric speciation.
Observed instances
• Mayr bird fauna
• The Australian bird Petroica multicolor
• Reproductive isolation occurs in populations of Drosophila subject to population
bottlenecking
The London Underground mosquito is a variant of the mosquito Culex pipiens that entered in the
London Underground in the nineteenth century. Evidence for its speciation include genetic divergence,
behavioral differences, and difficulty in mating.[7]

[edit] Parapatric
Main article: Parapatric speciation
In parapatric speciation, the zones of two diverging populations are separate but do overlap. There is
only partial separation afforded by geography, so individuals of each species may come in contact or cross
the barrier from time to time, but reduced fitness of the heterozygote leads to selection for behaviours or
mechanisms that prevent breeding between the two species.
Ecologists refer to parapatric and peripatric speciation in terms of ecological niches. A niche must be
available in order for a new species to be successful.
 Observed instances
• Ring species
• The Larus gulls form a ring species around the North Pole.
• The Ensatina salamanders, which form a ring round the Central Valley in California.
• The Greenish Warbler (Phylloscopus trochiloides), around the Himalayas.
• the grass Anthoxanthum has been known to undergo parapatric speciation in such cases as
mine contamination of an area.

[edit] Sympatric
This section needs additional citations for verification.
Please help improve this article by adding reliable references. Unsourced material may be
challenged and removed. (March 2008)
Main article: Sympatric speciation
Sympatric speciation refers to the formation of two or more descendant species from a single
ancestral species all occupying the same geographic location.
In sympatric speciation, species diverge while inhabiting the same place. Often cited examples of
sympatric speciation are found in insects that become dependent on different host plants in the same area.[8]
[9] However, the existence of sympatric speciation as a mechanism of speciation is still hotly contested.
People have argued that the evidences of sympatric speciation are in fact examples of micro-allopatric, or
heteropatric speciation. The most widely accepted example of sympatric speciation is that of the cichlids of
Lake Nabugabo in East Africa, which is thought to be due to sexual selection.
 Until recently, there has a been a dearth of hard evidence that supports this form of speciation, with a
general feeling that interbreeding would soon eliminate any genetic differences that might appear. But there
has been at least one recent study that suggests that sympatric speciation has occurred in Tennessee cave
salamanders.[10]
The three-spined sticklebacks, freshwater fishes, that have been studied by Dolph Schluter (who
received his Ph.D. for his work on Darwin's finches with Peter Grant) and his current colleagues in British
Columbia, were once thought to provide an intriguing example best explained by sympatric speciation.
Schluter and colleagues have found:
• Two different species of three-spined sticklebacks in each of five different lakes.
• a large benthic species with a large mouth that feeds on large prey in the littoral zone
• a smaller limnetic species — with a smaller mouth — that feeds on the small plankton
in open water.
• DNA analysis indicates that each lake was colonized independently, presumably by a marine
ancestor, after the last ice age.
• DNA analysis also shows that the two species in each lake are more closely related to each
other than they are to any of the species in the other lakes.
• Nevertheless, the two species in each lake are reproductively isolated; neither mates with the
other.
• However, aquarium tests showed that
• the benthic species from one lake will spawn with the benthic species from the other
lakes and
• likewise the limnetic species from the different lakes will spawn with each other.
 • These benthic and limnetic species even display their mating preferences when
presented with sticklebacks from Japanese lakes; that is, a Canadian benthic prefers a
Japanese benthic over its close limnetic cousin from its own lake.
• Their conclusion: in each lake, what began as a single population faced such competition for
limited resources that
• disruptive selection — competition favoring fishes at either extreme of body size and
mouth size over those nearer the mean — coupled with
• assortative mating — each size preferred mates like it - favored a divergence into two
subpopulations exploiting different food in different parts of the lake.
• The fact that this pattern of speciation occurred the same way on three separate
occasions suggests strongly that ecological factors in a sympatric population can cause
speciation.
However, the DNA evidence cited above is from mitochondrial DNA (mtDNA), which can often move
easily between closely related species ("introgression") when they hybridize. A more recent study,[11] using
genetic markers from the nuclear genome, shows that limnetic forms in different lakes are more closely
related to each other (and to marine lineages) than to benthic forms in the same lake. The threespine
stickleback is now usually considered an example of "double invasion" (a form of allopatric speciation) in
which repeated invasions of marine forms have subsequently differentiated into benthic and limnetic forms.
The threesspine stickleback provides an example of how molecular biogeographic studies that rely solely on
mtDNA can be misleading, and that consideration of the genealogical history of alleles from multiple unlinked
markers (i.e. nuclear genes) is necessary to infer speciation histories.
Sympatric speciation driven by ecological factors may also account for the extraordinary diversity of
crustaceans living in the depths of Siberia's Lake Baikal.
 [edit] Speciation via polyploidization

Speciation via polyploidy: A diploid cell undergoes failed meiosis, producing diploid gametes, which
self-fertilize to produce a tetraploid zygote.
Polyploidy is a mechanism often attributed to causing some speciation events in sympatry. Not all
polyploids are reproductively isolated from their parental plants, so an increase in chromosome number may
not result in the complete cessation of gene flow between the incipient polyploids and their parental diploids
(see also hybrid speciation).
Polyploidy is observed in many species of both plants and animals, and results in rapid speciation
since offspring of, for example, tetraploid x diploid matings result in triploid sterile progeny.[12] In fact, it has
been proposed that many of the existing plant and most animal species have undergone an event of
polyploidization in their evolutionary history.[ citation needed] However, reproduction is often by
parthenogenesis since polyploid animals are often sterile. [needs further editing—not true in plants]. Rare
instances of polyploid mammals are known, but most often result in prenatal death.
[edit] Hawthorn fly
One example of evolution at work is the case of the hawthorn fly, Rhagoletis pomonella, also known
as the apple maggot fly, which appears to be undergoing sympatric speciation.[13] Different populations of
hawthorn fly feed on different fruits. A distinct population emerged in North America in the 19th century some
time after apples, a non-native species, were introduced. This apple-feeding population normally feeds only
on apples and not on the historically preferred fruit of hawthorns. The current hawthorn feeding population
does not normally feed on apples. Some evidence, such as the fact that six out of thirteen allozyme loci are
different, that hawthorn flies mature later in the season and take longer to mature than apple flies; and that
there is little evidence of interbreeding (researchers have documented a 4-6% hybridization rate) suggests
that sympatric speciation is occurring. The emergence of the new hawthorn fly is an example of evolution in
progress.[14]

[edit] Speciation via hybrid formation

See Hybrid speciation section under the Genetics heading below.

[edit] Reinforcement (Wallace effect)

Reinforcement is the process by which natural selection increases reproductive isolation.[15] It may
occur after two populations of the same species are separated and then come back into contact. If their
reproductive isolation was complete, then they will have already developed into two separate incompatible
species. If their reproductive isolation is incomplete, then further mating between the populations will produce
hybrids, which may or may not be fertile. If the hybrids are infertile, or fertile but less fit than their ancestors,
then there will be no further reproductive isolation and speciation has essentially occurred (e.g., as in horses
and donkeys.) The reasoning behind this is that if the parents of the hybrid offspring each have naturally
selected traits for their own certain environments, the hybrid offspring will bear traits from both, therefore
would not fit either ecological niche as well as the parents did. The low fitness of the hybrids would cause
selection to favor assortative mating, which would control hybridization. This is sometimes called the Wallace
effect after the evolutionary biologist Alfred Russel Wallace who suggested in the late 19th century that it
might be an important factor in speciation.[16] If the hybrid offspring are more fit than their ancestors, then
the populations will merge back into the same species within the area they are in contact.
Reinforcement is required for both parapatric and sympatric speciation. Without reinforcement, the
geographic area of contact between different forms of the same species, called their "hybrid zone," will not
develop into a boundary between the different species. Hybrid zones are regions where diverged populations
meet and interbreed. Hybrid offspring are very common in these regions, which are usually created by
diverged species coming into secondary contact. Without reinforcement the two species would have
uncontrollable inbreeding. Reinforcement may be induced in artificial selection experiments as described
below.

[edit] Artificial speciation

New species have been created by domesticated animal husbandry, but the initial dates and
methods of the initiation of such species are not clear. For example, domestic sheep were created by
hybridisation, and no longer produce viable offspring with Ovis orientalis, one species from which they are
descended.[17] Domestic cattle, on the other hand, can be considered the same species as several varieties
of wild ox, gaur, yak, etc., as they readily produce fertile offspring with them.[18]
The best-documented creations of new species in the laboratory were performed in the late 1980s.
William Rice and G.W. Salt bred fruit flies, Drosophila melanogaster, using a maze with three different
choices of habitat such as light/dark and wet/dry. Each generation was placed into the maze, and the groups
of flies that came out of two of the eight exits were set apart to breed with each other in their respective
groups. After thirty-five generations, the two groups and their offspring were isolated reproductively because
of their strong habitat preferences: they mated only within the areas they preferred, and so did not mate with
flies that preferred the other areas.[19] The history of such attempts is described in Rice and Hostert (1993).
[20]
Diane Dodd was also able to show how reproductive isolation can develop from mating preferences
in Drosophila pseudoobscura fruit flies after only eight generations using different food types, starch and
maltose.[21]
 Dodd's experiment has been easy for many others to replicate, including with other kinds of fruit flies
and foods.[22]

[edit] Genetics
Few speciation genes have been found. They usually involve the reinforcement process of late
stages of speciation. In 2008 a speciation gene causing reproductive isolation was reported.[23] It causes
hybrid sterility between related subspecies.
[edit] Hybrid speciation
Main article: Hybrid speciation
Hybridization between two different species sometimes leads to a distinct phenotype. This phenotype
can also be fitter than the parental lineage and as such natural selection may then favor these individuals.
Eventually, if reproductive isolation is achieved, it may lead to a separate species. However, reproductive
isolation between hybrids and their parents is particularly difficult to achieve and thus hybrid speciation is
considered an extremely rare event. The Mariana Mallard is known to have arisen from hybrid speciation.
Hybridization without change in chromosome number is called homoploid hybrid speciation. It is
considered very rare but has been shown in Heliconius butterflies [24] and sunflowers. Polyploid speciation,
which involves changes in chromosome number, is a more common phenomenon, especially in plant
species.

[edit] Gene transposition as a cause

Theodosius Dobzhansky, who studied fruit flies in the early days of genetic research in 1930s,
speculated that parts of chromosomes that switch from one location to another might cause a species to split
into two different species. He mapped out how it might be possible for sections of chromosomes to relocate
themselves in a genome. Those mobile sections can cause sterility in inter-species hybrids, which can act as
a speciation pressure. In theory, his idea was sound, but scientists long debated whether it actually
happened in nature. Eventually a competing theory involving the gradual accumulation of mutations was
shown to occur in nature so often that geneticists largely dismissed the moving gene hypothesis.[25]
 However, 2006 research shows that jumping of a gene from one chromosome to another can
contribute to the birth of new species.[26] This validates the reproductive isolation mechanism, a key
component of speciation.[27]

[edit] Interspersed repeats

Main article: Interspersed repeat
Interspersed repetitive DNA sequences function as isolating mechanisms. These repeats protect
newly evolving gene sequences from being overwritten by gene conversion, due to the creation of non-
homologies between otherwise homologous DNA sequences. The non-homologies create barriers to gene
conversion. This barrier allows nascent novel genes to evolve without being overwritten by the progenitors of
these genes. This uncoupling allows the evolution of new genes, both within gene families and also allelic
forms of a gene. The importance is that this allows the splitting of a gene pool without requiring physical
isolation of the organisms harboring those gene sequences.

[edit] Human speciation

Humans have genetic similarities with chimpanzees and gorillas, suggesting common ancestors.
Analysis of genetic drift and recombination using a Markov model suggests humans and chimpanzees
speciated apart 4.1 million years ago.[28]
[edit] See also
Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Extinction
• Species problem
• Heteropatry
• Chronospecies
• Koinophilia

[edit] References
1. ^ Cook, O. F. 1906. Factors of species-formation. Science 23:506-507.
2. ^ Cook, O. F. 1908. Evolution without isolation. American Naturalist 42:727-731.
3. ^ Observed Instances of Speciation by Joseph Boxhorn. Retrieved 8 June 2009.
4. ^ J.M. Baker (2005). "Adaptive speciation: The role of natural selection in mechanisms of
geographic and non-geographic speciation". Studies in History and Philosophy of Biological and
Biomedical Sciences 36 (2): 303–326. doi:10.1016/j.shpsc.2005.03.005. PMID 19260194.
5. ^ Kingsley, D.M. (January 2009) "From Atoms to Traits," Scientific American, p. 57
 6. ^ Frank J. Sulloway (1982). "The Beagle collections of Darwin's finches (Geospizinae)".
Bulletin of the British Museum (Natural History) Zoology Series 43 (2): 49–58. available online
7. ^ Katharine Byrne and Richard A Nichols (1999) "Culex pipiens in London Underground
tunnels: differentiation between surface and subterranean populations"
8. ^ Feder, J. L., X. Xie, J. Rull, S. Velez, A. Forbes, B. Leung, H. Dambroski, K. E. Filchak, and
M. Aluja. 2005. Mayr, Dobzhansky, and Bush and the complexities of sympatric speciation in
Rhagoletis. Proceedings of the National Academy of Sciences, USA 1902:6573-6580
9. ^ Berlocher, S. H., and J. L. Feder. 2002. Sympatric speciation in phytophagous insects:
moving beyond controversy? Annual Review of Entomology 47:773-815
10.^ MATTHEW L. NIEMILLER, BENJAMIN M. FITZPATRICK, BRIAN T. MILLER (2008).
"Recent divergence with gene flow in Tennessee cave salamanders (Plethodontidae: Gyrinophilus)
inferred from gene genealogies". Molecular Ecology 17 (9): 2258–2275. available online
11.^ E.B. TAYLOR, J.D. McPHAIL (2000). "Historical contingency and determinism interact to
prime speciation in sticklebacks". Proceedings of the Royal Society of London Series B 267 (1460):
2375–2384. doi:10.1098/rspb.2000.1294. PMID 11133026. [1] available online
12.^ Ramsey, J., and D. W. Schemske. 1998. Pathways, mechanisms, and rates of polyploid
formation in flowering plants. Annual Review of Ecology and Systematics 29:467-501.
13.^ Feder JL, Roethele JB, Filchak K, Niedbalski J, Romero-Severson J (1 March 2003).
"Evidence for inversion polymorphism related to sympatric host race formation in the apple maggot
fly, Rhagoletis pomonella". Genetics 163 (3): 939–53. PMID 12663534. PMC 1462491.
http://www.genetics.org/cgi/pmidlookup?view=long&pmid=12663534.
14.^ Berlocher SH, Bush GL (1982). "An electrophoretic analysis of Rhagoletis (Diptera:
Tephritidae) phylogeny". Systematic Zoology 31 (2): 136–55. doi:10.2307/2413033.
http://jstor.org/stable/2413033.
 15.^ Ridley, M. (2003) "Speciation - What is the role of reinforcement in speciation?" adapted
from Evolution 3rd edition (Boston: Blackwell Science) tutorial online
16.^ Ollerton, J. "Flowering time and the Wallace Effect" (PDF). Heredity, August 2005.
http://oldweb.northampton.ac.uk/aps/env/lbrg/journals/papers/OllertonHeredityCommentary2005.pdf.
Retrieved 2007-05-22.
17.^ Hiendleder S., et al. (2002) "Molecular analysis of wild and domestic sheep questions
current nomenclature and provides evidence for domestication from two different subspecies"
Proceedings of the Royal Society B: Biological Sciences 269:893-904
18.^ Nowak, R. (1999) Walker's Mammals of the World 6th ed. (Baltimore: Johns Hopkins
University Press)
19.^ Rice, W.R. and G.W. Salt (1988). "Speciation via disruptive selection on habitat preference:
experimental evidence". The American Naturalist 131: 911–917. doi:10.1086/284831.
20.^ W.R. Rice and E.E. Hostert (1993). "Laboratory experiments on speciation: What have we
learned in forty years?". Evolution 47 (6): 1637–1653. doi:10.2307/2410209.
http://jstor.org/stable/2410209.
21.^ Dodd, D.M.B. (1989) "Reproductive isolation as a consequence of adaptive divergence in
Drosophila pseudoobscura." Evolution 43:1308–1311.
22.^ Kirkpatrick, M. and V. Ravigné (2002) "Speciation by Natural and Sexual Selection: Models
and Experiments" The American Naturalist 159:S22–S35 DOI
23.^ http://www.sciencemag.org/cgi/content/short/323/5912/376
24.^ Mavarez, J.; Salazar, C.A., Bermingham, E., Salcedo, C., Jiggins, C.D. , Linares, M.
(2006). "Speciation by hybridization in Heliconius butterflies". Nature 441 (7095): 868.
doi:10.1038/nature04738. PMID 16778888.
25.^ University of Rochester Press Releases
 26.^ Masly, John P., Corbin D. Jones, Mohamed A. F. Noor, John Locke, and H. Allen Orr
(September 2006). "Gene Transposition as a Cause of Hybrid Sterility in Drosophila". Science 313
(5792): 1448–1450. doi:10.1126/science.1128721. PMID 16960009.
http://www.sciencemag.org/cgi/content/short/313/5792/1448. Retrieved 2007-03-18.
27.^ Minkel, J.R. (September 8, 2006) "Wandering Fly Gene Supports New Model of Speciation"
Science News
28.^ Hobolth A, Christensen OF, Mailund T, Schierup MH (2007) "Genomic Relationships and
Speciation Times of Human, Chimpanzee, and Gorilla Inferred from a Coalescent Hidden Markov
Model." PLoS Genet 3(2): e7 (doi:10.1371/journal.pgen.0030007)

[edit] Further reading

• Coyne, J. A. & Orr, H. A. (2004). Speciation. Sunderlands, Massachusetts: Sinauer
Associates, Inc. ISBN 0-87893-089-2.
• Grant, V. (1981). Plant Speciation (2nd Edit. ed.). New York: Columbia University Press.
ISBN 0-231-05113-1.
• Mayr, E. (1963). Animal Species and Evolution. Harvard University Press. ISBN 0-674-
03750-2
• Marko, P. B. (2008). Allopatry. Oxford: Elsevier. ISBN 978-0-444-52033-3.
• White, M. J. D. (1978). Modes of Speciation. San Francisco, California: W. H. Freeman and
Company. ISBN 0-716-70284-3.
• Dedicated issue of Philosophical Transactions B on Speciation in microorganisms is freely
available.
 [edit] External links
• Observed Instances of Speciation from the Talk.Origins Frequently Asked Questions
• Speciation, and
• Evidence for Speciation from Understanding Evolution by the University of California
Museum of Paleontology
• Speciation from John Hawks' Anthropology Weblog - paleoanthropology, genetics, and
evolution
• Speciation in the context of evolution

[hide]
v•d•e
Speciation

Basic concepts Species · Cline · Chronospecies · Speciation

Modes of Allopatric · Heteropatric · Peripatric · Parapatric · Sympatric ·

speciation Polyploidy · Paleopolyploidy

Auxiliary
mechanisms Sexual selection · Assortative mating · Punctuated equilibrium
 Intermediate stages Hybrid · Species complex · Ring species · Haldane's rule

[show]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Speciation"

Categories: Ecology | Evolutionary biology | Speciation

• This page was last modified on 7 August 2010 at 15:26.
 W000

Spéciation
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Cet article ne cite pas suffisamment ses sources (janvier 2009).
Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
bas de page. (Modifier l'article)
Les mécanismes de
l'évolution biologique
Mécanismes non aléatoires :
• sélection naturelle
• sélection
utilitaire
• sélection
sexuelle
• sélection de
parentèle
• sélection de
groupe
• sélection
stratégique
• sélection artificielle

Mécanismes aléatoires :
• mutation génétique
• recombinaison
• dérive génétique

Conséquences de l'évolution :
• spéciation
• adaptation des
espèces
• radiation évolutive

Comparaison des différents modes de spéciation.

 La spéciation est, en biologie, le processus évolutif par lequel de nouvelles espèces vivantes
apparaissent. En chimie, la spéciation désigne l'ensemble des espèces chimiques sous lesquelles un
élément se présente dans un environnement donné, cf. spéciation chimique. L'article traite ici de la
spéciation en biologie.
Une espèce n'apparaît pas instantanément par une mutation conduisant à l'apparition d'un individu
d'un type nouveau. Les espèces s'individualisent à partir de populations appartenant à une espèce d'origine
(sauf pour le cas de certains végétaux, voir plus bas). Il s'agit donc de l'évolution de populations
interfécondes - composant une même espèce, par définition - en populations non-interfécondes, c'est-à-dire
isolées sur le plan reproducteur. Elles prennent ainsi le statut d'espèces biologiques vraies. La spéciation
résulte de la sélection naturelle et/ou de la dérive génétique, qui sont les deux moteurs de l'évolution. Le
concept de spéciation a été essentiellement développé par Ernst Mayr.
On distingue plusieurs types de spéciation, basés sur la répartition géographique des populations en
divergence au cours du processus :
Sommaire
[masquer]
• 1 Spéciation allopatrique
• 2 Spéciation sympatrique
• 3 Vitesse de spéciation
• 4 Notes et références
• 5 Voir aussi
• 5.1 Articles connexes

Spéciation allopatrique [modifier]

Selon ce mode de spéciation, des populations initialement interfécondes évoluent en espèces
distinctes car elles sont isolées géographiquement. C'est le mode de spéciation de loin le plus fréquent chez
les animaux. Il se décline en trois modalités :
• Spéciation vicariante : une barrière géographique (rivière, montagne, vallée, océan,
glacier...) coupe l'aire de répartition d'une espèce en plusieurs zones. Dans chacune des zones,
chaque population évolue indépendamment des autres, pouvant donner naissance à une nouvelle
espèce.
• Spéciation péripatrique ou spéciation par effet fondateur : un petit nombre d'individus fonde
une nouvelle population en marge de l'aire de répartition de l'espèce d'origine, par exemple suite à la
 colonisation d'une île près de la côte. Cette nouvelle population de petite taille peut évoluer
rapidement en une nouvelle espèce.
• Spéciation parapatrique : des populations en divergence ne sont pas totalement isolées
géographiquement mais possèdent une zone de contact étroite. Les migrations entre populations
sont cependant limitées puisque ces dernières se perpétuent dans des conditions
environnementales différentes (gradient climatique par exemple). La sélection naturelle a donc un
rôle important dans ce mode de spéciation.
Ces populations peuvent aussi revenir en sympatrie (contact secondaire), et peuvent même donner
une descendance si les évolutions n'ont pas été trop importantes. Dans ce cas, il peut tout de même y avoir
une spéciation s'il y a une faiblesse hybride, c'est-à-dire que la survie/reproduction de l'hybride est plus faible
(même légèrement) que celle des deux populations. Dans ce cas les allèles conférant la capacité de
reproduction avec les individus de l'autre population seront donc désavantagés et disparaîtront. On parle
alors de mécanismes de renforcement qui apparaissent dès qu'il y a une amorce de reproduction (faiblesse
hybride).
Les espèces Panthera leo et Panthera tigris illustrent parfaitement ce concept. Actuellement, ce sont
deux espèces différentes, elles ne peuvent se reproduire l'une avec l'autre qu'en captivité. Lors du contact
secondaire, les lions et les tigres pouvaient donc encore se reproduire, mais les tigrons et les ligres (hybride)
présentent une faiblesse (système immunitaire défaillant pour le tigron...). Les lions se reproduisant avec des
tigres ont donc eu moins de descendants et ceux-ci ont progressivement disparu. Actuellement aucun lion ne
se reproduit avec un tigre en conditions naturelles. Il s'agit bien d'une spéciation.
Dans l'espèce Canis familiaris, on comprendra aisément que la fécondation réussie de femelles
chihuahua par des mâles saint-Bernard pose une double difficulté exclusivement physique :
 • la possibilité pratique de l'accouplement,
• celui d'une arrivée de la gestation à bon terme,
même si les gamètes sont compatibles in vitro : la raciation commencerait dans ce cas précis - sans
l'existence d'intermédiaires - son cheminement vers une possible spéciation.
A contrario, certaines races de Canis lupus sont interfécondes avec Canis familiaris et les
croisements donnent une descendance féconde. Les notions de race et d'espèce contiennent donc parfois
une réduction simplificatrice de la complexité du vivant. (voir Classification). Il faut noter qu'actuellement cet
exemple ne semble plus pertinent, les chiens ( Canis familiaris) étant considérés comme une sous-espèce de
Canis lupus sous le nom de Canis lupus familiaris.

Spéciation sympatrique [modifier]

Des populations non isolées géographiquement peuvent évoluer en espèces distinctes. Ici, la
sélection naturelle joue un rôle crucial dans la divergence des populations. Ce phénomène reste
controversé, mais semble prouvé chez plusieurs poissons (notamment les cichlidés), ou chez certains
insectes phytophages. La différenciation des espèces pourrait aussi résulter du conflit sexuel.
Variante : chez certains végétaux supérieurs (angiospermes), deux espèces normalement non-
interfécondes peuvent engendrer des hybrides féconds, sous certaines conditions liées au doublement du
nombre de chromosomes présent dans l'hybride formé (diploïdie). Il peut alors lui-même se perpétuer tout en
étant incapable de se reproduire avec des individus appartenant aux espèces de ses parents. C'est donc le
premier représentant d'une nouvelle espèce, sans faire intervenir un isolement géographique. On parle donc
là aussi de spéciation sympatrique, qui semble très fréquente chez les espèces herbacées.
 Un exemple plus rare de spéciation sympatrique est la divergence entre "résident" et "transient" chez
l'Orque dans le pacifique[1]. Les deux formes habitent le même océan mais ne chassent pas les mêmes
proies, n'ont pas les mêmes émissions vocales et ne se reproduisent pas entre eux. Des divergences entre
les espèces peuvent aussi provenir des contraintes de micro-habitats contrastés. Ainsi, le putois présente un
phénotype rare dit dark qui est directement dépendant des ruisseaux boisés et établit une convergence avec
la physionomie du très rare vison d'Europe.[2]

Vitesse de spéciation [modifier]

L’histoire de l’évolution montre que les spéciations nécessitent généralement plusieurs centaines de
milliers d’années. Toutefois, certaines nouvelles espèces de souris comme celles de Madère n'ont nécessité
qu'entre 500 et 1000 ans pour apparaître (en fonction de l’origine viking ou portugaise qu’on leur attribue) ce
qui représente environ 2000 à 4000 générations soit une vitesse extrêmement rapide.
En opposition avec cette vision ("gradualisme phylétique"), la théorie des équilibres ponctués
(Stephen Jay Gould, Niles Eldredge) postule que cette histoire comprendrait de longues périodes d'équilibre,
ponctuées de brèves périodes de changements importants comme la spéciation ou les extinctions.

Notes et références [modifier]

1. ↑ Burden et al.: Resident And Transient-Type Killer Wales SC/56/SM15. [archive]
2. ↑ Thierry Lodé "Genetic divergence without spatial isolation in polecat Mustela putorius
populations". J Evol Biol 14:228-236, 2001
 Voir aussi [modifier]
Articles connexes [modifier]
• Évolution
• Écologie du paysage
• Fragmentation écopaysagère
• Insularisation écologique
• Variation clinale

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive
 Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·
naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens
 Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des
Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Sp%C3%A9ciation ».

Catégorie : Processus évolutif | [+]

 W000

Adaptation
From Wikipedia, the free encyclopedia

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This article is about the evolutionary process. For other uses, see Adaptation (disambiguation).
Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
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Introduction
Evidence
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History
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Biology portal · v • d • e
Adaptation is the evolutionary process whereby a population becomes better suited to its habitat.[1]
[2] This process takes place over many generations,[3] and is one of the basic phenomena of biology.[4]
The term adaptation may also refer to a feature which is especially important for an organism's
survival.[5] For example, the adaptation of horses' teeth to the grinding of grass, or their ability to run fast and
escape predators. Such adaptations are produced in a variable population by the better suited forms
reproducing more successfully, that is, by natural selection.
Contents
[hide]
• 1 General principles
• 1.1 Definitions
• 1.2 Adaptedness and fitness
• 2 Brief history
• 3 Types of adaptation
• 3.1 Changes in habitat
• 3.1.1 Habitat tracking
• 3.1.2 Genetic change
• 3.2 Intimate relationships: co-adaptations
• 3.2.1 Mimicry
• 3.3 The basic machinery: internal adaptations
• 3.4 Compromise and conflict between adaptations
• 4 Shifts in function
• 4.1 Pre-adaptations
• 4.2 Co-option of existing traits: exaptation
• 5 Related issues
• 5.1 Non-adaptive traits
• 5.1.1 Fitness landscapes; drift
• 5.1.2 Vestigial organs
[edit] General principles
The significance of an adaptation can only be understood in relation to the total biology of the
species. Julian Huxley[6]

Adaptation is, first of all, a process, rather than a physical part of a body.[7] The distinction may be
seen in an internal parasite (such as a fluke), where the bodily structure is greatly simplified, but nevertheless
the organism is highly adapted to its unusual environment. From this we see that adaptation is not just a
matter of visible traits: in such parasites critical adaptations take place in the life-cycle, which is often quite
complex.[8] However, as a practical term, adaptation is often used for the product: those features of a
species which result from the process. Many aspects of an animal or plant can be correctly called
adaptations, though there are always some features whose function is in doubt. By using the term adaptation
for the evolutionary process, and adaptive trait for the bodily part or function (the product), the two senses of
the word may be distinguished.
Adaptation may be seen as one aspect of a two-stage process. First, there is speciation (species-
splitting or cladogenesis), caused by geographical isolation or some other mechanism.[9][10] Second, there
follows adaptation, driven by natural selection. Something like this must have happened with Darwin's
finches, and there are many other examples. The present favourite is the evolution of cichlid fish in African
lakes, where the question of reproductive isolation is much more complex.[11][12]
Another great principle is that an organism must be viable at all stages of its development and at all
stages of its evolution. This is obviously true, and it follows that there are constraints on the evolution of
development, behaviour and structure of organisms. The main constraint, over which there has been much
debate, is the requirement that changes in the system during evolution should be relatively small changes,
because the body systems are so complex and interlinked. This is a sound principle, though there may be
rare exceptions: polyploidy in plants is common,[13] and the symbiosis of micro-organisms that formed the
eukaryota is a more exotic example.[14]
All adaptations help organisms survive in their ecological niches.[15] These adaptative traits may be
structural, behavioral or physiological. Structural adaptations are physical features of an organism (shape,
body covering, armament; and also the internal organization). Behavioural adaptations are composed of
inherited behaviour chains and/or the ability to learn: behaviours may be inherited in detail (instincts), or a
tendency for learning may be inherited (see neuropsychology). Examples: searching for food, mating,
vocalizations. Physiological adaptations permit the organism to perform special functions (for instance,
making venom, secreting slime, phototropism); but also more general functions such as growth and
development, temperature regulation, ionic balance and other aspects of homeostasis. Adaptation, then,
affects all aspects of the life of an organism.

[edit] Definitions
The following definitions are mainly due to Theodosius Dobzhansky.
1. Adaptation is the evolutionary process whereby an organism becomes better able to live in
its habitat or habitats.[16]
2. Adaptedness is the state of being adapted: the degree to which an organism is able to live
and reproduce in a given set of habitats.[17]
3. An adaptive trait is an aspect of the developmental pattern of the organism which enables or
enhances the probability of that organism surviving and reproducing.[18]
[edit] Adaptedness and fitness
Main article: Fitness (biology)
From the above definitions, it is clear that there is a relationship between adaptedness and fitness (a
key population genetics concept). Fitness is an estimate and a predictor of the rate of natural selection. What
natural selection does is change the relative frequencies of alternative phenotypes, insofar as they are
heritable.[19] Although the two are connected, the one does not imply the other: a phenotype with high
adaptedness may not have high fitness. Dobzhansky mentioned the example of the Californian redwood,
which is highly adapted, but a relic species in danger of extinction.[16] Elliott Sober commented that
adaptation was a retrospective concept since it implied something about the history of a trait, whereas fitness
predicts a trait's future.[20]
1. Fitness. The degree of demographic difference among phenotypes. Usually a relative
measure: the average contribution to a breeding population by a phenotype or a class of phenotypes.
This is also known as Darwinian fitness, relative fitness, selective coefficient, and other terms.
2. Adaptedness. Usually an absolute measure: the average absolute contribution to the
breeding population by a carrier of a phenotype or a class of phenotypes. Also known as absolute
fitness, and as the Malthusian parameter when applied to species as a whole.[21]

[edit] Brief history

Main article: History of evolutionary thought
 Adaptation as a fact of life has been accepted by all the great thinkers who have tackled the world of
living organisms. It is their explanations of how adaptation arises that separates these thinkers. A few of the
most significant ideas:[22]
• Empedocles did not believe that adaptation required a final cause (~ purpose), but "came
about naturally, since such things survived". Aristotle, however, did believe in final causes.
• In natural theology, adaptation was interpreted as the work of a deity, even as evidence for
the existence of God.[23] William Paley believed that organisms were perfectly adapted to the lives
they lead, an argument that shadowed Leibniz, who had argued that God had brought about the best
of all possible worlds. Voltaire's Dr Pangloss[24] is a parody of this optimistic idea, and Hume also
argued against design.[25] The Bridgewater Treatises are a product of natural theology, though
some of the authors managed to present their work in a fairly neutral manner. The series was
lampooned by Robert Knox, who held quasi-evolutionary views, as the Bilgewater Treatises. Darwin
broke with the tradition by emphasising the flaws and limitations which occurred in the animal and
plant worlds.[26]
 Lamarck
• Lamarck. His is a proto-evolutionary theory of the inheritance of acquired traits, whose main
purpose is to explain adaptations by natural means.[27] He proposed a tendency for organisms to
become more complex, moving up a ladder of progress, plus "the influence of circumstances",
usually expressed as use and disuse. His evolutionary ideas, and those of Geoffroy, fail because
they cannot be reconciled with heredity. This was known even before Mendel by medical men
interested in human races (Wells, Lawrence), and especially by Weismann.
Many other students of natural history, such as Buffon, accepted adaptation, and some also
accepted evolution, without voicing their opinions as to the mechanism. This illustrates the real merit of
Darwin and Wallace, and secondary figures such as Bates, for putting forward a mechanism whose
significance had only been glimpsed previously. A century later, experimental field studies and breeding
experiments by such as Ford and Dobzhansky produced evidence that natural selection was not only the
'engine' behind adaptation, but was a much stronger force than had previously been thought.[28][29][30]

[edit] Types of adaptation

Adaptation is the heart and soul of evolution. Niles Eldredge[31]

[edit] Changes in habitat

Before Darwin, adaptation was seen as a fixed relationship between an organism and its habitat. It
was not appreciated that as the climate changed, so did the habitat; and as the habitat changed, so did the
biota. Also, habitats are subject to changes in their biota: for example, invasions of species from other areas.
The relative numbers of species in a given habitat are always changing. Change is the rule, though much
depends on the speed and degree of the change.
When the habitat changes, three main things may happen to a resident population: habitat tracking,
genetic change or extinction. In fact, all three things may occur in sequence. Of these three effects, only
genetic change brings about adaptation.

[edit] Habitat tracking

When a habitat changes, the most common thing to happen is that the resident population moves to
another locale which suits it; this is the typical response of flying insects or oceanic organisms, who have
wide (though not unlimited) opportunity for movement.[32] This common response is called habitat tracking. It
is one explanation put forward for the periods of apparent stasis in the fossil record (the punctuated
equilibrium thesis).[33]

[edit] Genetic change

Genetic change is what occurs in a population when natural selection acts on the genetic variability
of the population. By this means, the population adapts genetically to its circumstances.[34] Genetic changes
may result in visible structures, or may adjust physiological activity in a way that suits the changed habitat.
It is now clear that habitats and biota do frequently change. Therefore, it follows that the process of
adaptation is never finally complete.[35] Over time, it may happen that the environment changes little, and
the species comes to fit its surroundings better and better. On the other hand, it may happen that changes in
the environment occur relatively rapidly, and then the species becomes less and less well adapted. Seen like
this, adaptation is a genetic tracking process, which goes on all the time to some extent, but especially when
the population cannot or does not move to another, less hostile area. Also, to a greater or lesser extent, the
process affects every species in a particular ecosystem.[36][37]
Van Valen thought that even in a stable environment, competing species had to constantly adapt to
maintain their relative standing. This became known as the Red Queen hypothesis.

[edit] Intimate relationships: co-adaptations

Main article: Co-adaptation
In co-evolution, where the existence of one species is tightly bound up with the life of another
species, new or 'improved' adaptations which occur in one species are often followed by the appearance and
spread of corresponding features in the other species. There are many examples of this; the idea
emphasises that the life and death of living things is intimately connected, not just with the physical
environment, but with the life of other species. These relationships are intrinsically dynamic, and may
continue on a trajectory for millions of years, as has the relationship between flowering plants and insects
(pollination).
Pollinator constancy: these honeybees selectively visit flowers from only one species, as can be
seen by the colour of the pollen in their baskets:

• Co-extinction
• Infection-resistance
• Mimicry
• Mutualism
• Parasite-host
• Pollination syndrome
• Predator-prey
• Symbiosis
 The gut contents, wing structures, and mouthpart morphologies of fossilized beetles and flies
suggest that they acted as early pollinators. The association between beetles and angiosperms during the
early Cretaceous period led to parallel radiations of angiosperms and insects into the late Cretaceous. The
evolution of nectaries in late Cretaceous flowers signals the beginning of the mutualism between
hymenopterans and angiosperms.[38]

[edit] Mimicry
Main article: Mimicry
 A and B show real wasps; the rest are mimics: three hoverflies and one beetle.
Henry Walter Bates' work on Amazonian butterflies led him to develop the first scientific account of
mimicry, especially the kind of mimicry which bears his name: Batesian mimicry.[39] This is the mimicry by a
palatable species of an unpalatable or noxious species. A common example seen in temperate gardens is
the hover-fly, many of which – though bearing no sting – mimic the warning colouration of hymenoptera
(wasps and bees). Such mimicry does not need to be perfect to improve the survival of the palatable species.
[40]
Bates, Wallace and Müller believed that Batesian and Müllerian mimicry provided evidence for the
action of natural selection, a view which is now standard amongst biologists.[41] All aspects of this situation
can be, and have been, the subject of research.[42] Field and experimental work on these ideas continues to
this day; the topic connects strongly to speciation, genetics and development.[43]
• More on mimicry: Warning Colour and Mimicry Lecture outline from University College
London

[edit] The basic machinery: internal adaptations

There are some important adaptations to do with the overall coordination of the systems in the body.
Such adaptations may have significant consequences. Examples, in vertebrates, would be temperature
regulation, or improvements in brain function, or an effective immune system. An example in plants would be
the development of the reproductive system in flowering plants.[44] Such adaptations may make the clade
(monophyletic group) more viable in a wide range of habitats. The acquisition of such major adaptations has
often served as the spark for adaptive radiation, and huge success for long periods of time for a whole group
of animals or plants.

[edit] Compromise and conflict between adaptations

It is a profound truth that Nature does not know best; that genetical evolution... is a story of
waste, makeshift, compromise and blunder. Peter Medawar[45]
 All adaptations have a downside: horse legs are great for running on grass, but they can't scratch
their backs; mammals' hair helps temperature, but offers a niche for ectoparasites; the only flying penguins
do is under water. Adaptations serving different functions may be mutually destructive. Compromise and
make-shift occur widely, not perfection. Selection pressures pull in different directions, and the adaptation
that results is some kind of compromise.[46]
Since the phenotype as a whole is the target of selection, it is impossible to improve
simultaneously all aspects of the phenotype to the same degree. Ernst Mayr.[47]
Consider the antlers of the Irish elk, (often supposed to be far too large; in deer antler size has an
allometric relationship to body size). Obviously antlers serve positively for defence against predators, and to
score victories in the annual rut. But they are costly in terms of resource. Their size during the last glacial
period presumably depended on the relative gain and loss of reproductive capacity in the population of elks
during that time.[48] Another example: camouflage to avoid detection is destroyed when vivid colors are
displayed at mating time. Here the risk to life is counterbalanced by the necessity for reproduction.
 An Indian Peacock's train
in full display
The peacock's ornamental train (grown anew in time for each mating season) is a famous adaptation.
It must reduce his maneuverability and flight, and is hugely conspicuous; also, its growth costs food
resources. Darwin's explanation of its advantage was in terms of sexual selection: "it depends on the
advantage which certain individuals have over other individuals of the same sex and species, in exclusive
relation to reproduction."[49] The kind of sexual selection represented by the peacock is called 'mate choice',
with an implication that the process selects the more fit over the less fit, and so has survival value.[50] The
recognition of sexual selection was for a long time in abeyance, but has been rehabilitated.[51] In practice,
the blue peafowl Pavo cristatus is a pretty successful species, with a big natural range in India, so the overall
outcome of their mating system is quite viable.
The conflict between the size of the human foetal brain at birth, (which cannot be larger than about
400ccs, else it will not get through the mother's pelvis) and the size needed for an adult brain (about
1400ccs), means the brain of a newborn child is quite immature. The most vital things in human life
(locomotion, speech) just have to wait while the brain grows and matures. That is the result of the birth
compromise. Much of the problem comes from our upright bipedal stance, without which our pelvis could be
shaped more suitably for birth. Neanderthals had a similar problem.[52][53][54]

[edit] Shifts in function

Adaptation and function are two aspects of one problem. Julian Huxley[55]
[edit] Pre-adaptations
This occurs when a species or population has characteristics which (by chance) are suited for
conditions which have not yet arisen. For example, the polyploid rice-grass Spartina townsendii is better
adapted than either of its parent species to their own habitat of saline marsh and mud-flats.[56] White
Leghorn fowl are markedly more resistant to vitamin B deficiency than other breeds.[57] On a plentiful diet
there is no difference, but on a restricted diet this preadaptation could be decisive.
Pre-adaptation may occur because a natural population carries a huge quantity of genetic variability.
[58] In diploid eukaryotes, this is a consequence of the system of sexual reproduction, where mutant alleles
get partially shielded, for example, by the selective advantage of heterozygotes. Micro-organisms, with their
huge populations, also carry a great deal of genetic variability.
The first experimental evidence of the pre-adaptive nature of genetic variants in micro-organisms
was provided by Salvador Luria and Max Delbrück who developed fluctuation analysis, a method to show the
random fluctuation of pre-existing genetic changes that conferred resistance to phage in the bacterium
Escherichia coli.

[edit] Co-option of existing traits: exaptation

Main article: Exaptation
The classic example is the ear ossicles of mammals, which we know from palaeontological and
embrological studies originated in the upper and lower jaws and the hyoid of their Synapsid ancestors, and
further back still were part of the gill arches of early fish.[59][60] We owe this esoteric knowledge to the
comparative anatomists, who, a century ago, were at the cutting edge of evolutionary studies.[61] The word
exaptation was coined to cover these shifts in function, which are surprisingly common in evolutionary
history.[62] The origin of wings from feathers that were originally used for temperature regulation is a more
recent discovery (see feathered dinosaurs).

[edit] Related issues

[edit] Non-adaptive traits
Some traits appear to be not adaptive, that is, selectively neutral. There may be various causes: the
utility of a trait is lost and does not now appear adaptive; the utility of a trait is unknown; the trait is a
consequence of another trait that is adaptive (i.e. spandrels). Because genes have pleiotropic effects, not all
traits may be functional. Of course, a trait may have been adaptive at some point in an organism's
evolutionary history, but habitats change, leading to adaptations becoming redundant or even a hindrance
(maladaptations). Such adaptations are termed vestigial. So, the utility of adaptations may ebb and flow.

[edit] Fitness landscapes; drift

Main article: Fitness landscape
Main article: Genetic drift
Sewall Wright's explanation for evolutionary stasis was that organisms come to occupy adaptive
peaks. In order to evolve to another, higher peak, the species would first have to pass through a valley of
maladaptive intermediate stages. This could happen by genetic drift if the population were small enough.
This was Wright's shifting balance theory of evolution.[63] There has been much skepticism among
evolutionary biologists as to whether these rather delicate conditions hold often in natural populations.[58]
Ronald Fisher felt that most populations in nature were too large for these effects of genetic drift to be
important.[28]

[edit] Vestigial organs

Main article: Vestigiality
Many organisms have vestigial organs, which are the remnants of fully functional structures in their
ancestors. As a result of changes in lifestyle the organs became redundant, and are either not functional or
reduced in functionality. With the loss of function goes the loss of positive selection, and the subsequent
accumulation of deleterious mutations. Since any structure represents some kind of cost to the general
economy of the body, an advantage may accrue from their elimination once they are not functional.
Examples: wisdom teeth in humans; the loss of pigment and functional eyes in cave fauna; the loss of
structure in endoparasites.[64]

[edit] Extinction
Main article: Extinction
If a population cannot move or change sufficiently to preserve its long-term viability, then obviously, it
will become extinct, at least in that locale. The species may or may not survive in other locales. Species
extinction occurs when the death rate over the entire species (population, gene pool ...) exceeds the birth
rate for a long enough period for the species to disappear. It was an observation of Van Valen that groups of
species tend to have a characteristic and fairly regular rate of extinction.[65]
 [edit] Co-extinction
Main article: Co-extinction
Just as we have co-adaptation, there is also co-extinction. Co-extinction refers to the loss of a
species due to the extinction of another; for example, the extinction of parasitic insects following the loss of
their hosts. Co-extinction can also occur when a flowering plant loses its pollinator, or through the disruption
of a food chain.[66] "Species co-extinction is a manifestation of the interconnectedness of organisms in
complex ecosystems ... While co-extinction may not be the most important cause of species extinctions, it is
certainly an insidious one".[67]

[edit] Flexibility, acclimatization, learning

Flexibility deals with the relative capacity of an organism to maintain themselves in different habitats:
their degree of specialization. Acclimatization is a term used for automatic physiological adjustments during
life; learning is the term used for improvement in behavioral performance during life. In biology these terms
are preferred, not adaptation, for changes during life which improve the performance of individuals. These
adjustments are not inherited genetically by the next generation.
Adaptation, on the other hand, occurs over many generations; it is a gradual process caused by
natural selection which changes the genetic make-up of a population so the collective performs better in its
niche.
 [edit] Flexibility
Populations differ in their phenotypic plasticity, which is the ability of an organism with a given
genotype to change its phenotype in response to changes in its habitat, or to its move to a different habitat.
[68][69]
To a greater or lesser extent, all living things can adjust to circumstances. The degree of flexibility is
inherited, and varies to some extent between individuals. A highly specialized animal or plant lives only in a
well-defined habitat, eats a specific type of food, and cannot survive if its needs are not met. Many herbivores
are like this; extreme examples are koalas which depend on eucalyptus, and pandas which require bamboo.
A generalist, on the other hand, eats a range of food, and can survive in many different conditions. Examples
are humans, rats, crabs and many carnivores. The tendency to behave in a specialized or exploratory
manner is inherited – it is an adaptation.
Rather different is developmental flexibility: "An animal or plant is developmentally flexible if when it
is raised or transferred to new conditions it develops so that it is better fitted to survive in the new
circumstances".[70] Once again, there are huge differences between species, and the capacities to be
flexible are inherited.

[edit] Acclimatization
Main article: Acclimatization
If humans move to a higher altitude, respiration and physical exertion become a problem, but after
spending time in high altitude conditions they acclimatize to the pressure by increasing production of red
blood corpuscles. The ability to acclimatize is an adaptation, but not the acclimatization itself. Fecundity goes
down, but deaths from some tropical diseases also goes down.
Over a longer period of time, some people will reproduce better at these high altitudes than others.
They will contribute more heavily to later generations. Gradually the whole population becomes adapted to
the new conditions. This we know takes place, because the performance of long-term communities at higher
altitude is significantly better than the performance of new arrivals, even when the new arrivals have had time
to make physiological adjustments.[71]
Some kinds of acclimatization happen so rapidly that they are better called reflexes. The rapid colour
changes in some flatfish, cephalopods, chameleons are examples.[72]

[edit] Learning
Social learning is supreme for humans, and is possible for quite a few mammals and birds: of course,
that does not involve genetic transmission except to the extent that the capacities are inherited. Similarly, the
capacity to learn is an inherited adaptation, but not what is learnt; the capacity for human speech is inherited,
but not the details of language.

[edit] Function and teleonomy

Adaptation raises some issues concerning how biologists use key terms such as function.
[edit] Function
To say something has a function is to say something about what it does for the organism, obviously.
It also says something about its history: how it has come about. A heart pumps blood: that is its function. It
also emits sound, which is just an ancillary side-effect. That is not its function. The heart has a history (which
may be well or poorly understood), and that history is about how natural selection formed and maintained the
heart as a pump. Every aspect of an organism that has a function has a history. Now, an adaptation must
have a functional history: therefore we expect it must have undergone selection caused by relative survival in
its habitat. It would be quite wrong to use the word adaptation about a trait which arose as a by-product.[73]
[74]
It is widely regarded as unprofessional for a biologist to say something like "A wing is for flying",
although that is their normal function. A biologist would be conscious that sometime in the remote past
feathers on a small dinosaur had the function of retaining heat, and that later many wings were not used for
flying (e.g. penguins, ostriches). So, the biologist would rather say that the wings on a bird or an insect
usually had the function of aiding flight. That would carry the connotation of being an adaptation with a history
of evolution by natural selection.

[edit] Teleonomy
Main article: Teleonomy
Teleonomy is a term invented to describe the study of goal-directed functions which are not guided
by the conscious forethought of man or any supernatural entity. It is contrasted with Aristotle's teleology,
which has connotations of intention, purpose and foresight. Evolution is teleonomic; adaptation hoards
hindsight rather than foresight. The following is a definition for its use in biology:
 Teleonomy: The hypothesis that adaptations arise without the existence of a prior purpose, but
by the action of natural selection on genetic variability.[75]
The term may have been suggested by Colin Pittendrigh in 1958;[76] it grew out of cybernetics and
self-organising systems. Ernst Mayr, George C. Williams and Jacques Monod picked up the term and used it
in evolutionary biology.[77][78][79][80]
Philosophers of science have also commented on the term. Ernest Nagel analysed the concept of
goal-directedness in biology;[81] and David Hull commented on the use of teleology and teleonomy by
biologists:
Haldane can be found remarking, "Teleology is like a mistress to a biologist: he cannot live
without her but he’s unwilling to be seen with her in public". Today the mistress has become a lawfully
wedded wife. Biologists no longer feel obligated to apologize for their use of teleological language;
they flaunt it. The only concession which they make to its disreputable past is to rename it ‘teleonomy’.
[82]

[edit] See also

Evolutionary biology portal

• Adaptive radiation
• Co-adaptation
• Co-evolution
• Ecological trap
 • Exaptation
• Intragenomic conflict
• Maladaptation
• Mimicry
• Polymorphism (biology)

[edit] References
1. ^ The Oxford Dictionary of Science defines adaptation as "Any change in the structure or
functioning of an organism that makes it better suited to its environment".
2. ^ Bowler P.J. 2003. Evolution: the history of an idea. California. p10
3. ^ Patterson C. 1999. Evolution. Natural History Museum, London. p1
4. ^ Williams, George C. 1966. Adaptation and natural selection: a critique of some current
evolutionary thought. Princeton. "Evolutionary adaptation is a phenomenon of pervasive importance
in biology." p5
5. ^ Both uses of the term 'adaptation' are recognized by King R.C. Stansfield W.D. and
Mulligan P. 2006. A dictionary of genetics. Oxford, 7th ed.
6. ^ Huxley, Julian 1942. Evolution the modern synthesis. Allen & Unwin, London. p449
7. ^ Mayr, Ernst 1982. The growth of biological thought. Harvard. p483: "Adaptation... could no
longer be considered a static condition, a product of a creative past, and became instead a
continuing dynamic process."
8. ^ Price P.W. 1980. The evolutionary biology of parasites. Princeton.
9. ^ Mayr E. 1963. Animal species and evolution. Harvard.
 10.^ Mayr, Ernst 1982. The growth of biological thought: diversity, evolution and inheritance .
Harvard. p562–566
11.^ Salzburger W., Mack T., Verheyen E., Meyer A. (2005). "Out of Tanganyika: Genesis,
explosive speciation, key-innovations and phylogeography of the haplochromine cichlid fishes"
(PDF). BMC Evolutionary Biology 5 (1): 17. doi:10.1186/1471-2148-5-17. PMID 15723698.
PMC 554777. http://www.biomedcentral.com/content/pdf/1471-2148-5-17.pdf.
12.^ Kornfield, Irv; Smith, Peter (November 2000). "African Cichlid Fishes: Model Systems for
Evolutionary Biology". Annual Review of Ecology and Systematics 31: 163.
doi:10.1146/annurev.ecolsys.31.1.163.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.ecolsys.31.1.163.
13.^ Stebbins, G. Ledyard, Jr. 1950. Variation and evolution in plants. Columbia. Polyploidy,
chapters 8 and 9.
14.^ Margulis, Lynn (ed) 1991. Symbiosis as a source of evolutionary innovation: speciation and
morphogenesis MIT. ISBN 0-262-13269-9
15.^ Hutchinson G. Evelyn 1965. The ecological theatre and the evolutionary play. Yale. The
niche is the central concept in evolutionary ecology; see especially part II The niche: an abstractly
inhabited hypervolume. p26–78
16.^ a b Dobzhansky T. 1968. On some fundamental concepts of evolutionary biology.
Evolutionary biology 2, 1–34.
17.^ Dobzhansky T. 1970. Genetics of the evolutionary process. Columbia, N.Y. p4–6, 79–82, 84–
87
18.^ Dobzhansky T. 1956. Genetics of natural populations XXV. Genetic changes in populations
of Drosophila pseudoobscura and Drosphila persimilis in some locations in California. Evolution 10,
82–92.
 19.^ Endler, John A. 1986. Natural selection in the wild. Princeton. p33–51: 'Fitness and
adaptation'.
20.^ Sober, Elliott 1984. The nature of selection: a philosophical enquiry . M.I.T.
21.^ following discussion in Endler, John A. 1986. Natural selection in the wild. Princeton. p33–
51: 'Fitness and adaptation'.
22.^ references and details in their articles
23.^ Desmond, Adrian 1989. The politics of evolution. Chicago. p31/32, footnote 18.
24.^ In Candide, ou l'optimisme.
25.^ Sober, Elliott 1993. Philosophy of biology. Oxford. Chapter 2
26.^ Darwin, Charles. 1872. The origin of species. 6th edition, p397: Rudimentary, atrophied
and aborted organs.
27.^ see, for example, the discussion in Bowler, Peter H. 2003. Evolution: the history of an idea.
3rd ed, California. p86–95, especially "Whatever the true nature of Lamark's theory, it was his
mechanism of adaptation that caught the attention of later naturalists". (p90)
28.^ a b Provine, William 1986. Sewall Wright and evolutionary biology. University of Chicago
Press.
29.^ Ford E.B. 1975. Ecological genetics, 4th ed. Chapman and Hall, London.
30.^ Orr H. 2005. The genetic theory of adaptation: a brief history. Nature Rev. Genetics 6, 2,
p119–127.
31.^ Eldredge, Niles 1995. Reinventing Darwin: the great evolutionary debate . Wiley N.Y. p33
32.^ Eldredge, Niles 1986. Time frames: the rethinking of Darwinian evolution and the theory of
punctuated equilibria. p136, Of glaciers and beetles.
33.^ Eldredge, Niles 1995. Reinventing Darwin: the great evolutionary debate . Wiley, N.Y. p64
 34.^ Orr H. 2005. The genetic theory of adaptation: a brief history. Nature Rev. Genetics, 6, 119–
127.
35.^ Mayr, Ernst 1982. The growth of biological thought: diversity, evolution and inheritance.
Harvard. Harvard. p481 (and sequence) tells how Darwin's ideas on adaptation developed as he
came to appreciate it as "a continuing dynamic process" (bottom p483).
36.^ Sterelny K. & Griffiths P.E. 1999. Sex and death: an introduction to philosophy of biology .
University of Chicago Press. p217 ISBN O-226-77304-3
37.^ Freeman S. & Herron J.C. 2007. Evolutionary analysis. Pearson Education. p364 ISBN 0-
13-227584-8
38.^ Stebbins, G. Ledyard, Jr. 1974. Flowering plants: evolution above the species level.
Harvard.
39.^ Carpenter GDH and Ford EB 1933. Mimicry. Methuen, London.
40.^ Wickler W. 1968. Mimicry in plants and animals. World University Library, London.
41.^ Moon H.P. 1976. Henry Walter Bates FRS 1825-1892: explorer, scientist and darwinian .
Leicestershire Museums, Leicester.
42.^ Ruxton GD, Sherratt TN and Speed MP 2004. Avoiding attack: the evolutionary ecology of
crypsis, warning signals and mimicry. Oxford.
43.^ Mallet, James 2001. The speciation revolution. J Evolutionary Biology 14, 887-8.
44.^ Stebbins, G. Ledyard, Jr. 1974. Flowering plants: evolution above the species level.
Harvard. Contains an extensive analysis of the evolution of adaptations in the radiation of
Angiosperms.
45.^ Medawar, Peter 1960. The future of Man. Methuen, London.
46.^ Jacob, Francois 1977. Evolution and tinkering. Science 196 1161–1166.
 47.^ Mayr, Ernst 1982. The growth of biological thought: diversity, evolution and inheritance .
Harvard. p589
48.^ It is, of course, not possible to test selective pressures on extinct populations in any direct
way. GOULD, STEPHEN J. (1974): Origin and function of 'bizarre' structures - antler size and skull size in
'Irish Elk', Megaloceros giganteus. Evolution 28(2): 191-220. doi:10.2307/2407322 (First page text)
49.^ Darwin, Charles 1871. The Descent of Man and selection in relation to sex. Murray,
London.
50.^ The case was treated by Fisher R.A. 1930. Genetical theory of natural selection. Oxford.
p134–139.
51.^ Cronin, Helen 1991. The ant and the peacock: altruism and sexual selection from Darwin to
the present day. Cambridge.
52.^ Rosenberg K.R. 2005. The evolution of modern human childbirth. Am J. Physical
Anthropology 35, p89–124.
53.^ Friedlander, Nancy & Jordan, David K. 1995. Obstetric implications of Neanderthal
robusticity and bone density. Human Evolution (Florence) 9: 331-342.
54.^ Miller, Geoffrey 2007. Brain evolution. In Gangestad S.W. and Simpson J.A. (eds) The
evolution of mind: fundamental questions and controversies . Guildford.
55.^ Huxley, Julian 1942. Evolution the modern synthesis. Allen & Unwin, London. p417
56.^ Huskins C.L. 1931. The origin of Spartina townsendii. Genetica 12, 531.
57.^ Lamoreux W.F and Hutt F.B. 1939. Breed differences in resistance to a deficiency in
vitamin B1 in the fowl. J. Agric. Res. Washington 58, 307–315.
58.^ a b [Dobzhansky T.] 1981. Dobzhansky's genetics of natural populations. eds Lewontin RC,
Moore JA, Provine WB and Wallace B. Columbia University Press N.Y.
 59.^ Egdar F. Allin and James A. Hopson 1992. Evolution of the auditory system in Synapsida
("Mammal-like reptiles" and primitive mammals) as seen in the fossil record. Section IV (Mammals),
Chapter 28, pages 587-614 in The evolutionary biology of hearing edited by Douglas B. Webster,
Richard R. Fay, and Arthur N. Popper. Springer-Verlag. ISBN 0-387-97588-8.
60.^ Neil Shubin 2008. Your Inner Fish: a journey into the 3.5-billion-year history of the human
body Pantheon Books 2008. ISBN 978-0-375-42447-2. Chapter 10, "Ears"
61.^ Panchen, Alec. 1992. Classification, evolution and the nature of biology . Cambridge.
Chapter 4 Homology and the evidence for evolution.
62.^ Gould, Stephen Jay and Elizabeth S. Vrba 1982. Exaptation – a missing term in the science
of form. Paleobiology 8, 1, 4–15.
63.^ Wright, Sewall 1932. The roles of mutation, inbreeding, crossbreeding, and selection in
evolution. In Proceedings of the Sixth International Congress on Genetics , p355–366.
64.^ Charles Darwin was the first to put forward such ideas: Barrett P.H. (ed) 1987. Charles
Darwin's notebooks (1836–1844). Cambridge.
65.^ Van Valen L. 1973. A new evolutionary law. Evolutionary Theory 1, 1–30.
66.^ Darwin in the Origin of Species tells the story of "a web of complex relations" involving
heartsease (Viola tricolor), red clover (Trifolium pratense, humble-bees (bumblebees), mice and cats.
Origin, 6th edition, p57.
67.^ Koh, Lian Pih. 2004. Science, 305, 5690, 1632-1634, 10 September 2004.
68.^ Price TD, Qvarnström A & Irwin DE 2003. The role of phenotypic plasticity in driving genetic
evolution. Proc. Biol. Sci. 270 p1433–1440.
69.^ Price T.D. 2006. Phenotypic plasticity, sexual selection and the evolution of colour patterns.
J Exp Biol. 209 p2368–2376
70.^ Maynard Smith J. 1993. The theory of evolution. Cambridge. 3rd ed, p33.
 71.^ Moore Lorna G. and Regensteiner Judith G. 1983. Adaptation to high altitude. Ann. Rev.
Anthropology 12, p285–304.
72.^ Maynard Smith uses the term physiologically versatile for such animals. Maynard Smith J.
1993. The theory of evolution. Cambridge. 3rd ed, p32.
73.^ Sober, Elliott 1993. Philosophy of biology. Oxford. p85–86
74.^ Williams, George C. 1966. Adaptation and natural selection: a critique of some current
evolutionary thought. Princeton. p8–10
75.^ "The hypothesis that adaptations arise without the existence of a prior purpose, but by
chance may change the fitness of an organism." Oxford Dictionary of Zoology. But one might
question the word chance, since natural selection, by its operation in particular habitats, is not a
random process (it may be a stochastic or probabilistic process, however).
76.^ Pittendrigh C.S. 1958. Adaptation, natural selection and behavior. In A. Roe and George
Gaylord Simpson (eds) Behavior and evolution. Yale.
77.^ Mayr, Ernst 1965. Cause and effect in biology. In D. Lerner (ed) Cause and effect. Free
Press, New York. p33–50.
78.^ Mayr, Ernst 1988. Toward a new philosophy of biology. Chapter 3 "The multiple meanings
of teleological".
79.^ Williams, George C. 1966. Adaptation and natural selection; a critique of some current
evolutionary thought. Chapter 9. Princeton.
80.^ Monod, Jacques 1971. Chance and necessity: an essay on the natural philosophy of
modern biology. Knopf, New York. ISBN 0-394-46615-2
81.^ Nagel, E. 1977. Teleology revisited: goal-directed processes in biology. Journal of
Philosophy 74: 261–301.
82.^ Hull D. L. 1981. Philosophy and biology. In G. Fløistad (ed) Philosophy of Science Nijhoff.
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Adaptation (biologie)
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 L’adaptation est, en biologie de l'évolution, la modification d'un caractère anatomique, d'un
processus physiologique ou d'un trait comportemental dans une population d'individus sous l'effet de la
sélection naturelle, le nouvel état de ce caractère améliorant la survie et le succès reproductif des individus
qui en sont porteurs. Le terme d'adaptation peut désigner le processus ou le résultat du processus ; il est
parfois utilisé comme synonyme de « sélection ». Intuitivement, l'adaptation peut être vue comme
l'adéquation de l'être vivant à son milieu naturel ou de l'organe à sa fonction.
La notion d'adaptation est au cœur de la théorie de l'évolution par sélection naturelle inventée par
Charles Darwin (sous le terme anglais fitness). Elle est parfois critiquée pour être tautologique : l'adaptation
est à la fois ce qui conditionne la sélection des individus dans leur milieu et le résultat de cette sélection.

Sommaire
[masquer]
• 1 Histoire du concept
• 2 Adaptation
• 3 Notes et références
• 4 Bibliographie

Histoire du concept [modifier]

Jusqu'au XIXe siècle, les naturalistes n'employaient pas le terme d'adaptation ; ils lui préféraient les
mots de "convenance" et d'"harmonie". Ce dernier reflète les idées finalistes qui, alors étaient très
majoritaires. Et de fait, c'est pour échapper à la téléologie voyant une finalité (divine ou non) dans la nature
que le concept d'adaptation a été développé portant le débat aux frontières de la métaphysique.
C'est pourquoi Cuénot, un des premiers darwinien français écrivait en ouverture de son ouvrage
L'Adaptation (1925) :
« L'adaptation est une effrayante question. […] Une adaptation est en réalité la solution d'un
problème, exactement comme une machine ou un outil fabriqués par l'homme. […]
Reconnaître les adaptations en tant que faits n'est pas très difficile : c'est question de critique,
d'observations ou d'expériences bien conduites ; mais ensuite l'esprit demande impérieusement à
comprendre le mécanisme par lequel les êtres vivants ont été pourvus de ces adaptations. Depuis les
premiers philosophes grecs les explications se sont succédé, causes finales de l'école aristotélienne,
réaction utile de l'être au milieu de Lamarck, sélection naturelle de Darwin, etc. ; assurément, tout le
monde est d'accord, maintenant, pour rechercher aux adaptations une explication causale, dans le
domaine de l'investigation scientifique, mais même si nous connaissions une loi générale qui en rendît
un compte satisfaisant, comme on l'a cru longtemps pour la théorie darwinienne, il se poserait encore
une question suprême, que l'Homme ne peut éluder. Pourquoi cette loi générale ? pourquoi tout se
passe-t-il comme si la Nature voulait la perpétuation de la Vie ? pourquoi cette finalité spéciale que la
Vie impose à la Matière ? Par ces questions, nous entrons dans le domaine de la métaphysique. » pp.
3-4.
Cuénot n'a pas prétendu apporter une réponse à cette épineuse question, il se contente de signaler
son existence. Toutefois, l'école néo-darwinienne se défend d'être finaliste et assure que l'adaptation, bien
qu'elle ressemble à un phénomène dirigé, est exempte de toute téléologie. Le jeu des forces naturelles qui
interviennent dans la sélection naturelle suffit à en rendre un compte exact et précis.
 Évidemment, quand un être vit, prospère et se perpétue dans un milieu donné, sa structure et ses
fonctions sont telles qu'elles permettent la vie ; autrement dit, il n'existe pas de désaccord entre elles et le
milieu extérieur. Cette approximation autorise, à elle seule, à affirmer qu'il existe un minimum d'adaptation
entre l'être organisé et son milieu. Considérons la faune d'un biotope limité, une mare, une plage marine,
etc., nous voyons que les animaux qui la composent appartiennent à des types d'organisation très variés.
Des solutions tout à fait différentes permettent donc l'ajustement de l'être vivant à son milieu et
l'épanouissement de la vie. L'adaptation est rarement une notion ayant une valeur absolue ; elle présente
toujours un caractère relatif.
Cuénot distingue trois types d'adaptations successives :
1. l'accommodation ou adaptation ponctuelle de l'individu à un milieu,
2. l'acclimatation ou adaptation d'un groupe établi de manière durable dans un milieu,
3. la naturalisation ou l'adaptation de l'espèce à un milieu où elle s'est établie de manière
définitive.
En outre, il considère aussi l'adaptation statistique ou adaptation physiologique et éthologique qui se
traduit par une convergence des formes (par exemple, le requin et le dauphin) des organismes vivant dans
des milieux semblables ou des organes (par exemple, l'œil chez la pieuvre et chez les mammifères) ayant en
charge de remplir la même fonction, mais appartenant à des lignées différentes.
Il met aussi en évidence les Limites de l'adaptation, notamment à travers les Organes inutiles, les
Organes utilisés mais non nécessaires, ou encore les Organes mal faits et les fonctions nuisibles que sont
par exemples les Organes hypertéliques, c'est-à-dire démesurés et encombrants. Le grand Cerf
Mégaloceros du Quaternaire d'Irlande développa ainsi des bois surdimensionnés atteignant 2,50 mètres
d'envergure, mais en fait conformes au développement de la taille de son corps.
 Au sujet des limites de la notion d'adaptation, Cuénot conclu ainsi :
« Dans une machine industrielle bien étudiée, il n'y a pas de rouage indifférent ; chaque écrou
a son rôle éventuel ; la courbure des pièces, leur poids, leur épaisseur, ont été l'objet de recherches
bannissant tout ce qui est inutile ; il n'y a pas d'organes rudimentaires, à moins qu'on ne se soit servi
de vieilles pièces provenant d'autres machines, et gardant la trace de leur fonctionnement primitif ; il
n'y a pas non plus de superflu, à moins que l'artisan, voulant rendre son œuvre plus agréable, n'y ait
ajouté des ornements, des sculptures, comme dans les outils d'autrefois. La machine vivante, au
contraire, a un passé où elle était autre qu'actuellement, et qui a laissé des traces ; la Nature ne lui
demande que de vivre et de durer, tant bien que mal, et il lui importe peu que son fonctionnement soit
économique.
La position des biologistes modernes vis-à-vis la question de l'adaptation est donc, je pense à
juste titre, tout autre que celle des naturalistes qui les ont précédés, de Bernardin de Saint-Pierre à
Weismann : ces derniers, pour des raisons sans doute différentes mais qui aboutissaient au même
résultat, étaient persuadés que tout était adapté, que chaque détail des organismes devait avoir une
signification utile, un rôle à jouer : sans doute cette conviction a priori du cause-finalier ou du
sélectionniste a souvent amené les physiologistes à des découvertes capitales, en les incitant à
rechercher avec persévérance la fonction de petits organes jugés d'abord insignifiants, tels que le
corps thyroïde, l'hypophyse, le thymus, les capsules surrénales, le corps jaune ovarien, les îlots de
Langerhans du pancréas, etc., qui en effet ont un rôle important dans la coordination de l'organisme.
[…]
Mais la médaille a un revers : cette conviction a amené bien souvent les naturalistes à
rechercher et à attribuer des significations utiles à des structures qui n'en ont probablement aucune, et
à errer grandement au sujet des adaptations. » pp. 52-53.
 En effet, la notion d'adaptation est devenue en quelque sorte la tarte à la crème de la biologie
évolutive, elle est systématiquement convoquée, conjointement à la sélection naturelle, pour expliquer les
particularités des êtres vivants, alors les études éthologiques qui pourraient en confirmer la pertinence sont
inexistantes ou impossibles à mener (cas des fossiles).
Étienne Rabaud est un des biologiste qui ont critiqué la notion d'adaptation (et à travers elle le
mécanisme de la sélection naturelle) de la manière la plus radicale:
"L'hypothèse [darwinienne] ne résiste pas à la critique la plus élémentaire. Ne suffit-il pas de
constater que l'appréciation d'un avantage tourne dans un cercle vicieux ? Quand un organisme
persiste, nous décidons qu'il possède une disposition avantageuse, et nous déclarons avantageuse
une disposition quelconque, précisément parce que l'organisme persiste." [1]
Rabaud remarque également que les explications concernent souvent des organes isolés, alors que
l'organisme forme un tout, et que plus rarement encore des comparaisons sont faites entre les êtres vivants
ayant des dispositions analogues, afin de déterminer la réalité de l'avantage ou du rôle que joue l'organe
pour les êtres vivants concernés. Il constate également que les interprétations mises en avant pour justifier
l’existence d’une particularité chez une espèce ne tiennent généralement pas compte du fait que d’autres
espèces vivant dans le même milieu n’ont pas cette disposition supposée avantageuse, voire ont la
disposition opposée et ne s’en portent pas plus mal.
Il en conclu que la notion d'adaptation est trompeuse et qu'elle est un obstacle à l'étude plus fine et
plus précise des rapports effectifs des êtres vivants entre eux et avec leur milieu. Pour lui, la notion
d'adaptation induit à prendre les conséquences pour les causes et inversement : ce n'est pas parce que l'être
vit dans un milieu qu'il y est adapté, mais c'est plutôt parce qu'il y trouve de quoi vivre, qu'il est en adéquation
avec les conditions, qu'il habite dans ce milieu.
 Pour Rabaud, l'environnement n'est pas uniquement une contrainte qui s'impose à l'organisme, c'est
aussi et avant tout l'espace où peut se déploier son activité autonome : l'être vivant n'est pas adapté au
milieu ; c'est le contraire, il trouve dans le milieu les éléments spécifiques qui lui permettent d'assurer sa
subsistance. L'analogie du vivant avec une machine induit à négliger et tend à faire oublier le caractère actif
des êtres vivants dans la quête de leurs subsistances (particulièrement évidente chez les animaux), c'est-à-
dire l'autonomie du vivant par rapport à son milieu.

Adaptation [modifier]
Selon les modèles théoriques, le rôle de l'adaptation dans l'évolution biologique est plus ou moins
important. Selon la perspective du paradigme adaptationniste, il s'agit du principal facteur de transformation
des espèces.
On parle d'adaptabilité pour désigner la plasticité de certaines espèces face aux forces de l'évolution.

Notes et références [modifier]

1. ↑ Étienne Rabaud, Introduction aux sciences biologiques, 1941, p. 181.

Bibliographie [modifier]
Lucien Cuénot,
• L'adaptation, éd. Doin, 1925.
• Invention et finalité en biologie, éd. Flammarion, 1941.
Stephen Jay Gould
• & Richard C. Lewontin (1979). "The Spandrels of San Marco and the Panglossian Paradigm:
A Critique of the Adaptationist Programme" Proc. Roy. Soc. London B 205 (1979) pp. 581-598.
• & Richard C. Lewontin (1982). "L’adaptation biologique: les trompes de l'églises San Marco
et le paradigme panglossien", La Recherche 13(139).1494-1502.
• (1997). "The exaptive excellence of spandrels as a term and prototype" Proceedings of the
National Academy of Sciences USA. 94: 10750-10755.
Étienne Rabaud,
• L'adaptation et l'évolution, éd. Chiron, 1922.
• Zoologie biologique, éd. Gauthier-Villars, 1934.
• Transformisme et adaptation, éd. Flammarion, 1942.

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique Dérive génétique · Mutation génétique · Recombinaison

évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens
 Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des
Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Adaptation_(biologie) ».

Catégories : Sélection naturelle | Processus évolutif | [+]

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Cladistics
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Cladistics[1] is a method of classifying species of organisms into groups called clades, which consist
of 1) all the descendants of an ancestral organism and 2) the ancestor itself. For example, birds, dinosaurs,
crocodiles, and all descendants (living or extinct) of their most recent common ancestor form a clade.[2] In
the terms of biological systematics, a clade is a single "branch" on the "tree of life", a monophyletic group.
 Cladistics can be distinguished from other taxonomic systems, such as phenetics, by its focus on
shared derived characters (synapomorphies). Previous systems usually employed overall morphological
similarity to group species into genera, families and other higher level classification; cladistic classifications
(usually trees called cladograms) are intended to reflect the relative recency of common ancestry or the
sharing of homologous features. Cladistics is also distinguished by its emphasis on parsimony and
hypothesis testing (particularly falsificationism), rather than subjective decisions that some other taxonomic
systems rely upon.[3]
Cladistics originated in the work of the German entomologist Willi Hennig, who referred to it as
"phylogenetic systematics" (also the name of his 1966 book); the use of the terms "cladistics" and "clade"
was popularized by other researchers. The technique and sometimes the name have been successfully
applied in other disciplines: for example, to determine the relationships between the surviving manuscripts of
the Canterbury Tales.[4]
Cladists use cladograms, diagrams which show ancestral relations between species, to represent the
monophyletic relationships of species, termed sister-group relationships. This is interpreted as representing
phylogeny, or evolutionary relationships. Although traditionally such cladograms were generated largely on
the basis of morphological characters, genetic sequencing data and computational phylogenetics are now
very commonly used in the generation of cladograms.
Contents
[hide]
• 1 History of cladistics
• 1.1 Cladistics as a successor to phenetics
• 2 Clades
• 2.1 Terminology
• 2.2 Three definitions of clade
• 3 Cladograms
• 4 Cladistics in taxonomy
• 4.1 Phylogenetic nomenclature contrasted with
traditional taxonomy
• 4.2 Paraphyletic groups discouraged
• 4.3 Complexity of the Tree of Life
• 4.4 PhyloCode approach to naming species
• 4.5 Example
• 4.6 Summary of advantages of phylogenetic
nomenclature
• 4.7 Summary of criticisms of phylogenetic
nomenclature
• 5 Application to other disciplines
• 6 See also
[edit] History of cladistics
The term clade was introduced in 1958 by Julian Huxley, cladistic by Cain and Harrison in 1960, and
cladist (for an adherent of Hennig's school) by Mayr in 1965.[5] Hennig referred to his own approach as
phylogenetic systematics. From the time of his original formulation until the end of the 1980s cladistics
remained a minority approach to classification. However in the 1990s it rapidly became the dominant method
of classification in evolutionary biology. Computers made it possible to process large quantities of data about
organisms and their characteristics. At about the same time the development of effective polymerase chain
reaction techniques made it possible to apply cladistic methods of analysis to biochemical and molecular
genetic features of organisms as well as to anatomical ones.[6]

[edit] Cladistics as a successor to phenetics

For some decades in the mid to late twentieth century, a commonly used methodology was phenetics
("numerical taxonomy"). This can be seen as a predecessor[7] to some methods of today's cladistics (namely
distance matrix methods such as neighbor-joining), but made no attempt to resolve phylogeny, only
similarities.
[edit] Clades

The yellow group (sauropsids) is monophyletic, the blue group (reptiles) is paraphyletic, and the red
group (warm-blooded animals) is polyphyletic.
Main article: Clade
 A clade is a group of taxa consisting of an ancestor taxon and all of its descendant taxa. In the
diagram provided, it is hypothesized that the first vertebrate species is the common ancestor of all vertebrate
species, including fishes (Pisces). The first tetrapod is the ancestor of all tetrapods, including amphibians,
reptiles, mammals and birds. The tetrapod ancestor was a descendant of the original vertebrate ancestor, but
is not an ancestor of any fish living today. The tetrapod clade is thus nested within the vertebrate clade.
An important caution is that any cladogram is a provisional hypothesis. In a hypothetical example,
further genetic or morphological evidence might suggest that fish and amphibians share a common ancestor
that was not an ancestor of the other tetrapods. The new information would cause us to define a fish-and-
amphibian clade, altering the cladogram.

[edit] Terminology
The following terms are used to identify shared or distinct characters among groups:[8][9][10]
• Plesiomorphy ("close form") or ancestral state, also symplesiomorphy ("shared
plesiomorphy"), is a characteristic that is present at the base of the tree. For example, in the tree
shown, the presence of a backbone (shared by all vertebrates) can be hypothesized to have existed
in the common vertebrate ancestor.
• Apomorphy ("separate form") or derived state is a characteristic believed to have evolved
within the tree. For example, all tetrapods have four limbs; thus, having four limbs is an apomorphy
for vertebrates but a plesiomorphy for tetrapods.
• Synapomorphy ("shared apomorphy") is an apomorphy which is shared between taxa.
The application of the above terms to a group depends on one's perspective in the tree. They are
thus relative terms. For example, an apomorphy of one clade is the plesiomorphy of another contained within
it. These terms are equivalent to but more precise than the homology, allowing one to express the
hierarchical relationships among different homologies.
Three main types of groups can be identified by plotting their relationships in cladograms:[8][9][10]
• Monophyletic groups are groups containing only taxa descended from a given ancestor taxon
and all those descendants. In the diagram, all vertebrates are monophyletic, since all the taxa are
descended from a single ancestor (the common vertebrate species) and there are no others.
Monophyly is diagnosed by the synapomorphy relation.
• Paraphyletic groups are groups excluding one or more descendant taxa of the common
ancestor, and thus contain some but not all of its descendants. For instance, excluding birds from the
sample cladogram would create a paraphyletic group, reptiles. Paraphyletic groups are typically
diagnosed on the basis of shared plesiomorphy and the exclusion of groups that are diagnosable by
a synapomorphy. For example, birds are warm-blooded, excluding them from being reptiles, which
because of that exclusion is a paraphyletic group, as birds are also descended from amniotes. The
plesiomorphy is the amniotic fluid and the synapomorphy is warm-bloodedness.
• Polyphyletic groups are groups containing taxa from two or more different monophyletic
groups. For instance, a grouping of birds and mammals based on their warm-bloodedness is not
monophyletic. In the diagram, the Aves ancestor came from the Archosaurs, but the Mammalian
ancestor came from the Amniotes. Although there is an ultimate common ancestor among the
Tetrapods, the latest common ancestors are not the same and therefore the group is polyphyletic.
The warm-bloodedness is not a plesiomorphy but is a homoplasy, or convergence. Polyphyletic
groups are recognized by the homoplasy relation (that is, a group is polyphyletic because it is
diagnosed by a character that actually forms a homoplasy, see wastebasket taxon).
Clades relate to each other in these ways:
• A clade is basal to another clade if it contains that other clade as a subset within it. In this
case, the vertebrate clade is basal to the tetrapod and fish clades. The use of "basal" to mean a
clade that is less species-rich than a sister clade, with such a deficit being taken as an indication of
primitiveness, is incorrectly applied.[11]
• A clade located within a clade is said to be nested within that clade. The bird clade is nested
within the reptilian clade.
[edit] Three definitions of clade

The three ways to define a clade

There are three major ways to define a clade for use in a cladistic taxonomy.[12]
• Node-based: the last common ancestor of A and B, and all descendants of that ancestor.
Crown groups are a type of node-based clade.
 • Branch-based: the first ancestor of A which is not also an ancestor of C, and all descendants
of that ancestor. (This type of definition was originally called "stem-based", but this was changed to
avoid confusion with the term "stem group".) Total groups are a type of branch-based clade.
• Apomorphy-based: the first ancestor of A to possess derived trait M homologously (that is,
synapomorphically) with that trait in A, and all descendants of that ancestor. The process of
identifying and naming groups based on apomorphies is the method that most resembles classical
systematics, with the proviso that cladistic taxa always denote a clade.
See also: Phylogenetic definitions

[edit] Cladograms
Main articles: Cladogram and Polytomy
Cladists use cladograms, diagrams which show ancestral relations between taxa, to represent the
evolutionary tree of life. Although traditionally such cladograms were generated largely on the basis of
morphological characters, molecular sequencing data and computational phylogenetics are now very
commonly used in the generation of cladograms.
The starting point of cladistic analysis is a group of species and molecular, morphological, or other
data characterizing those species. The end result is a tree-like relationship diagram called a cladogram,[13]
or sometimes a dendrogram (Greek for "tree drawing").[14] The cladogram graphically represents a
hypothetical evolutionary process. Cladograms are subject to revision as additional data become available.
The terms "evolutionary tree", and sometimes "phylogenetic tree" are often used synonymously with
cladogram,[15] but others treat phylogenetic tree as a broader term that includes trees generated with a
nonevolutionary emphasis. In cladograms, all species lie at the leaves. The two taxa on either side of a split,
with a common ancestor and no additional descendents, are called "sister taxa" or "sister groups".[16] Each
subtree, whether it contains only two or a hundred thousand items, is called a "clade". Many cladists require
that all forks in a cladogram be 2-way forks. Some cladograms include 3-way or 4-way forks when there are
insufficient data to resolve the forking to a higher level of detail (see under phylogenetic tree).
For a given set of species, the number of distinct cladograms that can be drawn (ignoring which
cladogram best matches the species characteristics) is:[17]

Number of 2 3 4 5 6 7 8
species

Number of 1 3 15 105 945 10,395 135,135

cladograms

This superexponential growth of the number of possible cladograms explains why manual creation of
cladograms becomes very difficult when the number of species is large. If a cladogram represents N species,
the number of levels (the "depth") in the cladogram is on the order of log 2(N).[18] For example, if there are 32
species of deer, a cladogram representing deer could be around 5 levels deep (because 2 5 = 32), although
this is really just the lower limit. A cladogram representing the complete tree of life, with about 10 million
species, could be about 23 levels deep. This formula gives a lower limit, with the actual depth generally a
larger value, because the various branches of the cladogram will not be uniformly deep. Conversely, the
depth may be shallower if forks larger than 2-way forks are permitted.
 A cladogram tree has an implicit time axis,[19] with time running forward from the base of the tree to
the leaves of the tree. If the approximate date (for example, expressed as millions of years ago) of all the
evolutionary forks were known, those dates could be captured in the cladogram. Thus, the time axis of the
cladogram could be assigned a time scale (e.g. 1 cm = 1 million years), and the forks of the tree could be
graphically located along the time axis. Such cladograms are called scaled cladograms. Many cladograms
are not of this type, for a variety of reasons:
• They are built from species characteristics that cannot be readily dated (e.g. morphological
data in the absence of fossils or other dating information)
• When the characteristic data are DNA/RNA sequences, it is feasible to use sequence
differences to establish the relative ages of the forks, but converting those ages into actual years
requires a significant approximation of the rate of change[20]
• Even when the dating information is available, positioning the cladogram's forks along the
time axis in proportion to their dates may cause the cladogram to become difficult to understand or
hard to fit within a human-readable format
Cladistics makes no distinction between extinct and extant species,[21] and it is appropriate to
include extinct species in the group of organisms being analyzed. Cladograms that are based on DNA/RNA
generally do not include extinct species because DNA/RNA samples from extinct species are rare.
Cladograms based on morphology, especially morphological characteristics that are preserved in fossils, are
more likely to include extinct species.

[edit] Cladistics in taxonomy

Main article: Phylogenetic nomenclature
[edit] Phylogenetic nomenclature contrasted with traditional taxonomy

A highly resolved, automatically generated tree of life based on completely sequenced genomes[22]
Most taxonomists have used the traditional approaches of Linnaean taxonomy and later Evolutionary
taxonomy to organize life forms. These approaches use several fixed levels of a hierarchy, such as kingdom,
phylum, class, order, and family. Phylogenetic nomenclature does not feature those terms, because the
evolutionary tree is so deep and so complex that it is inadvisable to set a fixed number of levels.
Evolutionary taxonomy insists that groups reflect phylogenies. In contrast, Linnaean taxonomy allows
both monophyletic and paraphyletic groups as taxa. Since the early 20th century, Linnaean taxonomists have
generally attempted to make at least family- and lower-level taxa (i.e. those regulated by the codes of
nomenclature) monophyletic. Ernst Mayr in 1985 drew a distinction between the terms cladistics and
phylogeny:[23]
"It would seem to me to be quite evident that the two concepts of phylogeny (and their role in the
construction of classifications) are sufficiently different to require terminological distinction. The
term phylogeny should be retained for the broad concept of phylogeny, promoted by Darwin and
adopted by most students of phylogeny in the ensuing 90 years. The concept of phylogeny as
mere genealogy should be terminologically distinguished as cladistics. To lump the two
concepts together terminologically could not help but produce harmful equivocation."

Willi Hennig's pioneering work provoked a spirited debate[24] about the relative merits of
phylogenetic nomenclature versus Linnaean or evolutionary taxonomy, which has continued down to the
present,[25] however Hennig did not advocate abandoning the Linnaean nomenclatural system. Some of the
debates in which the cladists were engaged had been running since the 19th century, but they were renewed
fervor,[26] as can be seen from the Foreword to Hennig (1979) by Rosen, Nelson, and Patterson:[27]
"Encumbered with vague and slippery ideas about adaptation, fitness, biological species and
natural selection, neo-Darwinism (summed up in the "evolutionary" systematics of Mayr and
 Simpson) not only lacked a definable investigatory method, but came to depend, both for
evolutionary interpretation and classification, on consensus or authority."

Phylogenetic nomenclature strictly and exclusively follows phylogeny and has arbitrarily deep trees
with binary branching: each taxon corresponds to a clade. Linnaean taxonomy, while since the advent of
evolutionary theory following phylogeny, also may subjectively consider similarity and has a fixed hierarchy of
taxonomic ranks, and its taxa are not required to correspond to clades.

[edit] Paraphyletic groups discouraged

Many cladists discourage the use of paraphyletic groups in classification of organisms, because they
detract from cladistics' emphasis on clades (monophyletic groups). In contrast, proponents of the use of
paraphyletic groups argue that any dividing line in a cladogram creates both a monophyletic section above
and a paraphyletic section below. They also contend that paraphyletic taxa are necessary for classifying
earlier sections of the tree – for instance, the early vertebrates that would someday evolve into the family
Hominidae cannot be placed in any other monophyletic family. They also argue that paraphyletic taxa provide
information about significant changes in organisms' morphology, ecology, or life history – in short, that both
paraphyletic groups and clades are valuable notions with separate purposes.[ citation needed]

[edit] Complexity of the Tree of Life

The cladistic tree of life is a fractal:[28]
"The tree of life is inherently fractal-like in its complexity, .... Look closely at the 'lineage' of a
phylogeny ... and it dissolves into many smaller lineages, and so on, down to a very fine scale."
 The overall shape of a dichotomous (bifurcating) tree is recursive; as a viewpoint zooms into the tree
of life, the same type of tree appears no matter what the scale. When extinct species are considered (both
known and unknown), the complexity and depth of the tree can be very large. Moreover the tree continues to
recreate itself by bifurcation, a series of events called fractal evolution.[29]. Every single speciation event,
including all the species that are now extinct, represents an additional fork on the hypothetical, complete
cladogram of the tree of life.
The tree of life is a quasi-self-similar fractal; that is, the deep reconstruction is not as regular as the
shallow reconstruction.[30] By shallow Mishler means the most recent branching toward and at the tips, and
by deep the more ancient branches further back, which are harder to reconstruct and are missing unknown
extinct lines. In the shallow part of the tree, branching events are relatively regular; it is often possible to
estimate the times between them. In the deep part of the tree, "homology assessments" are "difficult" and the
times vary widely.[31] At this level Eldredge's and Gould's punctuated equilibrium applies, which
hypothesizes long periods of stability followed by punctuations of rapid speciation, based on the fossil record.

[edit] PhyloCode approach to naming species

A formal code of phylogenetic nomenclature, the PhyloCode,[32] is currently under development. It is
intended for use by both those who would like to abandon Linnaean taxonomy and those who would like to
use taxa and clades side by side. In several instances (see for example Hesperornithes) it has been
employed to clarify uncertainties in Linnaean systematics so that in combination they yield a taxonomy that
unambiguously places problematic groups in the evolutionary tree in a way that is consistent with current
knowledge.
[edit] Example
For example, Linnaean taxonomy contains the taxon Tetrapoda, defined morphologically as
vertebrates with four limbs (as well as animals with four-limbed ancestors, such as snakes), which is often
given the rank of superclass, and divides into the classes Amphibia, Reptilia, Aves, Mammalia.
Phylogenetic nomenclature also contains the taxon Tetrapoda (see the diagram under Clades
above), whose living members can be classified phylogenically as "the clade defined by the common
ancestor of amphibians and mammals", or more precisely the clade defined by the common ancestor of a
specific amphibian and mammal (or bird or snake). This definition gives us the Crown group tetrapods (or
Crown-Tetrapoda). A few primitive four legged ancestors (the Ichthyostegalia) fall outside Crown-Tetrapoda.
[33] An alternative is to define tetrapoda as all animals more closely related to mammals than to lungfish (our
nearest living non-tetrapod relatives). In this definition, the ichthyostegalians are included, together with a
host of fossil animals usually classed as crossopterygian fish. This wider definition is termed Pan-Tetrapoda.
A third option is to define Tetrapoda according to their apomorphy (their unique trait, i.e. having legs rather
than fins), a definition that yield the same group as the Linnaean taxon.
Non of the phylogenetic taxa as described above have a rank, and neither do its subtaxa. All the
subclades are contained within one another. The clades are not divided into several non-overlapping taxa (as
in traditional taxonomy), rather the clade is split into two clades at the first branching, a process repeated
througout. With regards to the traditional classes, Aves and Mammalia are subclades, contained in the
subclade Amniota, while Reptilia and Amphibia are paraphyletic taxa, not clades.[34]. Instead of classifying
non-mammalian, non-avian amniotes as reptiles, Amniota is divided into the two clades Sauropsida (which
contains birds and all living amniotes other than mammals, including all living traditional reptiles) and
Theropsida (mammals and the extinct mammal-like reptiles). Similarly, Amphibia can be split into the
Batrachomorpha (fossil amphibians more closely related to modern amphibians) and Reptiliomorpha, the
latter of which the amiotes is a sub-clade.[35]. Ichthyostegalians and other Stem-tetrapods represent sister
groups from splits predating the Batrachomorpha/Reptilopmorpha split.[33]
Further information: Reptiles, History of Classification

[edit] Summary of advantages of phylogenetic nomenclature

Proponents of phylogenetic nomenclature enumerate key distinctions between phylogenetic
nomenclature and Linnaean taxonomy as follows:[36]

Phylogenetic Nomenclature Linnaean Taxonomy

Handles arbitrarily deep trees. Often must invent new level names (such as
superorder, suborder, infraorder, parvorder,
magnorder) to accommodate new discoveries. Biased
towards trees about 4 to 12 levels deep.

Discourages naming or use of groups that Acceptable to name and use paraphyletic
are not monophyletic groups

Primary goal is to reflect actual process of Primary goal is to group species based on
evolution morphological similarities
 Assumes that the shape of the tree will New discoveries often require renaming or
change frequently, with new discoveries releveling of Classes, Orders, and Kingdoms

[edit] Summary of criticisms of phylogenetic nomenclature

Critics of phylogenetic nomenclature include Ashlock,[37] Mayr,[38] Williams.[39] Some of their
criticisms include:

Phylogenetic Nomenclature Linnaean Taxonomy

Limited to entities related by evolution or Supports groupings without reference to

ancestry evolution or ancestry

Does not include a process for naming Includes a process for giving unique names to
species species

Clade definitions emphasize ancestry at the Taxa definitions based on tangible

expense of descriptive characteristics characteristics

Ignores sensible, clearly defined paraphyletic Permits clearly defined groups such as
groups such as reptiles reptiles

Difficult to determine if a given species is in a Straightforward process to determine if a

clade or not (e.g. if clade X is defined as "most given species is in a taxon or not
recent common ancestor of A and B along with its
descendants", then the only way to determine if
species Y is in the clade is to perform a complex
evolutionary analysis)

Limited to organisms that evolved by Applicable to all organisms, regardless of

inherited traits; not applicable to organisms that evolutionary mechanism
evolved via complex gene sharing or lateral transfer

[edit] Application to other disciplines

The comparisons used to acquire data on which cladograms can be based are not limited to the field
of biology.[40] Any group of individuals or classes, hypothesized to have a common ancestor, and to which a
set of common characteristics may or may not apply, can be compared pairwise. Cladograms can be used to
depict the hypothetical descent relationships within groups of items in many different academic realms. The
only requirement is that the items have characteristics that can be identified and measured.
Recent attempts to use cladistic methods outside of biology address the reconstruction of lineages
in:
• Anthropology and archeology.[41] Compares cultures or artifacts using groups of cultural
traits or artifact features.
• Linguistics.[42] Compares languages using groups of linguistic features.
 • Textual criticism or Stemmatics.[43][44] Compares manuscripts of the same work (original
lost) using groups of distinctive copying errors.
• Ethology.[45] Compares animal species using behavioral traits presumed hereditary.

[edit] See also

• Bauplan • Important • Phylogenomics
• Bioinformatics publications in • Phylogeography
• Biomathematics phylogenetics • Phylogenetic
• Coalescent theory • Language family comparative methods
• Computational • Maximum • Scientific
phylogenetics parsimony Classification
• Dendrogram • Molecular • Stratocladistics
• Evolution of phylogeny • Systematics
Mollusca • PhyloCode • Tree structure
• Last common • Phylogenetics
ancestor • Phylogenetic
network
• Phylogenetics
software
[edit] References
1. ^ (Ancient Greek: κλάδος, klados, "branch")
2. ^ Glossary entry "clade" Understanding Evolution. 2010. University of California Museum of
Paleontology.
3. ^ Phillips, Ben (2000). "Natural History Collections: Cladistics". University of Edinburgh.
http://www.nhc.ed.ac.uk/index.php?page=236.273.444. Retrieved 4 July 2009.
4. ^ "Canterbury Tales Project". http://www.canterburytalesproject.org. Retrieved 4 July 2009.
5. ^ Dupuis 1984
6. ^ Baron, C.; Høeg, J.T. (2005). "Gould, Scharm and the paleontologocal perspective in
evolutionary biology". in Koenemann, S.; Jenner, R.A.. Crustacea and Arthropod Relationships. CRC
Press. pp. 3–14. ISBN 0849334985. http://books.google.com/?id=LalmQ4346O0C&dq=Nielsen,+C.
+2001+%22Animal+evolution%22+Chelicerata. Retrieved 2008-10-15.
7. ^ Mayr 1982, p. 221
8. ^ a b Patterson 1982, pp. 21–74
9. ^ a b Patterson, Colin (1988). "Homology in classical and molecular biology". Molecular
Biology and Evolution 5 (6): 603–625. PMID 3065587.
10.^ a b de Pinna, M.G.G (1991). "Concepts and tests of homology in the cladistic paradigm".
Cladistics 7: 367–394. doi:10.1111/j.1096-0031.1991.tb00045.x.
11.^ Krell, F.-T.; Cranston, P. (2004). "Which side of the tree is more basal?". Systematic
Entomology 29 (3): 279–281. doi:10.1111/j.0307-6970.2004.00262.x.
12.^ de Queiroz, K.; Gauthier, J. (1994). "Toward a phylogenetic system of biological
nomenclature". Trends in Research in Ecology and Evolution 9 (1): 27–31. doi:10.1016/0169-
5347(94)90231-3.
 13.^ Cracraft, Joel; Donaghue, Michael J., eds (2004). Assembling the Tree of Life. Oxford,
England: Oxford University Press. pp. 45, 78, 555.
14.^ Weygoldt, P. (February 1998). "Evolution and systematics of the Chelicerata".
Experimental and Applied Acarology 22 (2): 63–79. doi:10.1023/A:1006037525704.
15.^ Singh, Gurcharan (2004). Plant Systematics: An Integrated Approach. Science. pp. 203–4.
ISBN 1578083516.
16.^ Gould, Steven Jay (1983). Hen's Teeth and Horse's Toes. New York: Norton. p. 357.
ISBN 0393017168.
17.^ Lowe, Andrew (2004). Ecological Genetics: Design, Analysis, and Application . Blackwell
Publishing. p. 164. ISBN 1405100338.
18.^ Aldous, David (1996). "Probability Distributions on Cladograms". Random Discrete
Structures. Springer. pp. 13. ISBN 0387946233.
19.^ Freeman, Scott (1998). Evolutionary Analysis. Prentice Hall. p. 380. ISBN 0135680239.
20.^ Carroll, Robert Lynn (1997). Patterns and Processes of Vertebrate Evolution. Cambridge
University Press. p. 80. ISBN 052147809X.
21.^ Scott-Ram, N. R. (1990). Transformed Cladistics, Taxonomy and Evolution. Cambridge
University Press. p. 83. ISBN 0521340861.
22.^ Letunic, I.; Bork, P. (2007). "Interactive Tree Of Life (iTOL): an online tool for phylogenetic
tree display and annotation" (Pubmed). Bioinformatics 23 (1): 127–8.
doi:10.1093/bioinformatics/btl529. PMID 17050570.
23.^ Mayr, E (1985). "Darwin and the Definition of Phylogeny". Systematic Zoology 34 (1): 97–
98. doi:10.2307/2413350. http://jstor.org/stable/2413350. .
24.^ Wheeler, Quentin (2000). Species Concepts and Phylogenetic Theory: A Debate . Columbia
University Press. ISBN 0231101430.
 25.^ Benton, M. (2000). "Stems, nodes, crown clades, and rank-free lists: is Linnaeus dead?".
Biological Reviews 75 (4): 633–648. PMID 11117201.
26.^ Hull, David (1988). Science as a Process. University of Chicago Press. pp. 232–276.
ISBN 0226360512.
27.^ Hennig 1979, p. ix
28.^ Mishler 2005, p. 57
29.^ Gordon, Richard (1999). The Hierarchical Genome and Differentiation Waves. World
Scientific. p. 664. ISBN 9810222688.
30.^ Mishler 2005, p. 62
31.^ Mishler 2005, p. 62: "In deep reconstruction problems, the branch events at issue
happened a relatively long time ago and the ... relative lengths of ... branches are often quite
different...."
32.^ Pennisi, E. (2001). "Evolutionary Biology: Preparing the Ground for a Modern 'Tree of Life'".
Science 293 (5537): 1979–1980. doi:10.1126/science.293.5537.1979. PMID 11557858.
33.^ a b Clarck, J.A. (1997): Ichthyostega, Tree of Life website
34.^ Tudge, Colin (2002). The Variety of Life: A survey and a Celebration of all the Creatures
that Have Ever Lived. Oxford: Oxford University Press. p. 83. ISBN 0-19-860426-2.
35.^ Benton, M. J. (2000), Vertebrate Paleontology, 2nd Ed. Blackwell Science Ltd 3rd ed. 2004
– see also taxonomic hierarchy of the vertebrates, according to Benton 2004
36.^ Hennig 1975.
37.^ Ashlock 1971, Ashlock 1972, Ashlock 1974, Ashlock 1979.
38.^ Mayr 1974, Mayr 1978, Mayr, E; Bock, WJ (2002). "Classifications and other ordering
systems". Journal of Zoological Systematics and Evolutionary Research 40: 169–194.
doi:10.1046/j.1439-0469.2002.00211.x.
 39.^ Williams, P.A. (1992). "Confusion in cladism". Synthese 01: 135–132.
40.^ Mace, Ruth; Clare, Clare J.; Shennan, Stephen, eds (2005). The Evolution of Cultural
Diversity: A Phylogenetic Approach. Cavendish Publishing. p. 1.
41.^ Lipo, Carl; O'Brien, Michael J.; Collard, Mark et al., eds (2006). Mapping Our Ancestors:
Phylogenetic Approaches in Anthropology and Prehistory. Piscataway: Transaction Publishers.
42.^ Oppenheimer, Stephen (2006). The Origins of the British. London: Robinson. pp. 290–300,
340–56. ISBN 0786718900.
43.^ Ehrman, Bart D. (2005). The text of the New Testament: its tranmission, corruption and
restoration (4th ed.). New York: Oxford University Press. p. 207 ff.
44.^ Robinson, Peter M.W.; O’Hara, Robert J. (1996). "Cladistic analysis of an Old Norse
manuscript tradition". Research in Humanities Computing 4: 115–137. http://rjohara.net/cv/1996-rhc.
45.^ Jerison, Harry J. (2003). "On Theory in Comparative Psychology". in Sternberg, Robert J.;
Kaufman, James C.. The evolution of intelligence. Mahwah, NJ: Lawrence Erlbaum Associates, Inc..
p. 254. ISBN 0123852501 .

[edit] Bibliography
• Ashlock, Peter D. (1971). "Monophyly and associated terms". Systematic Zoology 20 (1): 63–
69. doi:10.2307/2412223. http://jstor.org/stable/2412223.
• Ashlock, Peter D. (1972). "Monophyly again". Systematic Zoology 21 (4): 430–438.
doi:10.2307/2412435. http://jstor.org/stable/2412435.
• Ashlock, Peter D. (1974). "The uses of cladistics". Annual Review of Ecology and
Systematics 5: 81–99. doi:10.1146/annurev.es.05.110174.000501. ISSN 0066-4162.
 • Ashlock, Peter D. (1979). "An evolutionary systematist’s view of classification". Systematic
Zoology 28 (4): 441–450. doi:10.2307/2412559. http://jstor.org/stable/2412559.
• Cuénot, Lucien (1940). "Remarques sur un essai d'arbre généalogique du règne animal".
Comptes Rendus de l'Académie des Sciences de Paris 210: 23–27. Available free online at
http://gallica.bnf.fr (No direct URL). This is the paper credited by Hennig (1979) for the first use of the
term 'clade'.
• Dupuis, Claude (1984). "Willi Hennig's impact on taxonomic thought". Annual Review of
Ecology and Systematics 15: 1–24. ISSN 0066-4162.
• Hennig, Willi (1975). "'Cladistic analysis or cladistic classification': a reply to Ernst Mayr".
Systematic Zoology 24 (2): 244–256. doi:10.2307/2412765. http://jstor.org/stable/2412765. The
paper to which he was responding is reprinted in Mayr 1976.
• Hennig, Willi (1966). Phylogenetic systematics (tr. D. Dwight Davis and Rainer Zangerl) .
Urbana, IL: Univ. of Illinois Press (reprinted 1979 and 1999). ISBN 0-252-06814-9.
• Hennig, Willi (1979). Phylogenetic systematics (3rd edition of 1966 book). Urbana: University
of Illinois Press. ISBN 0-252-06814-9. Translated from manuscript and so never published in
German.
• Hull, David L. (1979). "The limits of cladism". Systematic Zoology 28 (4): 416–440.
doi:10.2307/2412558. http://jstor.org/stable/2412558.
• Mayr, Ernst (1974). "Cladistic analysis or cladistic classification?". Zeitschrift fűr Zoologische
Systematik und Evolutionforschung 12: 94–128.
• Mayr, Ernst (1976). Evolution and the diversity of life (Selected essays). Cambridge, MA:
Harvard University Press. ISBN 0-674-27105-X. Reissued 1997 in paperback. Includes a reprint of
 Mayr's 1974 anti-cladistics paper at pp. 433–476, "Cladistic analysis or cladistic classification." This is
the paper to which Hennig 1975 is a response.
• Mayr, Ernst (1978). "Origin and history of some terms in systematic and evolutionary
biology". Systematic Zoology 27 (1): 83–88. doi:10.2307/2412818. http://jstor.org/stable/2412818.
• Mayr, Ernst (1982). The growth of biological thought: diversity, evolution and inheritance .
Cambridge, MA: Harvard University Press. ISBN 0-674-36446-5.
• Mishler, Brent D. (2005). "The logic of the data matrix in phylogenetic analysis". in Albert,
Victor. Parsimony, Phylogeny, and Genomics. Oxford University Press. pp. 57–70.
ISBN 0199297304 .
• Patterson, Colin (1982). "Morphological characters and homology". in Joysey, Kenneth A;
Friday, A. E.. Problems in Phylogenetic Reconstruction. Systematics Association Special Volume 21.
London: Academic Press. ISBN 0-12-391250-4 .
• Schuh, Randall T. and Andrew V. Z. Brower. 2009. Biological Systematics: principles and
applications (2nd edn.) Cornell University Press, ISBN 978-0-8014-4799-0

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 • Collins, Allen G.; Guralnick, Rob; Smith, Dave (1994-2005). "Journey into Phylogenetic
Systematics". University of California Museum of Paleontology.
http://www.ucmp.berkeley.edu/clad/clad4.html. Retrieved 21 January 2010.
• Felsenstein, Joe. "Phylogeny Programs". Seattle: University of Washington.
http://evolution.gs.washington.edu/phylip/software.html. Retrieved 21 January 2010.
• O'Neil, Dennis (1998-2008). "Classification of Living Things". San Marcos CA: Palomar
College. http://anthro.palomar.edu/animal/default.htm. Retrieved 21 January 2010.
• Robinson, Peter; O'Hara, Robert J. (1992). "Report on the Textual Criticism Challenge 1991".
rjohara.net. http://rjohara.net/darwin/files/bmcr. Retrieved 21 January 2010.
• Theobald, Douglas (1999-2004). "Phylogenetics Primer". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/comdesc/phylo.html. Retrieved 21 January 2010.
• Wiley, E.O.; Siegel-Causey, D.; Brooks, D.R.; Funk, V.A. (1991). "Chapter 1: Introduction,
Terms and Concepts". The Compleat Cladist: A Primer of Phylogenetic Procedures . The University
of Kansas Museum of Natural History Special Publication No. 19. American Museum of Natural
History. ISBN 0893380350. http://www.amnh.org/learn/pd/fish_2/pdf/compleat_cladist.pdf.

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Topics in phylogenetics

Relevant fields Computational phylogenetics · Molecular phylogeny · Cladistics

 Basic concepts Phylogenetic tree · Phylogenetic network · Long branch attraction · Clade ·
Ghost lineage

Inference Maximum parsimony · Maximum likelihood · Neighbor-joining · UPGMA ·

methods Bayesian inference · Least squares

Current topics PhyloCode · DNA barcoding

-morphy Symplesiomorphy · Apomorphy · Plesiomorphy · Synapomorphy ·

Autapomorphy

-phyly Monophyly/Holophyly · Paraphyly · Polyphyly

List of evolutionary biology topics

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 W000

Cladistique
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Ne doit pas être confondu avec Cladistia.

La cladistique, du grec ancien κλάδος (klados) signifiant « branche », aussi appelée systématique
phylogénétique, est une théorie concernant la classification phylogénétique, qui est l'étude de la
classification des êtres vivants selon leurs relations de parenté, dans un cadre évolutionniste. Elle repose sur
la construction de groupes monophylétiques (ou clades), c'est-à-dire des groupes incluant un ancêtre et
l'ensemble de sa descendance.
Exemples de cladogrammes
Sommaire
[masquer]
• 1 Description générale
• 2 Concepts importants de la cladistique
• 2.1 Le clade comme unité de classification
• 2.2 Paraphylie
• 2.3 Homologie vs. homoplasie
• 3 Lecture d'un cladogramme
• 4 Enracinement et groupe externe
• 5 Cladistique et évolution
• 6 Parcimonie, compatibilité et 3ia
• 6.1 La parcimonie
• 6.2 L'analyse à trois éléments
• 6.3 La compatibilité
• 7 Voir aussi
• 7.1 Bibliographie
• 7.2 Autres références
• 7.3 Liens externes
Description générale [modifier]

Willi Hennig (1972)

La cladistique se distingue des autres domaines de la systématique car elle cherche à établir les
relations de parenté strictes entre les taxons en distinguant caractères dérivés (apomorphies) et primitifs
(plésiomorphies). Un groupe monophylétique ne se fondera que sur le partage de caractères dérivés
propres, également appelés synapomorphies.
Cette méthode de reconstruction phylogénétique a été introduite dans les années 1950 par
l'entomologiste allemand Willi Hennig.
Les résultats de ce type de reconstruction peuvent être représentés sous forme de graphes
connexes non cycliques, ou arbres, appelés ici cladogrammes. Ces arbres récapitulent ainsi les relations de
parenté entre les taxons considérés. Les données morphologiques, moléculaires et comportementales sont
utilisées dans les travaux cladistiques. La reconstruction d'arbres est largement assistée par des logiciels
informatiques.
Concepts importants de la cladistique [modifier]

Le clade comme unité de classification [modifier]

Dans cet arbre phylogénétique, le groupe des sauropsides, constitué des reptiles, des oiseaux et de
leur ancêtre hypothétique commun, est considéré comme monophylétique.
La cladistique reconnait une unité de classement bien particulière, le clade. Un clade est un groupe
de taxons formant un groupe monophylétique, une totalité de descendance, un ancêtre commun et tous ses
descendants. Ce concept s'oppose à celui du grade, rapprochement de taxons reposant sur d'autres critères
(e.g. ressemblance générale, somme de modifications adaptatives). La cladistique qualifiera souvent ces
grades de paraphylétiques ou polyphylétiques selon que le rapprochement soit effectué sur la base de
plésiomorphies ou d'homoplasies. Certains grades sont cependant monophylétiques. Par exemple, les
algues forment un grade polyphylétique, les reptiles forment un grade paraphylétique et les mammifères
forment un grade monophylétique.
La notion de clade peut aussi être définie par un ensemble de taxons plus proches qu'ils ne le sont
entre eux que de tout autre taxon. Par exemple, les reptiles ne forment pas un clade car certains sont plus
apparentés aux oiseaux qu'aux autres reptiles (les Crocodiliens sont inclus dans les Archosauriens comme
les oiseaux). Il est à noter qu'un clade représente uniquement une unité évolutive. On parle aussi parfois de
groupe holophylétique, ce qui est synonyme de groupe monophylétique.
Paraphylie [modifier]

Le groupe paraphylétique des reptiles apparaît en bleu.

Si on parle de groupe monophylétique pour une totalité de descendance, la notion de paraphylie
s'applique à un groupe défini sur un ou des caractères plésiomorphes. Ce n'est donc pas une totalité de
descendance puisque les organismes portant les états dérivés en sont exclus. Un groupe paraphylétique
n'est donc pas monophylétique. Il est donc important en cladistique de bien séparer caractère plésiomorphe
et apomorphe pour déterminer si un groupe est monophylétique ou non. La méthode cladistique ne permet
donc pas l'existence de groupes tel que les reptiles, les poissons, les invertébrés, les procaryotes, etc. Dans
un groupe paraphylétique, certains taxons sont plus proches d'autres taxons hors du groupe que de taxons
au sein du groupe paraphylétique. Dans l'exemple des poissons, si on considère les Tétrapodes comme
groupe dérivé et les poissons comme groupe primitif par rapport aux Tétrapodes, on constate que certains
poissons sont plus proches des Tétrapodes qu'ils ne le sont d'autres poissons : par exemple le Cœlacanthe
est plus proche des Tétrapodes que du requin.
La notion de polarisation et de sens de transformation est ici importante. Un caractère pouvant
exister sous deux états: primitif (plésiomorphe) ou dérivé (apomorphe), il faut pouvoir déterminer quel état
est apomorphe et lequel est plésiomorphe. Pour cela on peut utiliser plusieurs critères dont le plus utilisé est
celui du groupe externe. D'autres critères sont utilisés comme le critère ontogénétique et à moindre mesure
les critères paléontologique et chorologique. Soient deux états a et b d'un même caractère ; on ne connait
pas le sens de transformation entre a et b on a donc a↔b. On peut donc admettre deux sens de
transformation : a→b ou b→a. Dans le cas a→b, a se transforme en b, donc a est plésiomorphe par rapport à
b et seul b nous renseigne sur les relations de parenté. Dans le cas b→a, b se transforme en a, b est donc
plésiomorphe par rapport à a et seul a nous renseigne sur les relations de parenté.
En cladistique un ensemble d'espèces ne peut pas être primitif : il n'y a pas de groupe ancestral. Si
un clade contient plusieurs groupes, c'est que chacun de ces groupes a des caractères dérivés qui les
distinguent des autres (autapomorphies), ce qui implique qu'un groupe ancestral ne peut pas exister en
cladistique. La notion d'ancêtre pourrait s'appliquer à l'espèce, cependant une espèce ancestrale est
nécessairement définie par des caractères primitifs donc plésiomorphes par rapport à ses descendants, elle
est donc forcément paraphylétique. En cladistique l'ancêtre est donc une hypothèse et n'est pas identifié, ce
qui, évidemment, n'implique pas qu'il n'existe pas mais simplement qu'il ne peut pas être défini en tant que
taxon ou espèce mais simplement en tant qu'hypothèse d'un ensemble de caractères primitifs (on parle alors
de morphotype ancestral).
Homologie vs. homoplasie [modifier]

Le groupe des "animaux à sang chaud" est polyphylétique.

On distingue deux types de ressemblances, l'homologie et l'homoplasie. Des caractères homologues
sont hérités d'un ancêtre commun, la relation liant deux homologues étant appelée homologie. À l'inverse,
l'homoplasie, terme introduit par Lankester en 1870, caractérise des ressemblances entre caractères n'étant
pas hérités d'un ancêtre commun. Différencier l'homoplasie de l'homologie est une tâche compliquée et
plusieurs critères définissent l'homologie dont le plus utilisé est le principe des connexions d'Étienne
Geoffroy Saint-Hilaire appelé aussi identité des connexions. Ce principe sera repris par Richard Owen.
d'après ce principe, sont homologues deux organes si quelles que soient leurs formes ou fonctions, ils ont
les même connexions avec d'autres organes. Ce principe peut aisément s'étendre aux séquences
moléculaires et l'alignement de séquences correspond à l'application de ce principe.
En réalité ce principe ne permet que de formuler des hypothèses dites d'homologie primaire. En effet
rien ne nous assure que le caractère supposé homologue l'est effectivement et que ce n'est pas de
l'homoplasie tant que l'hypothèse d'homologie n'est pas testée. Le test effectué est le test de congruence
selon Hennig ou de parcimonie. En cladistique on cherche à minimiser le nombre d'hypothèses
surnuméraires ou ad hoc. Une hypothèse ad hoc en reconstruction phylogénétique étant l'apparition
indépendante d'un caractère dans deux lignées différentes, c'est-à-dire de l'homoplasie. À l'inverse,
l'hypothèse de base est que le même caractère observé chez deux taxons indique une relation de parenté.
En maximisant alors l'homologie (ou en minimisant l'homoplasie, ce qui constitue deux démarches
équivalentes), on obtiendra un test des hypothèses primaires d'homologie. On retiendra alors le
cladogramme maximisant l'homologie. Les caractères apparaissant une fois dans le cladogramme résultant
de l'analyse de parcimonie seront considérés comme effectivement homologues et donc hérités d'un ancêtre
commun. L'homologie secondaire n'est donc plus seulement une homologie déduite du principe des
connexions mais une homologie de parenté ou de descendance.
À l'inverse, certains caractères peuvent se révéler homoplastiques une fois l'arbre reconstitué. Ces
caractères impliquent au moins deux pas évolutifs ou deux hypothèses de transformation dans le
cladogramme résultant. L'homoplasie est donc une hypothèse d'homologie primaire rejetée. Il existe en fait
deux types d'homoplasie: la Convergence évolutive et la réversion, décelables entre elles seulement sur
l'arbre enraciné le plus parcimonieux. La convergence indique que le même caractère (au sens des
connexions et non au sens évolutif) est apparu au moins deux fois dans le groupe considéré. À l'inverse, la
réversion est la perte secondaire d'un caractère, c'est-à-dire le retour à l'état primitif. Par exemple l'état
primitif des membres pairs chez les Vertébrés est l'absence. L'état dérivé chez les Gnathostomes est leur
présence. Or ce caractère est perdu chez les Gymnophiones ou les Serpents. Une réversion est une
homoplasie qui s'effectue sur une même morphocline (suite de transformations) alors que la convergence
s'effectue sur plusieurs morphoclines distinctes. L'homoplasie n'étant pas un caractère hérité par un ancêtre
commun, elle ne nous renseigne pas sur les relations de parenté. Un groupe formé sur une homoplasie est
appelé groupe polyphylétique.
Dans certains cas il n'y a aucun élément pour savoir si l'homoplasie est une réversion ou une
convergence, on parle alors de synapomorphie ambigüe[1]. À partir du cladogramme, le choix de savoir si le
caractère est issu d'une convergence ou d'une réversion est arbitraire[2]. L'hypothèse ACCTRAN
(Accelerated transformation) suppose des réversions. L'hypothèse DELTRAN (Delayed transformation)
suppose des convergences. Si le choix de l'un ou l'autre n'est pas important en moléculaire, le choix est bien
moins anodin pour les phylogénies morphologiques car le choix de l'une ou l'autre des hypothèses change
l'interprétation des caractères. Cependant le choix de l'un ou l'autre n'influence pas les relations de parentés.
L'image ci dessous l'illustre pour deux arbres ayant la même répartition des caractères, on a deux
configurations des transformations différentes.
 Illustration d'une synapomorphie ambigüe.

Lecture d'un cladogramme [modifier]

En général les possessions de caractères sont résumées dans un tableau appelé matrice
taxons/caractères donnant pour chaque caractère de chaque taxon son état. Ici nous allons utiliser un
exemple hypothétique. Par convention, pour chaque caractère "x", l'état primitif est noté x et l'état dérivé x'
(ça aurait pu être l'inverse). Tous les caractères en x sont donc plésiomorphes (exemple b) et les caractères
en x' son apomorphes (exemple b'). Voici la matrice hypothétique:
 taxon taxon taxon taxon taxon taxon
caractère
A B C D E F

Caractère
a a a' a' a' a'
"a"

Caractère
b b b b b' b
"b"

Caractère
c' c c c c' c'
"c"

Caractère
d d' d' d' d d'
"d"

Caractère
e e e e e e
"e"

On considère l'arbre suivant pour les taxons A, B, C, D, E et F: (A(B((C, D)(E, F)))). Cet arbre est choisit
arbitrairement. Ici R=A+B+C+D+E+F: c'est la racine ou le nœud qui contient tout. H=B+C+D+E+F
G=C+D+E+F I=C+D et J=E+F, donc G=I+J. Les taxons H, G, I et J sont des nœuds internes (Ce sont des
taxons au même titre qu'A, B, C etc.). L'ordre dans lequel les taxons sont disposés ne change pas la lecture
de l'arbre. En effet l'arbre (A(B((C, D)(E, F)))) est équivalent à l'arbre ((B((D, C)(E, F)))A)par exemple.
 La position la plus parcimonieuse des caractères (en minimisant le nombre de fois où ils
apparaissent) est représentée dans l'image associée par des barres rouges où x →x' indique le passage de
l'état x à x' et x'→x le passage de l'état x' à l'état x. Les barres rouges représentent des transformations sur
l'arbre ou "pas évolutifs".
 Exemple de cladogramme.

Selon la disposition la plus parcimonieuse des caractères, le caractère a' est commun à C, D, E, et F, c'est
donc une synapomorphie de G.
Le caractère b' n'apparaît que pour un seul taxon terminal : C. C'est donc une autapomorphie de C,
c'est-à-dire un caractère dérivé du taxon C uniquement, qui ne nous renseigne pas sur ses relations de
parentés avec les autres taxons.
Le caractère "e" reste à l'état e pour tous les taxons, c'est donc une symplésiomorphie de R. Les
caractères "c" et "d" subissent deux transformations (deux pas) dans l'arbre, ce sont donc des homoplasies.
Sur la ligne bleue (allant de la racine à E), le caractère "d" est impliqué deux fois sous l'état de
caractère d, c'est donc une réversion. Pour le caractère "d" on a alors la morphocline, ou suite de
transformations suivante : d→d'→d. L'état de caractère d possédé par E n'est donc pas réellement primitif
mais semblable à l'état primitif.
Le caractère "c" apparaît dans deux morphoclines différentes sous l'état c', c'est donc une
convergence. Les deux états de "c" pour A et J ne sont donc pas des homologies secondaires : ce n'est pas
le même caractère hérité d'un ancêtre commun.
Sur cet arbre on peut compter le nombre de transformations, il y en a 6. On compte donc 6 pas.
Deux groupes frères sont les groupes les plus apparentés entre eux qu'ils ne le sont de n'importe
quel autre. Ici par exemple C est groupe frère de D, B groupe frère de G, J groupe frère de I, etc.
 Un cladogramme représente une hiérarchie, au sens mathématique du terme, c'est à dire un
ensemble de classes strictement incluses les unes dans les autres. En effet, ici C et D appartiennent à I et
tous les groupes appartiennent à R (R étant la racine donc la classe comprenant tout). Un cladogramme peut
donc être représenté en diagramme de Venn. L'image ci-dessous représente le cladogramme utilisé dans
l'exemple sous forme de diagramme de Venn.
 Cladogramme précédent sous forme de diagramme de Venn illustrant la structure hiérarchique des
relations entre taxons
Enracinement et groupe externe [modifier]
La principale manière de polariser un caractère qui est utilisée est l'utilisation d'un groupe externe ou
extra-groupe (out-group) [3]. On estime que tout état de caractère présent en dehors du groupe interne (ou
in-group) est antérieur au groupe interne car déjà présent ailleurs. On va donc considérer ce caractère
comme primitif. Tout état de caractère présent en dehors du groupe d'étude est donc considéré comme
primitif et tout état qui n'est présent que dans le groupe d'étude est considéré comme dérivé, car propre au
groupe d'étude. Au lieu d'utiliser cette procédure pour chaque caractère, on utilise un groupe entier en
considérant tous ces caractères. Ce groupe doit être considéré à priori comme externe du groupe d'étude.
Par exemple pour une phylogénie des tétrapodes on pourra utiliser un téléostéen ou pour une phylogénie
des gastéropodes on pourra utiliser un céphalopode ou un bivalve. Le groupe externe ne doit pas être trop
éloigné du groupe d'étude sinon les caractères ne pourront même pas être comparés. A l'inverse, s'il est trop
proche le groupe externe peut en fait faire partie du groupe interne. De plus, si certains états de caractères
du groupe externe sont homoplastiques avec ceux du groupe interne, la polarisation de certains caractères
peut être fausse. Par exemple si on polarise une phylogénie des mammifères avec seulement des oiseaux,
on peut considérer que l'état de caractère "aile" est primitif pour le membre antérieur des mammifères, ce qui
est bien évidement faux. Le bon choix du groupe externe est donc crucial et un mauvais choix peut entrainer
des incohérences.
Le groupe externe permet alors d'enraciner l'arbre en l'orientant. Dans l'image ci-dessous, l'arbre de
départ n'est pas enraciné alors que les deux arbres d'en dessous (avec soit A, soit B en groupe externe) sont
enracinés. Les relations de parentés et le sens des transformations de caractères ne peuvent donc être
inférées qu'à partir d'un arbre enraciné. On remarque que la topologie de l'arbre change considérablement
selon le groupe externe utilisé. Les représentations en diagramme de Venn (en bas) sont les même que les
cladogrammes en forme d'arbres (en haut) respectivement à droite et à gauche. On constate que les
structures d'inclusions sont différentes. L'arbre de gauche et de droite peuvent aussi être représentés en
parenthétique: (A(B(C(D, E)))) à gauche et (C(A, B)(D, E)) à droite.
 L'arbre du haut n'est pas enraciné. L'arbre de gauche est enraciné en prenant A comme groupe
externe. L'arbre de droite est enraciné en prenant C comme groupe externe.

Cladistique et évolution [modifier]

Il est important de bien distinguer deux concepts différents, le "pattern" et le "process". La cladistique
permet de comprendre le "Pattern" ou structure de parenté entre les espèces actuelles ou fossiles, c'est-à-
dire leurs relations de parentés, tandis que l'évolution répond à la notion de processus évolutif ou "Process",
c'est-à-dire comment les caractères des espèces évoluent et sont acquis (ex : la sélection naturelle). Dès
lors les cladogrammes représentent uniquement des classifications entre espèces ou taxons. Il est
cependant évident que les caractères, qui permettent de faire des clades, sont le résultat de mécanismes
évolutifs mais la cladistique ne cherche ni à les expliquer, ni à les connaître. Seuls les liens de parenté sont
recherchés en cladistique. Cependant le cladogramme résultant de l'analyse permettra d'interpréter
l'évolution des clades et des caractères. En effet, le clade étant une unité évolutive, connaître la structure de
parenté de taxon permet d'interpréter la distribution des caractères au cours du temps, et donc l'évolution de
ces caractères.

Parcimonie, compatibilité et 3ia [modifier]

Différents principes peuvent être appliqués dans le cadre de reconstructions phylogénétiques
cladistiques. Le principe de parcimonie, selon lequel l'explication la plus simple (ici l'arbre ayant le moins de
pas évolutifs) doit être retenue est très majoritairement utilisée, mais l'on compte aussi la méthode de
compatibilité (aujourd'hui tombée en désuétude) et l'analyse à trois éléments (three-items analysis, ou 3ia)
[4].

La parcimonie [modifier]
La philosophie derrière les méthodes de parcimonie se fonde sur un principe attribué à Guillaume
d'Ockham, selon lequel les entités ne doivent pas être multipliées par delà ce qui est nécessaire ( « Entia non
sunt multiplicanda praeter necessitatem »). Pour faire un choix entre deux arbres, l'on va donc considérer
que le plus court, i.e. l'arbre avec le moins de pas évolutifs, est l'arbre représentant l'hypothèse
phylogénétique la plus plausible. L'on distingue trois modèles de parcimonie, selon la considération des
convergences et des réversions :
• La parcimonie de Wagner, où convergences et réversions sont acceptées.
• La parcimonie de Camin-Sokal admet les convergences mais pas les réversions.
• La parcimonie de Dollo accepte les réversions mais pas les convergences.
À titre d'exemple, l'absence de fenêtre antéorbitaire chez les crocodiles actuels est considéré comme
une réversion (elle était présente chez leurs ancêtres putatifs) ; les topologies similaires du bassin des
oiseaux et des ornitischiens, ou encore les formes hydrodynamiques des delphinidés et de la plupart des
lamniformes sont considérées comme des convergences.

L'analyse à trois éléments [modifier]

L'analyse à trois éléments ou 3ia (Three item analysis, parfois aussi appelée TTS pour Three-Taxon
Statements) est une méthode cladistique qui reconstitue les hypothèses de parentés en décomposant les
hypothèses d'homologie en hypothèses minimales d'homologie, c'est-à-dire en sous-arbres minimaux (à trois
éléments en cherchant "si un premier taxon est plus proche d'un deuxième que d'un troisième", d'où le nom
de la méthode) appelés 3is pour "three item statements" ou assertions à trois éléments. Cette méthode a
d'abord été utilisée pour la biogéographie cladistique et a ensuite été utilisée pour l'étude cladistique des
taxons. La 3ia part d'hypothèses d'homologies sous forme d'arbres racinés représentant les relations entre
caractères homologues qui sont ensuite décomposés en sous arbres, cette méthode ne nécessite donc pas
de groupe externe mais demande de connaitre à priori les états dérivés ou primitifs des caractères. On parle
aussi parfois d'analyse à trois taxons.

La compatibilité [modifier]
Cette méthode, largement tombée dans l'obsolescence, va qualifier les caractères pour lesquelles il
existe un arbre montrant les différents états de caractère sans comporter d'homoplasie de caractères
mutuellement compatibles. Dans une analyse phylogénétique, l'ensemble des caractères mutuellement
compatibles formera une clique. L'arbre retenu sera construit à partir de la clique la plus importante et sera
dépourvu de caractères homoplastiques.

Voir aussi [modifier]

• Caractère ancestral et caractère dérivé
• Willi Hennig
• Monophylie
• Paraphylie
• Polyphylie
• Synapomorphie
 • Autapomorphie

Bibliographie [modifier]
• P. Tassy, La renaissance de la systématique, 2001 En ligne (fr)
• H. Le Guyader, G. Lecointre, Classification phylogénétique du vivant 3ème édition, Belin,
2001.
• P. Tassy L'arbre à remonter le temps, Christian Bourgois, Paris, 1991.
• P. Darlu & P. Tassy, La reconstruction phylogénétique. Concepts et Méthodes , Masson,
1993 En ligne (fr)
• W. Hennig, Phylogenetic Systematics, Illinois University Press, 1966, traduit par D. Dwight
Davis & R. Zangerl.
• I.J. Kitching, P.L. Forey, C.J. Humphries et D.M. Williams, Cladistics, Oxford, Oxford
University Press, 1998 (2e éd.).
• C. Patterson, Morphological characters and homology, in K.A. Joysey et A.E. Friday (éd.),
Problems in Phylogenetic Reconstruction, Londres, Academic Press, 1982.
• K. de Queiroz et J.A. Gauthier, « Phylogenetic taxonomy », Annual Review of Ecology and
Systematics no 23, 1992, p. 449–480.
• D.L. Swofford, G.J. Olsen, P.J. Waddell et D.M. Hillis, Phylogenetic inference, in D.M. Hillis,
C. Moritz et B.K. Mable (éd.), Molecular Systematics, Sunderland (Massachusetts), Sinauer
Associates, 1996.
• E.O. Wiley, Phylogenetics : The Theory and Practice of Phylogenetic Systematics.-, New
York, Wiley Interscience, 1981.
Autres références [modifier]
1. ↑ M.G.G. de Pinna, Concepts and tests of homology in the cladistic paradigm, Cladistics,
1991.En ligne (en) [archive]
2. ↑ I. Agnarson & J. A. Miller Is ACCTRAN better than DELTRAN?, Cladistics, 2008 En ligne
(en) [archive]
3. ↑ V. Barriel & P. Tassy Rooting with Multiple Outgroup: Consensus Versus Parsimony ,
Cladistics, 1998
4. ↑ R. Zaragüeta-Bagils & E. Bourdon, Three-item analysis: Hierarchical representation and
treatment of missing and inapplicable data, Comptes Rendus Palevol, Volume 6, Issues 6-7,
November 2007, Pages 527-534

Liens externes [modifier]

• (en) The Willi Hennig Society website

• Portail de la biologie

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Cladistique ».

Catégorie : Phylogénétique | [+]

 W000

Evolutionary developmental biology

From Wikipedia, the free encyclopedia

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Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics

Biology portal · v • d • e
Evolutionary developmental biology (evolution of development or informally, evo-devo) is a field of
biology that compares the developmental processes of different organisms to determine the ancestral
relationship between them, and to discover how developmental processes evolved. It addresses the origin
and evolution of embryonic development; how modifications of development and developmental processes
lead to the production of novel features, such as the evolution of feathers;[1] the role of developmental
plasticity in evolution; how ecology impacts in development and evolutionary change; and the developmental
basis of homoplasy and homology.[2]
Although interest in the relationship between ontogeny and phylogeny extends back to the nineteenth
century, the contemporary field of evo-devo has gained impetus from the discovery of genes regulating
embryonic development in model organisms. General hypotheses remain hard to test because organisms
differ so much in shape and form.[3] Nevertheless, it now appears that just as evolution tends to create new
genes from parts of old genes (molecular economy), evo-devo demonstrates that evolution alters
developmental processes (genes and gene networks) to create new and novel structures from the old gene
networks (such as bone structures of the jaw deviating to the ossicles of the middle ear) or will conserve
(molecular economy) a similar program in a host of organisms such as eye development genes in molluscs,
insects, and vertebrates [4] [5]. Initially the major interest has been in the evidence of homology in the cellular
and molecular mechanisms that regulate body plan and organ development. However more modern
approaches include developmental changes associated with speciation.[6]
Contents
[hide]
• 1 Basic principles
• 2 History
• 3 The developmental-genetic toolkit
• 4 Development and the origin of novelty
• 5 See also
• 6 References
• 6.1 Sources
• 7 Further reading
• 8 External links

[edit] Basic principles

Charles Darwin's theory of evolution is based on three principles: natural selection, heredity, and
variation. At the time that Darwin wrote, the principles underlying heredity and variation were poorly
understood. In the 1940s, however, biologists incorporated Gregor Mendel's principles of genetics to explain
both, resulting in the modern synthesis. It was not until the 1980s and 1990s, however, when more
comparative molecular sequence data between different kinds of organisms was amassed and detailed, that
an understanding of the molecular basis of the developmental mechanisms has arisen. Currently, it is well
understood how genetic mutation occurs. However, developmental mechanisms are not understood
sufficiently to explain which kinds of phenotypic variation can arise in each generation from variation at the
genetic level. Evolutionary developmental biology studies how the dynamics of development determine the
phenotypic variation arising from genetic variation and how that affects phenotypic evolution (specially its
direction). At the same time evolutionary developmental biology also studies how development itself evolves.
Thus, the origins of evolutionary developmental biology come from both an improvement in molecular biology
techniques as applied to development, and the full appreciation of the limitations of classic neo-Darwinism as
applied to phenotypic evolution. Some evo-devo researchers see themselves as extending and enhancing
the modern synthesis by incorporating into it findings of molecular genetics and developmental biology.
Others, drawing on findings of discordances between genotype and phenotype and epigenetic mechanisms
of development, are mounting an explicit challenge to neo-Darwinism.
Evolutionary developmental biology is not yet a unified discipline, but can be distinguished from
earlier approaches to evolutionary theory by its focus on a few crucial ideas. One of these is modularity: as
has been long recognized, plants and animal bodies are modular: they are organized into developmentally
and anatomically distinct parts. Often these parts are repeated, such as fingers, ribs, and body segments.
Evo-devo seeks the genetic and evolutionary basis for the division of the embryo into distinct modules, and
for the partly independent development of such modules.
Another central idea is that some gene products function as switches whereas others act as diffusible
signals. Genes specify proteins, some of which act as structural components of cells and others as enzymes
that regulate various biochemical pathways within an organism. Most biologists working within the modern
synthesis assumed that an organism is a straightforward reflection of its component genes. The modification
of existing, or evolution of new, biochemical pathways (and, ultimately, the evolution of new species of
organisms) depended on specific genetic mutations. In 1961, however, Jacques Monod, Jean-Pierre
Changeux and François Jacob discovered within the bacterium Escherichia coli a gene that functioned only
when "switched on" by an environmental stimulus.[7] Later, scientists discovered specific genes in animals,
including a subgroup of the genes which contain the homeobox DNA motif, called Hox genes, that act as
switches for other genes, and could be induced by other gene products, morphogens, that act analogously to
the external stimuli in bacteria. These discoveries drew biologists' attention to the fact that genes can be
selectively turned on and off, rather than being always active, and that highly disparate organisms (for
example, fruit flies and human beings) may use the same genes for embryogenesis (e.g., the genes of the
"developmental-genetic toolkit", see below), just regulating them differently.
Similarly, organismal form can be influenced by mutations in promoter regions of genes, those DNA
sequences at which the products of some genes bind to and control the activity of the same or other genes,
not only protein-specifying sequences. In addition to providing new support for Darwin's assertion that all
organisms are descended from a common ancestor, this finding suggested that the crucial distinction
between different species (even different orders or phyla) may be due less to differences in their content of
gene products than to differences in spatial and temporal expression of conserved genes. The implication
that large evolutionary changes in body morphology are associated with changes in gene regulation, rather
than the evolution of new genes, suggested that the action of natural selection on promoters responsive to
Hox and other "switch" genes may play a major role in evolution.
Another focus of evo-devo is developmental plasticity, the basis of the recognition that organismal
phenotypes are not uniquely determined by their genotypes. If generation of phenotypes is conditional, and
dependent on external or environmental inputs, evolution can proceed by a "phenotype-first" route,[3][8] with
genetic change following, rather than initiating, the formation of morphological and other phenotypic
novelties. The case for this was argued for by Mary Jane West-Eberhard in her 2003 book Developmental
plasticity and evolution.[8]
[edit] History
An early version of recapitulation theory, also called the biogenetic law or embryological parallelism,
was put forward by Étienne Serres in 1824–26 as what became known as the "Meckel-Serres Law" which
attempted to provide a link between comparative embryology and a "pattern of unification" in the organic
world. It was supported by Étienne Geoffroy Saint-Hilaire as part of his ideas of idealism, and became a
prominent part of his version of Lamarckism leading to disagreements with Georges Cuvier. It was widely
supported in the Edinburgh and London schools of higher anatomy around 1830, notably by Robert Edmond
Grant, but was opposed by Karl Ernst von Baer's embryology of divergence in which embryonic parallels only
applied to early stages where the embryo took a general form, after which more specialised forms diverged
from this shared unity in a branching pattern. The anatomist Richard Owen used this to support his idealist
concept of species as showing the unrolling of a divine plan from an archetype, and in the 1830s attacked the
transmutation of species proposed by Lamarck, Geoffroy and Grant.[9] In the 1850s Owen began to support
an evolutionary view that the history of life was the gradual unfolding of a teleological divine plan,[10] in a
continuous "ordained becoming", with new species appearing by natural birth.[11]
In On the Origin of Species (1859), Charles Darwin proposed evolution through natural selection, a
theory central to modern biology. Darwin recognised the importance of embryonic development in the
understanding of evolution, and the way in which von Baer's branching pattern matched his own idea of
descent with modification[12]:
“ We can see why characters derived from the embryo should be of equal
importance with those derived from the adult, for a natural classification of course
includes all ages.[13] ”
 Ernst Haeckel (1866), in his endeavour to produce a synthesis of Darwin's theory with Lamarckism
and Naturphilosophie, proposed that "ontogeny recapitulates phylogeny," that is, the development of the
embryo of every species (ontogeny) fully repeats the evolutionary development of that species (phylogeny),
in Geoffroy's linear model rather than Darwin's idea of branching evolution.[12] Haeckel's concept explained,
for example, why humans, and indeed all vertebrates, have gill slits and tails early in embryonic
development. His theory has since been discredited. However, it served as a backdrop for a renewed interest
in the evolution of development after the modern evolutionary synthesis was established (roughly 1936 to
1947).
Stephen Jay Gould called this approach to explaining evolution as terminal addition; as if every
evolutionary advance was added as new stage by reducing the duration of the older stages. The idea was
based on observations of neoteny.[14] This was extended by the more general idea of heterochrony
(changes in timing of development) as a mechanism for evolutionary change.[15]
D'Arcy Thompson postulated that differential growth rates could produce variations in form in his
1917 book On Growth and Form. He showed the underlying similarities in body plans and how geometric
transformations could be used to explain the variations.
Edward B. Lewis discovered homeotic genes, rooting the emerging discipline of evo-devo in
molecular genetics. In 2000, a special section of the Proceedings of the National Academy of Sciences
(PNAS) was devoted to "evo-devo",[16] and an entire 2005 issue of the Journal of Experimental Zoology Part
B: Molecular and Developmental Evolution was devoted to the key evo-devo topics of evolutionary innovation
and morphological novelty.[17]
[edit] The developmental-genetic toolkit
The developmental-genetic toolkit consists of a small fraction of the genes in an organism's genome
whose products control its development. These genes are highly conserved among Phyla. Differences in
deployment of toolkit genes affect the body plan and the number, identity, and pattern of body parts. The
majority of toolkit genes are components of signaling pathways, and encode for the production of
transcription factors, cell adhesion proteins, cell surface receptor proteins, and secreted morphogens, all of
these participate in defining the fate of undifferentiated cells, generating spatial and temporal patterns, which
in turn form the body plan of the organism. Among the most important of the toolkit genes are those of the
Hox gene cluster, or complex. Hox genes, transcription factors containing the more broadly distributed
homeobox protein-binding DNA motif, function in patterning the body axis. Thus, by combinatorial specifying
the identity of particular body regions, Hox genes determine where limbs and other body segments will grow
in a developing embryo or larva. A paragon of a toolbox gene is Pax6/eyeless, which controls eye formation
in all animals. It has been found to produce eyes in mice and Drosophila, even if mouse Pax6/eyeless was
expressed in Drosophila.[18]
This means that a big part of the morphological evolution undergone by organisms is a product of
variation in the genetic toolkit, either by the genes changing their expression pattern or acquiring new
functions. A good example of the first is the enlargement of the beak in Darwin's Large Ground-finch
(Geospiza magnirostris), in which the gene BMP is responsible for the larger beak of this bird, relative to the
other finches.[19]
The loss of legs in snakes and other squamates is another good example of genes changing their
expression pattern. In this case the gene Distal-less is very under-expressed, or not expressed at all, in the
regions where limbs would form in other tetrapods.[20] This same gene determines the spot pattern in
butterfly wings,[21] which shows that the toolbox genes can change their function.
 Toolbox genes, as well as being highly conserved, also tend to evolve the same function
convergently or in parallel. Classic examples of this are the already mentioned Distal-less gene, which is
responsible for appendage formation in both tetrapods and insects, or, at a finer scale, the generation of wing
patterns in the butterflies Heliconius erato and Heliconius melpomene. These butterflies are Müllerian mimics
whose coloration pattern arose in different evolutionary events, but is controlled by the same genes.[22] The
previous supports Kirschner and Gerhardt's theory of Facilitated Variation, which states that morphological
evolutionary novelty is generated by regulatory changes in various members of a large set of conserved
mechanisms of development and physiology.[23]

[edit] Development and the origin of novelty

Among the more surprising and, perhaps, counterintuitive (from a neo-Darwinian viewpoint) results of
recent research in evolutionary developmental biology is that the diversity of body plans and morphology in
organisms across many phyla are not necessarily reflected in diversity at the level of the sequences of
genes, including those of the developmental genetic toolkit and other genes involved in development.
Indeed, as Gerhart and Kirschner have noted, there is an apparent paradox: "where we most expect to find
variation, we find conservation, a lack of change".[24]
Even within a species, the occurrence of novel forms within a population does not generally correlate
with levels of genetic variation sufficient to account for all morphological diversity. For example, there is
significant variation in limb morphologies amongst salamanders and in differences in segment number in
centipedes, even when the respective genetic variation is low.
A major question then, for evo-devo studies, is: If the morphological novelty we observe at the level
of different clades is not always reflected in the genome, where does it come from? Apart from neo-
Darwinian mechanisms such as mutation, translocation and duplication of genes, novelty may also arise by
mutation-driven changes in gene regulation. The finding that much biodiversity is not due to differences in
genes, but rather to alterations in gene regulation, has introduced an important new element into evolutionary
theory.[25] Diverse organisms may have highly conserved developmental genes, but highly divergent
regulatory mechanisms for these genes. Changes in gene regulation are "second-order" effects of genes,
resulting from the interaction and timing of activity of gene networks, as distinct from the functioning of the
individual genes in the network.
The discovery of the homeotic Hox gene family in vertebrates in the 1980s allowed researchers in
developmental biology to empirically assess the relative roles of gene duplication and gene regulation with
respect to their importance in the evolution of morphological diversity. Several biologists, including Sean B.
Carroll of the University of Wisconsin–Madison suggest that "changes in the cis-regulatory systems of genes"
are more significant than "changes in gene number or protein function".[26] These researchers argue that the
combinatorial nature of transcriptional regulation allows a rich substrate for morphological diversity, since
variations in the level, pattern, or timing of gene expression may provide more variation for natural selection
to act upon than changes in the gene product alone.
Epigenetic alterations of gene regulation or phenotype generation that are subsequently consolidated
by changes at the gene level constitute another class of mechanisms for evolutionary innovation. Epigenetic
changes include modification of the genetic material due to methylation and other reversible chemical
alteration [27], as well as nonprogrammed remolding of the organism by physical and other environmental
effects due to the inherent plasticity of developmental mechanisms.[8] The biologists Stuart A. Newman and
Gerd B. Müller have suggested that organisms early in the history of multicellular life were more susceptible
to this second category of epigenetic determination than are modern organisms, providing a basis for early
macroevolutionary changes.[28]
[edit] See also
Evolutionary biology portal

• Developmental biology
• Animal evolution
• Plant Evolutionary Developmental Biology
• Baldwin effect
• Developmental systems theory
• Evolution of multicellularity
• Genetic assimilation
• Ontogeny
• Ontogeny recapitulates phylogeny
• List of gene families
• Important publications in evolutionary developmental biology
• Evolution and Development Leading journal
• Body plan
• Cell signaling
• Signal transduction
• Cell signaling networks
• Transcription factor
• Enhancer
 • Enhanceosome
• promoter (biology)
• Gene regulatory network

[edit] References
1. ^ Prum, R.O., Brush, A.H. (March 2003). "Which Came First, the Feather or the Bird?".
Scientific American 288 (3): 84–93. doi:10.1038/scientificamerican0303-84. PMID 12616863.
2. ^ Hall, Brian K. (2000). "Evo-devo or devo-evo—does it matter". Evolution & Development 2:
177–178. doi:10.1046/j.1525-142x.2000.00003e.x.
3. ^ a b Palmer, RA (2004). "Symmetry breaking and the evolution of development". Science
306 (5697): 828–833. doi:10.1126/science.1103707. PMID 15514148.
4. ^ Tomarev, Stanislav I.; Callaerts, Patrick; Kos, Lidia; Zinovieva, Rina; Halder, Georg;
Gehring, Walter; Piatigorsky, Joram (1997). "Squid Pax-6 and eye development". Proceedings of the
National Academy of Sciences 94 (6): 2421–2426. doi:10.1073/pnas.94.6.2421.
http://www.pnas.org/content/94/6/2421.full.
5. ^ Pichaud, Franck; Desplan, Claude (August 2002). "Pax genes and eye organogenesis".
Current opinion in genetics and development 12 (4): 430–434. doi:10.1016/S0959-437X(02)00321-0.
PMID 12100888.
6. ^ Pennisi, E (2002). "EVOLUTIONARY BIOLOGY:Evo-Devo Enthusiasts Get Down to
Details". Science 298 (5595): 953–955.. doi:10.1126/science.298.5595.953. PMID 12411686.
7. ^ Monod, J; Changeux, JP; Jacob, F (1963). "Allosteric proteins and cellular control
systems". Journal of Molecular Biology 6: 306–329. doi:10.1016/S0022-2836(63)80091-1.
PMID 13936070.
 8. ^ a b c West-Eberhard, M-J. (2003). Developmental plasticity and evolution. New York:
Oxford University Press. ISBN 978-0-19-512235-0.
9. ^ Desmond 1989, pp. 53–53, 86–88, 337–340
Secord 2003, pp. 252–253
10.^ Bowler 2003, pp. 120–128, 208
Secord 2003, pp. 424, 512
11.^ Desmond & Moore 1991, pp. 490–491
12.^ a b Bowler 2003, p. 170, 190-191
13.^ Darwin, Charles (1859). On the Origin of Species. London: John Murray. pp. 439–430.
ISBN 0801413192.
14.^ Ridley, Mark (2003). Evolution. Wiley-Blackwell. ISBN 978-1-4051-0345-9.
http://www.blackwellpublishing.com/ridley/.
15.^ Gould, Stephen Jay (1977). Ontogeny and Phylogeny. Cambridge, Massachusetts:
Harvard University Press. ISBN 0-674-63940-5.
16.^ Goodman CS and Coughlin BS (Eds). (2000). "Special feature: The evolution of evo-devo
biology". Proceedings of the National Academy of Sciences 97 (9): 4424–4456.
doi:10.1073/pnas.97.9.4424. PMID 10781035. PMC 18255.
http://www.pnas.org/cgi/content/full/97/9/4424.
17.^ Müller GB and Newman SA (Eds.) (2005). "Special issue: Evolutionary Innovation and
Morphological Novelty". Journal of Exp. Zool. Part B: Molecular and Developmental Evolution 304B:
485–631. http://www3.interscience.wiley.com/cgi-bin/jissue/112149101.
18.^ Xu, P.X., Woo, I., Her, H., Beier, D.R., Maas, R.L. (1997). "Mouse Eya homologues of the
Drosophila eyes absent gene require Pax6 for expression in lens and nasal placode". Development
124 (1): 219–231. PMID 9006082.
 19.^ Abzhanov, A.; Protas, M.; Grant, B.R.; Grant, P.R.; Tabin, C.J. (2004). "Bmp4 and
Morphological Variation of Beaks in Darwin's Finches". Science 305 (5689): 1462–1465.
doi:10.1126/science.1098095. PMID 15353802.
20.^ Cohn, M.J.; Tickle, C. (1999). "Developmental basis of limblessness and axial patterning in
snakes.". Nature 399 (6735): 474–479. doi:10.1038/20944. PMID 10365960.
21.^ Beldade, P.; Brakefield, P.M.; Long, A.D. (2002). "Contribution of Distal-less to quantitative
variation in butterfly eyespots". Nature 415 (6869): 315–318. doi:10.1038/415315a. PMID 11797007.
22.^ Baxter, S.W.; Papa, R.; Chamberlain, N.; Humphray, S.J.; Joron, M.; Morrison, C.; ffrench-
Constant, R.H.; McMillan, W.O.; Jiggins, C.D. (2008). "Convergent Evolution in the Genetic Basis of
Mullerian Mimicry in Heliconius Butterflies". Genetics 180 (3): 1567–1577.
doi:10.1534/genetics.107.082982. PMID 18791259.
23.^ Gerhart, John; Kirschner, Marc (2007). "The theory of facilitated variation". Proceedings of
the National Academy of Sciences 104 (suppl1): 8582–8589. doi:10.1073/pnas.0701035104.
PMID 17494755.
24.^ Gerhart, John; Kirschner, Marc (1997). Cells, Embryos and Evolution. Blackwell Science.
ISBN 978-0865425743.
25.^ Carroll, Sean B.; Grenier, Jennifer K.; Weatherbee, Scott D. (2005). From DNA to Diversity:
Molecular Genetics and the Evolution of Animal Design — Second Edition. Blackwell Publishing.
ISBN 1405119500.
26.^ Carroll, Sean B. (2000). "Endless forms: the evolution of gene regulation and morphological
diversity". Cell 101: 577–80. doi:10.1016/S0092-8674(00)80868-5.
27.^ Jablonka, Eva; Lamb, Marion (1995). Epigenetic Inheritance and Evolution: The
Lamarckian Dimension. Oxford, New York: Oxford University Press. ISBN 978-0198540632.
 28.^ Müller, Gerd B. and Newman, Stuart A., ed (2003). Origination of Organismal Form:
Beyond the Gene in Developmental and Evolutionary Biology . MIT Press.

[edit] Sources
• Bowler, Peter J. (2003). Evolution: the history of an idea. Berkeley: University of California
Press. ISBN 0-520-23693-9.
• Desmond, Adrian J. (1989). The politics of evolution: morphology, medicine, and reform in
radical London. Chicago: University of Chicago Press. ISBN 0-226-14374-0.
• Desmond, Adrian J.; Moore, James William (1991). Darwin. London: Michael Joseph.
ISBN 0-7181-3430-3.
• Secord, James A. (2003). Victorian sensation: the extraordinary publication, reception, and
secret authorship of Vestiges of the natural history of creation . Chicago: University of Chicago Press.
ISBN 0-226-74411-6.

[edit] Further reading

• Buss, Leo W. (1987). The Evolution of Individuality. Princeton University Press. ISBN 978-
0691084688.
• Carroll, Sean B. (2005). Endless Forms Most Beautiful: The New Science of Evo Devo and
the Making of the Animal Kingdom. Norton. ISBN 9780393060164.
• Goodwin, Brian (1994). How the Leopard Changed its Spots. Phoenix Giants.
ISBN 9780691088099.
 • Hall, Brian K. & Olsen, Wendy M., ed (2007). Keywords and Concepts in Evolutionary
Developmental Biology. New Delhi, India: Discovery Publishing House. ISBN 978-81-8356-256-0.
• Kirschner, Marc; Gerhart, John (2005). The Plausibility of Life: Resolving Darwin's Dilemma .
Yale University Press. ISBN 9780300108651.
• Laubichler, Manfred D. and Maienschein, Jane, ed (2007). From Embryology to Evo-Devo: A
History of Developmental Evolution. The MIT Press. ISBN 978-0-262-12283-2.
• Minelli, Alessandro (2003). The Development of Animal Form: Ontogeny, Morphology, and
Evolution. Cambridge University Press. ISBN 978-0521808514.
• Orr, H. Allen (2005-10-24). "Turned on: A revolution in the field of evolution?". The New
Yorker. http://www.newyorker.com/critics/books/articles/051024crbo_books1. Discussion of Carroll,
Endless Forms Most Beautiful
• Raff, Rudolf A. (1996). The Shape of Life: Genes, Development, and the Evolution of Animal
Form. The University of Chicago Press. ISBN 978-0226702667.
• Sommer, Ralf J. (2009). "The future of evo–devo: model systems and evolutionary theory".
Nature Reviews Genetics 10 (6): 416–422. doi:10.1038/nrg2567. PMID 19369972.

[edit] External links

• Scott F. Gilbert, The morphogenesis of evolutionary developmental biology
• Tardigrades (water bears) as evo-devo models, a short video from NPR's Science Friday
 [hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_developmental_biology"

Categories: Evolutionary biology | Developmental biology

• This page was last modified on 26 August 2010 at 05:38.

 W000

Génétique évolutive du développement

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

La génétique évolutive du développement souvent plus connue sous le nom d'évo-dévo (Evo-devo
pour Evolutionary developmental biology en anglais) est un champ disciplinaire en biologie de l'évolution qui
a pour objectif de comprendre l'origine de la complexité morphologique des organismes (plantes ou
animaux) à travers l'étude comparée des gènes qui régulent leur développement.
À l'intersection de la génomique, de la phylogénie moléculaire et de la génétique du développement,
l'évodévo est l'un des courants actuels les plus dynamiques en biologie de l'évolution.
Sommaire
[masquer]
• 1
Histoire
• 2 Voir
aussi
• 2
.1
Article
s
conne
xes
• 2
.2
Bibliog
raphie
• 2
.3
Liens
extern
es
• 2
Histoire [modifier]
Cette discipline est née dans les dernières décennies du XX e siècle mais elle s'inscrit dans la
continuité de l'embryologie comparative fondée au XIX e siècle.
Elle a conduit à la remise en cause de nombreuses conceptions issues en particulier de la nouvelle
synthèse.
Selon Alain Prochiantz, professeur au Collège de France, la découverte des gènes du
développement (le premier a été cloné en 1984) « a été un moment-clef de l’histoire de la biologie parce que
c’est une catégorie de gènes qui a permis de faire le lien entre la génétique, le développement et l’évolution.
En effet, les mutations de ces gènes permettent des changements massifs de forme des vivants. (...) Ces
gènes de développement permettent donc à la fois de comprendre l'ontogénèse et la phylogénèse. Grâce à
eux, une nouvelle discipline que l’on appelle Evo-dévo » [1].

Voir aussi [modifier]

Articles connexes [modifier]
• Génétique
• Génétique évolutive du développement des plantes
• Phylogénétique

Bibliographie [modifier]
• {...}
 Liens externes [modifier]
• {...}

Notes et références [modifier]

• Pour une discussion générale du cadre conceptuel dans lequel s'inscrit la génétique
évolutive du développement, on peut se reporter à l'ouvrage de Stephen Jay Gould, La Structure de
la théorie de l'évolution (Belknap/Harvard 2002) et en particulier au chapitre 10.
1. ↑ Ariane Poulantzas, entretien avec Alain Prochiantz, « Les bactéries pensent
aussi » [archive], La Vie des idées, 5 juin 2009
Ce document provient de « http://fr.wikipedia.org/wiki/G%C3%A9n%C3%A9tique_
%C3%A9volutive_du_d%C3%A9veloppement ».

Catégories : Embryologie | Génétique | Discipline de la biologie de l'évolution

 w000

Molecular evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics

Biology portal · v • d • e
Molecular evolution is the process of evolution at the scale of DNA, RNA, and proteins. Molecular
evolution emerged as a scientific field in the 1960s as researchers from molecular biology, evolutionary
biology and population genetics sought to understand recent discoveries on the structure and function of
nucleic acids and protein. Some of the key topics that spurred development of the field have been the
evolution of enzyme function, the use of nucleic acid divergence as a "molecular clock" to study species
divergence, and the origin of non-functional or junk DNA. Recent advances in genomics, including whole-
genome sequencing, high-throughput protein characterization, and bioinformatics have led to a dramatic
increase in studies on the topic. In the 2000s, some of the active topics have been the role of gene
duplication in the emergence of novel gene function, the extent of adaptive molecular evolution versus
neutral drift, and the identification of molecular changes responsible for various human characteristics
especially those pertaining to infection, disease, and cognition.

Contents
[hide]
• 1 Principles of molecular evolution
• 1.1 Mutations
• 1.2 Causes of change in allele frequency
• 1.3 Molecular study of phylogeny
• 2 The driving forces of evolution
• 3 Related fields
• 4 Key researchers in molecular evolution
• 5 Journals and societies
• 6 See also
• 7 References
• 8 Further reading
[edit] Principles of molecular evolution
[edit] Mutations
Main article: Mutation
Mutations are permanent, transmissible changes to the genetic material (usually DNA or RNA) of a
cell. Mutations can be caused by copying errors in the genetic material during cell division and by exposure
to radiation, chemicals, or viruses, or can occur deliberately under cellular control during the processes such
as meiosis or hypermutation. Mutations are considered the driving force of evolution, where less favorable (or
deleterious) mutations are removed from the gene pool by natural selection, while more favorable (or
beneficial) ones tend to accumulate. Neutral mutations do not affect the organism's chances of survival in its
natural environment and can accumulate over time, which might result in what is known as punctuated
equilibrium; the modern interpretation of classic evolutionary theory.

[edit] Causes of change in allele frequency

Main article: Population genetics
There are three known processes that affect the survival of a characteristic; or, more specifically, the
frequency of an allele (variant of a gene):
• Genetic drift describes changes in gene frequency that cannot be ascribed to selective
pressures, but are due instead to events that are unrelated to inherited traits. This is especially
important in small mating populations, which simply cannot have enough offspring to maintain the
same gene distribution as the parental generation.
 • Gene flow or Migration: or gene admixture is the only one of the agents that makes
populations closer genetically while building larger gene pools.
• Selection, in particular natural selection produced by differential mortality and fertility.
Differential mortality is the survival rate of individuals before their reproductive age. If they survive,
they are then selected further by differential fertility – that is, their total genetic contribution to the next
generation. In this way, the alleles that these surviving individuals contribute to the gene pool will
increase the frequency of those alleles. Sexual selection, the attraction between mates that results
from two genes, one for a feature and the other determining a preference for that feature, is also very
important.

[edit] Molecular study of phylogeny

Main articles: Molecular systematics and Phylogenetics
Molecular systematics is a product of the traditional field of systematics and molecular genetics. It is
the process of using data on the molecular constitution of biological organisms' DNA, RNA, or both, in order
to resolve questions in systematics, i.e. about their correct scientific classification or taxonomy from the point
of view of evolutionary biology.
Molecular systematics has been made possible by the availability of techniques for DNA sequencing,
which allow the determination of the exact sequence of nucleotides or bases in either DNA or RNA. At
present it is still a long and expensive process to sequence the entire genome of an organism, and this has
been done for only a few species. However, it is quite feasible to determine the sequence of a defined area of
a particular chromosome. Typical molecular systematic analyses require the sequencing of around 1000
base pairs.
[edit] The driving forces of evolution
Main articles: Neutral theory of molecular evolution, Modern evolutionary synthesis, and Mutationism
Depending on the relative importance assigned to the various forces of evolution, three perspectives
provide evolutionary explanations for molecular evolution.[1]
While recognizing the importance of random drift for silent mutations,[2] selectionists hypotheses
argue that balancing and positive selection are the driving forces of molecular evolution. Those hypotheses
are often based on the broader view called panselectionism, the idea that selection is the only force strong
enough to explain evolution, relaying random drift and mutations to minor roles.[1]
Neutralists hypotheses emphasize the importance of mutation, purifying selection and random
genetic drift.[3] The introduction of the neutral theory by Kimura,[4] quickly followed by King and Jukes' own
findings,[5] lead to a fierce debate about the relevance of neodarwinism at the molecular level. The Neutral
theory of molecular evolution states that most mutations are deleterious and quickly removed by natural
selection, but of the remaining ones, the vast majority are neutral with respect to fitness while the amount of
advantageous mutations is vanishingly small. The fate of neutral mutations are governed by genetic drift, and
contribute to both nucleotide polymorphism and fixed differences between species. [6][7][8]
Mutationists hypotheses emphasize random drift and biases in mutation patterns.[9] Sueoka was the
first to propose a modern mutationist view. He proposed that the variation in GC content was not the result of
positive selection, but a consequence of the GC mutational pressure.[10]
[edit] Related fields
An important area within the study of molecular evolution is the use of molecular data to determine
the correct biological classification of organisms. This is called molecular systematics or molecular
phylogenetics.
Tools and concepts developed in the study of molecular evolution are now commonly used for
comparative genomics and molecular genetics, while the influx of new data from these fields has been
spurring advancement in molecular evolution.

[edit] Key researchers in molecular evolution

Some researchers who have made key contributions to the development of the field:
• Motoo Kimura — Neutral theory
• Masatoshi Nei — Adaptive evolution
• Walter M. Fitch — Phylogenetic reconstruction
• Walter Gilbert — RNA world
• Joe Felsenstein — Phylogenetic methods
• Susumu Ohno — Gene duplication
• John H. Gillespie — Mathematics of adaptation
 [edit] Journals and societies
Journals dedicated to molecular evolution include Molecular Biology and Evolution, Journal of
Molecular Evolution, and Molecular Phylogenetics and Evolution. Research in molecular evolution is also
published in journals of genetics, molecular biology, genomics, systematics, or evolutionary biology. The
Society for Molecular Biology and Evolution publishes the journal "Molecular Biology and Evolution" and
holds an annual international meeting.

[edit] See also

• History of molecular • Genomic • Neutral theory of
evolution organization molecular evolution
• Chemical evolution • Horizontal gene • Nucleotide diversity
• Evolution transfer • Parsimony
• Genetic drift • Human evolution • Population genetics
• E. coli long-term • Evolution of • Selection
evolution experiment dietary antioxidants
• Evolutionary • Molecular clock
physiology • Comparative
phylogenetics

[edit] References
1. ^ a b Graur, D. and Li, W.-H. (2000). Fundamentals of molecular evolution. Sinauer.
 2. ^ Gillespie, J. H (1991). The Causes of Molecular Evolution. Oxford University Press, New
York. ISBN 0-19-506883-1.
3. ^ Kimura, M. (1983). The Neutral Theory of Molecular Evolution. Cambridge University
Press, Cambridge. ISBN 0-521-23109-4.
4. ^ Kimura, Motoo (1968). "Evolutionary rate at the molecular level". Nature 217 (5129): 624–
626. doi:10.1038/217624a0. PMID 5637732.
http://www2.hawaii.edu/~khayes/Journal_Club/fall2006/Kimura_1968_Nature.pdf.
5. ^ King, J.L. and Jukes, T.H. (1969). "Non-Darwinian Evolution". Science 164 (881): 788–798.
doi:10.1126/science.164.3881.788. PMID 5767777.
http://www.blackwellpublishing.com/ridley/classictexts/king.pdf.
6. ^ Nachman M. (2006). "Detecting selection at the molecular level" in: Evolutionary Genetics:
concepts and case studies. pp. 103–118.
7. ^ The nearly neutral theory expanded the neutralist perspective, suggesting that several
mutations are nearly neutral, which means both random drift and natural selection is relevant to their
dynamics.
8. ^ Ohta, T (1992). "The nearly neutral theory of molecular evolution". Annual Review of
Ecology and Systematics 23: 263–286. doi:10.1146/annurev.es.23.110192.001403.
9. ^ Nei, M. (2005). "Selectionism and Neutralism in Molecular Evolution". Molecular Biology
and Evolution 22(12) (12): 2318–2342. doi:10.1093/molbev/msi242. PMID 16120807.
10.^ Sueoka, N. (1964). "On the evolution of informational macromolecules". in In: Bryson, V.
and Vogel, H.J.. Evolving genes and proteins. Academic Press, New-York. pp. 479–496.
 [edit] Further reading
• Li, W.-H. (2006). Molecular Evolution. Sinauer. ISBN 0878934804.
• Lynch, M. (2007). The Origins of Genome Architecture. Sinauer. ISBN 0878934847.

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts
 Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Molecular_evolution"

Categories: Molecular evolution

 W000

Phylogenetics
From Wikipedia, the free encyclopedia

Jump to: navigation, search

"Phylogenesis" redirects here. For the science fiction novel, see Phylogenesis (novel).
Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics

Biology portal · v • d • e
In biology, phylogenetics is the study of evolutionary relatedness among various groups of organisms
(for example, species or populations), which is discovered through molecular sequencing data and
morphological data matrices. The term phylogenetics is of Greek origin from the terms phyle/phylon
(φυλή/φῦλον), meaning "tribe, race," and genetikos (γενετικός), meaning "relative to birth" from genesis
(γένεσις, "birth"). Taxonomy, the classification, identification, and naming of organisms, has been richly
informed by phylogenetics but remains methodologically and logically distinct.[1] The fields overlap however
in the science of phylogenetic systematics – often called "cladism" or "cladistics" –, where only phylogenetic
trees are used to delimit taxa, which represent groups of lineage-connected individuals.[2] In biological
systematics as a whole, phylogenetic analyses have become essential in researching the evolutionary tree of
life.

Contents
[hide]
• 1 Construction of a phylogenetic tree
• 2 Grouping of organisms
• 3 Molecular phylogenetics
• 4 Ernst Haeckel's recapitulation theory
• 5 Gene transfer
• 6 Taxon sampling and phylogenetic signal
• 7 See also
• 8 References
• 9 Further reading
• 10 External links
[edit] Construction of a phylogenetic tree
Evolution is regarded as a branching process, whereby populations are altered over time and may
speciate into separate branches, hybridize together, or terminate by extinction. This may be visualized in a
phylogenetic tree.
The problem posed by phylogenetics is that genetic data are only available for the present, and fossil
records (osteometric data) are sporadic and less reliable. Our knowledge of how evolution operates is used
to reconstruct the full tree.[3] Thus, a phylogenetic tree is based on a hypothesis of the order in which
evolutionary events are assumed to have occurred.
Cladistics is the current method of choice to infer phylogenetic trees. The most commonly-used
methods to infer phylogenies include parsimony, maximum likelihood, and MCMC-based Bayesian inference.
Phenetics, popular in the mid-20th century but now largely obsolete, uses distance matrix-based methods to
construct trees based on overall similarity, which is often assumed to approximate phylogenetic relationships.
All methods depend upon an implicit or explicit mathematical model describing the evolution of characters
observed in the species included, and are usually used for molecular phylogeny, wherein the characters are
aligned nucleotide or amino acid sequences.
[edit] Grouping of organisms

Phylogenetic groups, or taxa, can be monophyletic, paraphyletic, or polyphyletic.

There are some terms that describe the nature of a grouping in such trees. For instance, all birds and
reptiles are believed to have descended from a single common ancestor, so this taxonomic grouping (yellow
in the diagram below) is called monophyletic. "Modern reptile" (cyan in the diagram) is a grouping that
contains a common ancestor, but does not contain all descendants of that ancestor (birds are excluded). This
is an example of a paraphyletic group. A grouping such as warm-blooded animals would include only
mammals and birds (red/orange in the diagram) and is called polyphyletic because the members of this
grouping do not include the most recent common ancestor.

[edit] Molecular phylogenetics

Main article: Molecular phylogenetics
The evolutionary connections between organisms are represented graphically through phylogenetic
trees. Due to the fact that evolution takes place over long periods of time that cannot be observed directly,
biologists must reconstruct phylogenies by inferring the evolutionary relationships among present-day
organisms. Fossils can aid with the reconstruction of phylogenies; however, fossil records are often too poor
to be of good help. Therefore, biologists tend to be restricted with analysing present-day organisms to identify
their evolutionary relationships. Phylogenetic relationships in the past were reconstructed by looking at
phenotypes, often anatomical characteristics. Today, molecular data, which includes protein and DNA
sequences, are used to construct phylogenetic trees.[4] The overall goal of National Science Foundation's
Assembling the Tree of Life activity (AToL) is to resolve evolutionary relationships for large groups of
organisms throughout the history of life, with the research often involving large teams working across
institutions and disciplines. Investigators are typically supported for projects in data acquisition, analysis,
algorithm development and dissemination in computational phylogenetics and phyloinformatics. For
example, RedToL aims at reconstructing the Red Algal Tree of Life.
[edit] Ernst Haeckel's recapitulation theory
 Genealogical tree suggested by Haeckel (1866)
During the late 19th century, Ernst Haeckel's recapitulation theory, or biogenetic law, was widely
accepted. This theory was often expressed as "ontogeny recapitulates phylogeny", i.e. the development of an
organism exactly mirrors the evolutionary development of the species. Haeckel's early version of this
hypothesis [that the embryo mirrors adult evolutionary ancestors] has since been rejected, and the
hypothesis amended as the embryo's development mirroring embryos of its evolutionary ancestors. He was
accused by five professors of falsifying his images of embryos (See Ernst Haeckel). Most modern biologists
recognize numerous connections between ontogeny and phylogeny, explain them using evolutionary theory,
or view them as supporting evidence for that theory. Donald I. Williamson suggested that larvae and embryos
represented adults in other taxa that have been transferred by hybridization (the larval transfer theory).[5][6]
However, Williamson's views do not represent mainstream thought in molecular biology[7], and there is a
significant body of evidence against the larval transfer theory.[8]

[edit] Gene transfer

In general, organisms can inherit genes in two ways: vertical gene transfer and horizontal gene
transfer. Vertical gene transfer is the passage of genes from parent to offspring, and horizontal gene transfer
or lateral gene transfer occurs when genes jump between unrelated organisms, a common phenomenon in
prokaryotes.
Horizontal gene transfer has complicated the determination of phylogenies of organisms, and
inconsistencies in phylogeny have been reported among specific groups of organisms depending on the
genes used to construct evolutionary trees.
 Carl Woese came up with the three-domain theory of life (eubacteria, archaea and eukaryotes)
based on his discovery that the genes encoding ribosomal RNA are ancient and distributed over all lineages
of life with little or no horizontal gene transfer. Therefore, rRNAs are commonly recommended as molecular
clocks for reconstructing phylogenies.
This has been particularly useful for the phylogeny of microorganisms, to which the species concept
does not apply and which are too morphologically simple to be classified based on phenotypic traits.

[edit] Taxon sampling and phylogenetic signal

Owing to the development of advanced sequencing techniques in molecular biology, it has become
feasible to gather large amounts of data (DNA or amino acid sequences) to infer phylogenetic hypotheses.
For example, it is not rare to find studies with character matrices based on whole mitochondrial genomes
(~16,000 nucleotides, in many animals). However, it has been proposed that it is more important to increase
the number of taxa in the matrix than to increase the number of characters, because the more taxa the more
robust is the resulting phylogenetic tree [9] . This may be partly due to the breaking up of long branches. It
has been argued that this is an important reason to incorporate data from fossils into phylogenies where
possible. Of course, phylogenetic data that include fossil taxa are generally based on morphology, rather
than DNA data. Using simulations, Derrick Zwickl and David Hillis[10] found that increasing taxon sampling in
phylogenetic inference has a positive effect on the accuracy of phylogenetic analyses.
Another important factor that affects the accuracy of tree reconstruction is whether the data analyzed
actually contain a useful phylogenetic signal, a term that is used generally to denote whether related
organisms tend to resemble each other with respect to their genetic material or phenotypic traits.[11]
Ultimately, however, there is no way to measure whether a particular phylogenetic hypothesis is accurate or
not, unless the "true" relationships among the taxa being examined are already known. The best result an
empirical systematist can hope to attain is a tree with branches well-supported by the available evidence.

[edit] See also

Evolutionary biology portal

• Bauplan
• Bioinformatics
• Biomathematics
• Cladistics
• Coalescent theory
• Computational phylogenetics
• EDGE of Existence Programme
• Important publications in phylogenetics
• Language family
• Maximum parsimony
• Molecular phylogeny
• PhyloCode
• Joe Felsenstein
• Systematics
• Phylogenetic tree
 • Phylogenetic network
• Phylogenetic nomenclature
• Phylogenetics software
• Phylogenetic tree viewers
• Phylogeography
• Phylodynamics
• Phylogenetic comparative methods
• Microbial phylogenetics

[edit] References
1. ^ Edwards AWF, Cavalli-Sforza LL Phylogenetics is that branch of life science,which deals
with the study of evolutionary relation among various groups of organisms,through molecular
sequencing data. (1964). Systematics Assoc. Publ. No. 6: Phenetic and Phylogenetic Classification.
ed. Reconstruction of evolutionary trees. pp. 67–76.
2. ^ Speer, Vrian (1998). "UCMP Glossary: Phylogenetics". UC Berkeley.
http://www.ucmp.berkeley.edu/glossary/glossary_1.html. Retrieved 2008-03-22.
3. ^ Cavalli-Sforza LL, Edwards AWF (Sep., 1967). "Phylogenetic analysis: Models and
estimation procedures". Evol. 21 (3): 550–570. doi:10.2307/2406616. http://links.jstor.org/sici?
sici=0014-3820%28196709%2921%3A3%3C550%3APAMAEP%3E2.0.CO%3B2-I.
4. ^ Pierce, Benjamin A. (2007-12-17). Genetics: A conceptual Approach (3rd ed.). W. H.
Freeman. ISBN 978-0716-77928-5.
5. ^ Williamson DI (2003-12-31). "xviii". The Origins of Larvae (2nd ed.). Springer. pp. 261.
ISBN 978-1402-01514-4.
 6. ^ Williamson DI (2006). "Hybridization in the evolution of animal form and life-cycle".
Zoological Journal of the Linnean Society 148: 585–602. doi:10.1111/j.1096-3642.2006.00236.x.
7. ^ John Timmer, "Examining science on the fringes: vital, but generally wrong", ARS
Technica, 9 November 2009
8. ^ Michael W. Hart, and Richard K. Grosberg, "Caterpillars did not evolve from
onychophorans by hybridogenesis", Proceedings of the National Academy of the Sciences, 30
October 2009 (doi: 10.1073/pnas.0910229106)
9. ^ Wiens J (2006). "Missing data and the design of phylogenetic analyses". Journal of
Biomedical Informatics 39 (1): 34–42. doi:10.1016/j.jbi.2005.04.001. PMID 15922672.
10.^ Zwickl DJ, Hillis DM (2002). "Increased taxon sampling greatly reduces phylogenetic error".
Systematic Biology 51 (4): 588–598. doi:10.1080/10635150290102339. PMID 12228001.
11.^ Blomberg SP, Garland T Jr, Ives AR (2003). "Testing for phylogenetic signal in comparative
data: behavioral traits are more labile". Evolution 57 (4): 717–745. PMID 12778543. PDF

[edit] Further reading

• Schuh, R. T. and A. V. Z. Brower. 2009. Biological Systematics: principles and applications
(2nd edn.) ISBN 978-0-8014-4799-0

[edit] External links

• The Tree of Life
• Interactive Tree of Life
• PhyloCode
 • ExploreTree
• UCMP Exhibit Halls: Phylogeny Wing
• Willi Hennig Society
• Filogenetica.org in Spanish
• PhyloPat, Phylogenetic Patterns
• SplitsTree, program for computing phylogenetic trees and unrooted phylogenetic networks
• Dendroscope, program for drawing phylogenetic trees and rooted phylogenetic networks
• Phylogenetic inferring on the T-REX server
• Mesquite
• Geneious Pro all-in-one phylogenetics software
• NCBI - Systematics and Molecular Phylogenetics
• What Genomes Can Tell Us About the Past - lecture on phylogenetics by Sydney Brenner
• Mikko's Phylogeny Archive
• Phylogenetic Reconstruction from Gene-Order Data

[show]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of Adaptation · Macroevolution · Microevolution · Speciation

evolution

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

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Relevant fields Computational phylogenetics · Molecular phylogeny · Cladistics

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-morphy Symplesiomorphy · Apomorphy · Plesiomorphy · Synapomorphy ·
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 W000

Phylogénie
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

 Voir
« phylogénie » sur le
Wiktionnaire.

Arbre de Haeckel, 1866

 La phylogénie est l'étude des parentés entre différents êtres vivants en vue de comprendre
l'évolution des organismes vivants. On peut étudier la phylogénie, d'un groupe d'espèces mais également, à
un niveau intraspécifique, la généalogie entre populations ou entre individus.
On représente couramment une phylogénie par un arbre phylogénétique. Le nombre de nœuds entre
les branches, qui représente autant d'ancêtres communs, indique le degré de parenté entre les taxons. Plus
il y a de nœuds et donc d'ancêtres entre deux espèces, plus leur parenté est éloignée, c'est-à-dire que leur
ancêtre commun est ancien.
Dans un arbre élaboré par phénétique, la longueur des branches représente la distance génétique
entre taxons; dans un arbre élaboré par cladistique (cladogramme), on place sur les branches les
évènements évolutifs (caractères dérivés) ayant eu lieu dans chaque lignée.
Sommaire
[masquer]
• 1 Présentation
• 2 La cladistique
• 3 La phénétique
• 4 Utilisation conjointe de la phénétique et de la cladistique
• 5 Une phylogénie n'est pas une généalogie
• 6 Notes et références
• 7 Voir aussi
• 8 Liens externes

Présentation [modifier]
La systématique, l'étude de la diversité biologique en vue de sa classification phylogénétique, se
concentre, à la lumière des découvertes récentes, sur une classification phylogénétique remplaçant à
présent la classification classique. La classification classique établit des groupes ou taxons en fonction d'un
simple critère de ressemblance globale. Une classification phylogénétique suppose que l'on regroupe les
êtres vivants en fonction de leurs liens de parenté. Tout groupe systématique (ou « taxon ») renferme donc
des êtres vivants proches entre eux génétiquement (ce qui n'est pas toujours corrélé à une ressemblance
phénotypique globale). Les liens de parenté entre deux membres d'un taxon sont toujours plus étroits que les
liens de parenté entre un membre quelconque du groupe et un être vivant extérieur au groupe (il arrive que
ce membre extérieur soit pourtant très ressemblant en raison du phénomène de convergence évolutive, il
s'agit alors d'analogie entre les espèces, ce qui ne permet pas de les classer). Pour reconstituer les liens de
parenté entre êtres vivants, la phylogénie procède selon deux techniques : la phénétique et la cladistique. Il
est donc vraiment important de saisir la différence entre analogue (caractère qui se ressemble) et homologue
(caractère semblable hérité d'un ancêtre commun et dû à une évolution).

La cladistique [modifier]
La cladistique initiée par Willi Hennig hiérarchise les caractères comparés. Ne sont en fait regroupés
dans un même taxon que les êtres vivants qui partagent des caractères homologues : lorsqu'une
ressemblance entre deux taxons peut être attribuée à une ascendance commune, on parle d'homologie. Les
membres antérieurs de tous les tétrapodes, qu'ils soient bras ou ailes, sont homologues.
Ainsi l'aile de la chauve-souris et de l'oiseau sont-ils homologues en tant que membres antérieurs, et
non en tant qu'ailes. L'ancêtre commun de l'oiseau et de la chauve-souris possédait en effet déjà quatre
pattes mais ses membres antérieurs n'étaient pas des ailes. Cet ancêtre commun est en effet aussi celui des
lézards, des crocodiliens. Le membre antérieur « aile » est apparu plus tard indépendamment dans les deux
lignées chiroptères et oiseaux...
Les homologies sont en fait vues comme des innovations évolutives partagées (synapomorphies) : si
un même caractère homologue est partagé par deux taxons c'est que les deux taxons l'ont hérité de leur
ancêtre commun. Ce caractère homologue est donc apparu dans la lignée menant à cet ancêtre commun.
Tout être vivant possédant ce caractère homologue descend donc de cet ancêtre commun. Tout être vivant
ne possédant pas ce caractère homologue ne descend pas de cet ancêtre commun et est donc éloigné
génétiquement.
 La cladistique repose donc sur l'identification (souvent difficile) de l'homologie des caractères. Elle
est pertinente au niveau morphologique (et est donc le seul moyen de classer les espèces fossiles dont
l'ADN est rarement conservé) comme au niveau moléculaire. Les résultats sont représentés dans un arbre
phylogénétique ou cladogramme dans lequel chaque nœud représente un ancêtre commun et où les
synapomorphies sont représentées sur les branches dont la longueur est arbitraire. Deux taxons sont
d'autant plus apparentés qu'ils partagent un ancêtre commun proche dans l'arbre. Ici aussi, donc, les taxons
se retrouvent regroupés en fonction de leurs liens de parenté.

La phénétique [modifier]
La phénétique repose sur le postulat de base que le degré de ressemblance est corrélé au degré de
parenté. Elle suppose donc de quantifier la ressemblance entre les êtres vivants à classer.
Cette méthode se révèle peu pertinente lorsqu'on l'applique aux caractères morphologiques en
raison des analogies : certaines ressemblances entre êtres vivants ou taxons ne peuvent en effet être
attribuées à une ascendance commune. On parle alors d'analogie. Le principe utilisé pour expliquer ce
phénomène est la convergence évolutive : deux taxons différents vivant dans des niches écologiques
semblables ou sur lesquels la sélection naturelle a eu un impact semblable pourront avoir des caractères
analogues. Les ailes des oiseaux et des chauves-souris sont des caractères analogues en tant qu'ailes, car
ces deux ailes ne sont pas héritées d'un ancêtre commun ailé. De plus il est très difficile de quantifier
numériquement des ressemblances morphologiques.
En revanche, la phénétique devient pertinente dès lors que l'on compare un très grand nombre (au
sens statistique) de caractères car le nombre de caractères analogues devient négligeable parmi tous les
caractères dont la ressemblance est effectivement due à la parenté. Ainsi cette technique est très puissante
lorsqu'on l'applique au niveau moléculaire. Les systématiciens ont donc de plus en plus recours à des
méthodes moléculaires pour comparer les taxons et reconstruire les phylogénies. Pour ce faire, ils
comparent différents constituants moléculaires du vivant comme l'ADN, l'ARN ou les protéines. En effet,
ADN, ARN et protéines sont des molécules polymères. Chaque résidu de la molécule (nucléotide pour l'ADN
et l'ARN ou acide aminé pour la protéine) peut être considéré comme un caractère. Il est donc possible de
comparer les séquences chez plusieurs êtres vivants et de quantifier leur ressemblance par un simple
pourcentage que l'on assimile à la distance génétique entre les deux taxons auxquels appartiennent les deux
êtres vivants. Les résultats sont représentés dans un arbre phylogénétique, que l'on pourrait nommer
phénogramme, où la longueur des branches dépend de la distance génétique et représente donc le degré de
parenté entre les taxons étudiés. Cette technique se base sur le calcul d'un indice de similitude globale (ISG)
qui est défini après l'analyse de nombreux caractères (morphologiques, anatomiques, moléculaires...). Toute
analyse se fait à partir d'une seule espèce (exemple: comparaison de séquences nucléotidiques spécifiques
de plusieurs organismes par rapport à un seul) et à partir de cette comparaison, on crée une matrice de
distance (tableau au nombre d'entrées égal au nombre d'organismes comparés comprenant notre organisme
de référence) puis on recherche les plus petites distances (organismes les plus proches pour le critère
étudié) afin de constituer un arbre phylogénétique.

Utilisation conjointe de la phénétique et de la cladistique [modifier]

Pendant longtemps des discussions parfois violentes ont opposé tenants de l'une ou de l'autre
technique. Aujourd'hui la phénétique et la cladistique sont souvent utilisées conjointement comme étant deux
méthodes indépendantes. Lorsque leurs résultats sont convergents, on obtient des phylogénies très solides.
L'utilisation conjointe de ces deux méthodes a révélé l'existence dans la Classification classique de
nombreux groupes non fondés sur les liens de parenté et qui sont donc considérés comme non légitimes et
ne doivent plus êtres utilisés en taxonomie. L'utilisation de la phénétique moléculaire et de la cladistique
ainsi que la confrontation des arbres obtenus a été largement permise par les méthodes modernes que sont
l'amplification par PCR et le séquençage, alliées à de puissants outils de calcul qui permettent d'automatiser
ces méthodes.
Exemple de changements dans l'arbre phylogénétique dus à l'utilisation de ces techniques :
• le groupe des reptiles. Étaient regroupés au sein de celui-ci les crocodiliens (en fait
génétiquement proches des oiseaux) et les lézards, serpents et tortues (éloignés génétiquement des
oiseaux).
Exemple de l'utilisation du gène 16s pour les études de phylogénie des procaryotes.

Une phylogénie n'est pas une généalogie [modifier]

Le partage entre espèces d'un caractère ou d'un certain nombre de caractères jette sur ces mêmes
espèces le soupçon d'une origine commune qui remonte jusqu'à l'existence d'un ancêtre commun, le premier
à avoir acquis ce caractère ou ensemble de caractères. L'existence de l'ancêtre peut donc être découverte
grâce à la méthode cladistique, mais pas son identité, qui reste cachée. Ainsi par exemple les oiseaux
partagent tous un ancêtre commun, mais la découverte en 1861 d'un fossile comme Archaeopteryx, qui est
le plus ancien oiseau connu, ne prouve pas que ce fossile en particulier soit l'ancêtre de tous les oiseaux.
Effectivement une découverte future pourrait mettre au jour un oiseau fossile plus ancien qu'Archaeopteryx,
mais à nouveau la certitude d'être en face d'un « ancêtre » est inexistante. Les rapports d'ancêtre à
descendants (la généalogie) ne peuvent être identifiés en tant que tels que si l'identité même de l'ancêtre et
des descendants est préalablement connue. Autrement dit, pour retracer la généalogie, la science de la
classification devrait avoir la certitude de connaître toutes les espèces existantes et ayant existé. Comme ce
n'est pas le cas, car la science est loin de pouvoir connaître la totalité des espèces vivantes et fossiles, la
généalogie, même si elle a réellement eu lieu dans le passé, ne peut être retracée. Ce que la science de la
classification peut retracer, avec ces mêmes éléments partiels que sont les quelques espèces fossiles et
actuelles connues, ce sont les rapports de parenté entre espèces. Telle est la différence entre une
généalogie (« qui est ancêtre de qui ») et une phylogénie (« qui est le plus proche parent de qui »). Les
rapports phylogénétiques entre espèces connues constituent ainsi le seul critère objectif possible de
classification[1].

Notes et références [modifier]

1. ↑ (fr)La classification phylogénétique est un dispositif de classification des êtres vivants.
Elle tend à remplacer la classification traditionnelle en se basant seulement sur les rapports de
proximité évolutive entre espèces. [archive] sur www.mon-genome.com. Consulté le 28 juin 2010

Voir aussi [modifier]

• L'arbre phylogénétique du vivant
• Cladistique
• Classification phylogénétique
• Phylogéographie
• Phylogénie moléculaire
• Règnes du vivant
Liens externes [modifier]
• Fabriquer un arbre phylogénétique
• Phylogenie.net

• Portail de la biologie

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Phylog%C3%A9nie ».

Catégorie : Phylogénétique | [+]

 W000

Population genetics
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
Population genetics
Systematics

Biology portal · v • d • e
Population genetics is the study of allele frequency distribution and change under the influence of the
four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into
account the factors of population subdivision and population structure. It attempts to explain such
phenomena as adaptation and speciation.
 Population genetics was a vital ingredient in the emergence of the modern evolutionary synthesis. Its
primary founders were Sewall Wright, J. B. S. Haldane and R. A. Fisher, who also laid the foundations for the
related discipline of quantitative genetics.
Contents
[hide]
• 1 Fundamentals
• 1.1 Hardy–Weinberg principle
• 2 Scope and theoretical considerations
• 3 The four processes
• 3.1 Natural selection
• 3.2 Genetic drift
• 3.3 Mutation
• 3.4 Gene flow
• 4 Genetic structure
• 5 Microbial population genetics
• 6 History
• 6.1 Population genetics
• 6.2 Modern evolutionary synthesis
• 7 See also
• 8 References
• 9 External links
[edit] Fundamentals
Population genetics concerns the genetic constitution of populations and how this constitution
changes with time. A population is a set of organisms in which any pair of members can breed together. This
implies that all members belong to the same species and live near each other.[1]
For example, all of the moths of the same species living in an isolated forest are a population. A gene
in this population may have several alternate forms, which account for variations between the phenotypes of
the organisms. An example might be a gene for coloration in moths that has two alleles: black and white. A
gene pool is the complete set of alleles for a gene in a single population; the allele frequency for an allele is
the fraction of the genes in the pool that is composed of that allele (for example, what fraction of moth
coloration genes are the black allele). Evolution occurs when there are changes in the frequencies of alleles
within a population of interbreeding organisms; for example, the allele for black color in a population of moths
becoming more common.
To understand the mechanisms that cause a population to evolve, it is useful to consider what
conditions are required for a population not to evolve. The Hardy-Weinberg principle states that the
frequencies of alleles (variations in a gene) in a sufficiently large population will remain constant if the only
forces acting on that population are the random reshuffling of alleles during the formation of the sperm or
egg, and the random combination of the alleles in these sex cells during fertilization.[2] Such a population is
said to be in Hardy-Weinberg equilibrium as it is not evolving.[3]
 Hardy–Weinberg principle for two alleles: the horizontal axis shows the two allele frequencies p and q
and the vertical axis shows the genotype frequencies. Each graph shows one of the three possible
genotypes.

[edit] Hardy–Weinberg principle

Main article: Hardy–Weinberg principle
 The Hardy–Weinberg principle states that both allele and genotype frequencies in a population
remain constant—that is, they are in equilibrium—from generation to generation unless specific disturbing
influences are introduced. Outside the lab, one or more of these "disturbing influences" are always in effect.
Hardy Weinberg equilibrium is impossible in nature. Genetic equilibrium is an ideal state that provides a
baseline to measure genetic change against.
Allele frequencies in a population remain static across generations, provided the following conditions
are at hand: random mating, no mutation (the alleles don't change), no migration or emigration (no exchange
of alleles between populations), infinitely large population size, and no selective pressure for or against any
traits.
In the simplest case of a single locus with two alleles: the dominant allele is denoted A and the
recessive a and their frequencies are denoted by p and q; freq(A) = p; freq(a) = q; p + q = 1. If the population
is in equilibrium, then we will have freq(AA) = p2 for the AA homozygotes in the population, freq(aa) = q2 for
the aa homozygotes, and freq(Aa) = 2pq for the heterozygotes.
Based on these equations, useful but difficult-to-measure facts about a population can be
determined. For example, a patient's child is a carrier of a recessive mutation that causes cystic fibrosis in
homozygous recessive children. The parent wants to know the probability of her grandchildren inheriting the
disease. In order to answer this question, the genetic counselor must know the chance that the child will
reproduce with a carrier of the recessive mutation. This fact may not be known, but disease frequency is
known. We know that the disease is caused by the homozygous recessive genotype; we can use the Hardy–
Weinberg principle to work backward from disease occurrence to the frequency of heterozygous recessive
individuals.
Part of a series on
 Genetic genealogy
Concepts

• Genographic Project
• Genealogical DNA test
• Human mitochondrial DNA
haplogroups
• Human Y-chromosome DNA
haplogroups
• Haplogroup/ Haplotype
• Most recent common ancestor
• Personal genomics
• Population genetics

Other

• Genomics
This box: view • talk • edit

[edit] Scope and theoretical considerations

The mathematics of population genetics were originally developed as part of the modern evolutionary
synthesis. According to Beatty (1986), it defines the core of the modern synthesis.
 According to Lewontin (1974), the theoretical task for population genetics is a process in two spaces:
a "genotypic space" and a "phenotypic space". The challenge of a complete theory of population genetics is
to provide a set of laws that predictably map a population of genotypes (G1) to a phenotype space (P1),
where selection takes place, and another set of laws that map the resulting population ( P2) back to genotype
space (G2) where Mendelian genetics can predict the next generation of genotypes, thus completing the
cycle. Even leaving aside for the moment the non-Mendelian aspects of molecular genetics, this is clearly a
gargantuan task. Visualizing this transformation schematically:

(adapted from Lewontin 1974, p. 12). XD

T1 represents the genetic and epigenetic laws, the aspects of functional biology, or development, that
transform a genotype into phenotype. We will refer to this as the "genotype-phenotype map". T2 is the
transformation due to natural selection, T3 are epigenetic relations that predict genotypes based on the
selected phenotypes and finally T4 the rules of Mendelian genetics.

In practice, there are two bodies of evolutionary theory that exist in parallel, traditional population
genetics operating in the genotype space and the biometric theory used in plant and animal breeding,
operating in phenotype space. The missing part is the mapping between the genotype and phenotype space.
This leads to a "sleight of hand" (as Lewontin terms it) whereby variables in the equations of one domain, are
considered parameters or constants, where, in a full-treatment they would be transformed themselves by the
evolutionary process and are in reality functions of the state variables in the other domain. The "sleight of
hand" is assuming that we know this mapping. Proceeding as if we do understand it is enough to analyze
many cases of interest. For example, if the phenotype is almost one-to-one with genotype (sickle-cell
disease) or the time-scale is sufficiently short, the "constants" can be treated as such; however, there are
many situations where it is inaccurate.

[edit] The four processes

[edit] Natural selection
Main article: Natural selection
Natural selection is the process by which heritable traits that make it more likely for an organism to
survive and successfully reproduce become more common in a population over successive generations.
The natural genetic variation within a population of organisms means that some individuals will
survive more successfully than others in their current environment. Factors which affect reproductive success
are also important, an issue which Charles Darwin developed in his ideas on sexual selection.
Natural selection acts on the phenotype, or the observable characteristics of an organism, but the
genetic (heritable) basis of any phenotype which gives a reproductive advantage will become more common
in a population (see allele frequency). Over time, this process can result in adaptations that specialize
organisms for particular ecological niches and may eventually result in the emergence of new species.
Natural selection is one of the cornerstones of modern biology. The term was introduced by Darwin
in his groundbreaking 1859 book On the Origin of Species,[4] in which natural selection was described by
analogy to artificial selection, a process by which animals and plants with traits considered desirable by
human breeders are systematically favored for reproduction. The concept of natural selection was originally
developed in the absence of a valid theory of heredity; at the time of Darwin's writing, nothing was known of
modern genetics. The union of traditional Darwinian evolution with subsequent discoveries in classical and
molecular genetics is termed the modern evolutionary synthesis. Natural selection remains the primary
explanation for adaptive evolution.

[edit] Genetic drift

Main article: Genetic drift
Genetic drift is the change in the relative frequency in which a gene variant (allele) occurs in a
population due to random sampling and chance. That is, the alleles in the offspring in the population are a
random sample of those in the parents. And chance has a role in determining whether a given individual
survives and reproduces. A population's allele frequency is the fraction or percentage of its gene copies
compared to the total number of gene alleles that share a particular form.[5]
Genetic drift is an important evolutionary process which leads to changes in allele frequencies over
time. It may cause gene variants to disappear completely, and thereby reduce genetic variability. In contrast
to natural selection, which makes gene variants more common or less common depending on their
reproductive success,[6] the changes due to genetic drift are not driven by environmental or adaptive
pressures, and may be beneficial, neutral, or detrimental to reproductive success.
The effect of genetic drift is larger in small populations, and smaller in large populations. Vigorous
debates wage among scientists over the relative importance of genetic drift compared with natural selection.
Ronald Fisher held the view that genetic drift plays at the most a minor role in evolution, and this remained
the dominant view for several decades. In 1968 Motoo Kimura rekindled the debate with his neutral theory of
molecular evolution which claims that most of the changes in the genetic material are caused by genetic drift.
[7]

[edit] Mutation
Main article: Mutation
Mutations are changes in the DNA sequence of a cell's genome and are caused by radiation, viruses,
transposons and mutagenic chemicals, as well as errors that occur during meiosis or DNA replication.[8][9]
[10] Errors are introduced particularly often in the process of DNA replication, in the polymerization of the
second strand. These errors can also be induced by the organism itself, by cellular processes such as
hypermutation.
Mutations can have an impact on the phenotype of an organism, especially if they occur within the
protein coding sequence of a gene. Error rates are usually very low (1 error in every 10 million–100 million
bases) due to the "proofreading" ability of DNA polymerases.[11][12] Without proofreading, error rates are a
thousand-fold higher. Chemical damage to DNA occurs naturally as well, and cells use DNA repair
mechanisms to repair mismatches and breaks in DNA. Nevertheless, the repair sometimes fails to return the
DNA to its original sequence.
In organisms that use chromosomal crossover to exchange DNA and recombine genes, errors in
alignment during meiosis can also cause mutations.[13] Errors in crossover are especially likely when similar
sequences cause partner chromosomes to adopt a mistaken alignment; this makes some regions in
genomes more prone to mutating in this way. These errors create large structural changes in DNA sequence
—duplications, inversions or deletions of entire regions, or the accidental exchanging of whole parts between
different chromosomes (called translocation).
 Mutation can result in several different types of change in DNA sequences; these can either have no
effect, alter the product of a gene, or prevent the gene from functioning. Studies in the fly Drosophila
melanogaster suggest that if a mutation changes a protein produced by a gene, this will probably be harmful,
with about 70 percent of these mutations having damaging effects, and the remainder being either neutral or
weakly beneficial.[14] Due to the damaging effects that mutations can have on cells, organisms have evolved
mechanisms such as DNA repair to remove mutations.[8] Therefore, the optimal mutation rate for a species
is a trade-off between costs of a high mutation rate, such as deleterious mutations, and the metabolic costs
of maintaining systems to reduce the mutation rate, such as DNA repair enzymes.[15] Viruses that use RNA
as their genetic material have rapid mutation rates,[16] which can be an advantage since these viruses will
evolve constantly and rapidly, and thus evade the defensive responses of e.g. the human immune system.
[17]
Mutations can involve large sections of DNA becoming duplicated, usually through genetic
recombination.[18] These duplications are a major source of raw material for evolving new genes, with tens
to hundreds of genes duplicated in animal genomes every million years.[19] Most genes belong to larger
families of genes of shared ancestry.[20] Novel genes are produced by several methods, commonly through
the duplication and mutation of an ancestral gene, or by recombining parts of different genes to form new
combinations with new functions.[21][22]
Here, domains act as modules, each with a particular and independent function, that can be mixed
together to produce genes encoding new proteins with novel properties.[23] For example, the human eye
uses four genes to make structures that sense light: three for color vision and one for night vision; all four
arose from a single ancestral gene.[24] Another advantage of duplicating a gene (or even an entire genome)
is that this increases redundancy; this allows one gene in the pair to acquire a new function while the other
copy performs the original function.[25][26] Other types of mutation occasionally create new genes from
previously noncoding DNA.[27][28]

[edit] Gene flow

Main article: Gene flow
Gene flow is the exchange of genes between populations, which are usually of the same species.[29]
Examples of gene flow within a species include the migration and then breeding of organisms, or the
exchange of pollen. Gene transfer between species includes the formation of hybrid organisms and
horizontal gene transfer.
Migration into or out of a population can change allele frequencies, as well as introducing genetic
variation into a population. Immigration may add new genetic material to the established gene pool of a
population. Conversely, emigration may remove genetic material. As barriers to reproduction between two
diverging populations are required for the populations to become new species, gene flow may slow this
process by spreading genetic differences between the populations. Gene flow is hindered by mountain
ranges, oceans and deserts or even man-made structures such as the Great Wall of China, which has
hindered the flow of plant genes.[30]
Depending on how far two species have diverged since their most recent common ancestor, it may
still be possible for them to produce offspring, as with horses and donkeys mating to produce mules.[31]
Such hybrids are generally infertile, due to the two different sets of chromosomes being unable to pair up
during meiosis. In this case, closely related species may regularly interbreed, but hybrids will be selected
against and the species will remain distinct. However, viable hybrids are occasionally formed and these new
species can either have properties intermediate between their parent species, or possess a totally new
phenotype.[32] The importance of hybridization in creating new species of animals is unclear, although cases
have been seen in many types of animals,[33] with the gray tree frog being a particularly well-studied
example.[34]
Hybridization is, however, an important means of speciation in plants, since polyploidy (having more
than two copies of each chromosome) is tolerated in plants more readily than in animals.[35][36] Polyploidy is
important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to
pair with an identical partner during meiosis.[37] Polyploids also have more genetic diversity, which allows
them to avoid inbreeding depression in small populations.[38]
Horizontal gene transfer is the transfer of genetic material from one organism to another organism
that is not its offspring; this is most common among bacteria.[39] In medicine, this contributes to the spread
of antibiotic resistance, as when one bacteria acquires resistance genes it can rapidly transfer them to other
species.[40] Horizontal transfer of genes from bacteria to eukaryotes such as the yeast Saccharomyces
cerevisiae and the adzuki bean beetle Callosobruchus chinensis may also have occurred.[41][42] An
example of larger-scale transfers are the eukaryotic bdelloid rotifers, which appear to have received a range
of genes from bacteria, fungi, and plants.[43] Viruses can also carry DNA between organisms, allowing
transfer of genes even across biological domains.[44] Large-scale gene transfer has also occurred between
the ancestors of eukaryotic cells and prokaryotes, during the acquisition of chloroplasts and mitochondria.
[45]
Gene flow is the transfer of alleles from one population to another.
Migration into or out of a population may be responsible for a marked change in allele frequencies.
Immigration may also result in the addition of new genetic variants to the established gene pool of a
particular species or population.
 There are a number of factors that affect the rate of gene flow between different populations. One of
the most significant factors is mobility, as greater mobility of an individual tends to give it greater migratory
potential. Animals tend to be more mobile than plants, although pollen and seeds may be carried great
distances by animals or wind.
Maintained gene flow between two populations can also lead to a combination of the two gene pools,
reducing the genetic variation between the two groups. It is for this reason that gene flow strongly acts
against speciation, by recombining the gene pools of the groups, and thus, repairing the developing
differences in genetic variation that would have led to full speciation and creation of daughter species.
For example, if a species of grass grows on both sides of a highway, pollen is likely to be transported
from one side to the other and vice versa. If this pollen is able to fertilise the plant where it ends up and
produce viable offspring, then the alleles in the pollen have effectively been able to move from the population
on one side of the highway to the other.

[edit] Genetic structure

Because of physical barriers to migration, along with limited vagility, and natal philopatry, natural
populations are rarely panmictic (Buston et al., 2007). There is usually a geographic range within which
individuals are more closely related to one another than those randomly selected from the general
population. This is described as the extent to which a population is genetically structured (Repaci et al.,
2007).
[edit] Microbial population genetics
Microbial population genetics is a rapidly advancing field of investigation with relevance to many
other theoretical and applied areas of scientific investigations. The population genetics of microorganisms
lays the foundations for tracking the origin and evolution of antibiotic resistance and deadly infectious
pathogens. Population genetics of microorganisms is also an essential factor for devising strategies for the
conservation and better utilization of beneficial microbes (Xu, 2010).

[edit] History
See also: Modern evolutionary synthesis

Biston betularia f. typica is the white-bodied form of the peppered moth.

 Biston betularia f. carbonaria is the black-bodied form of the peppered moth.

[edit] Population genetics

Population genetics was developed as a reconciliation of the Mendelian and biometrician models. A
key step was the work of the British biologist and statistician R.A. Fisher. In a series of papers starting in
1918 and culminating in his 1930 book The Genetical Theory of Natural Selection , Fisher showed that the
continuous variation measured by the biometricians could be produced by the combined action of many
discrete genes, and that natural selection could change gene frequencies in a population, resulting in
evolution (though lacking the knowledge of what an actual gene was at this time, it should be said in this
sense he understood phenotypic trait frequency, rather than specifically identifiable gene frequency). In a
series of papers beginning in 1924, another British geneticist, J.B.S. Haldane, applied statistical analysis to
real-world examples of natural selection, such as the evolution of industrial melanism in peppered moths,
and showed that natural selection worked at an even faster rate than Fisher assumed.[46][47]
 The American biologist Sewall Wright, who had a background in animal breeding experiments,
focused on combinations of interacting genes, and the effects of inbreeding on small, relatively isolated
populations that exhibited genetic drift. In 1932, Wright introduced the concept of an adaptive landscape and
argued that genetic drift and inbreeding could drive a small, isolated sub-population away from an adaptive
peak, allowing natural selection to drive it towards different adaptive peaks. Fisher and Wright had some
fundamental disagreements and a controversy about the relative roles of selection and drift continued for
much of the century between the Americans and the British. The Frenchman Gustave Malécot was also
important early in the development of the discipline.
The work of Fisher, Haldane and Wright founded the discipline of population genetics. This
integrated natural selection with Mendelian genetics, which was the critical first step in developing a unified
theory of how evolution worked.[46][47]
John Maynard Smith was Haldane's pupil, whilst W.D. Hamilton was heavily influenced by the
writings of Fisher. The American George R. Price worked with both Hamilton and Maynard Smith. American
Richard Lewontin and Japanese Motoo Kimura were heavily influenced by Wright.

[edit] Modern evolutionary synthesis

In the first few decades of the 20th century, most field naturalists continued to believe that
Lamarckian and orthogenic mechanisms of evolution provided the best explanation for the complexity they
observed in the living world. However, as the field of genetics continued to develop, those views became less
tenable.[48] Theodosius Dobzhansky, a postdoctoral worker in T. H. Morgan's lab, had been influenced by
the work on genetic diversity by Russian geneticists such as Sergei Chetverikov. He helped to bridge the
divide between the foundations of microevolution developed by the population geneticists and the patterns of
macroevolution observed by field biologists, with his 1937 book Genetics and the Origin of Species.
 Dobzhansky examined the genetic diversity of wild populations and showed that, contrary to the
assumptions of the population geneticists, these populations had large amounts of genetic diversity, with
marked differences between sub-populations. The book also took the highly mathematical work of the
population geneticists and put it into a more accessible form. In Great Britain E.B. Ford, the pioneer of
ecological genetics, continued throughout the 1930s and 1940s to demonstrate the power of selection due to
ecological factors including the ability to maintain genetic diversity through genetic polymorphisms such as
human blood types. Ford's work would contribute to a shift in emphasis during the course of the modern
synthesis towards natural selection over genetic drift.[46][47][49][50]

[edit] See also

Evolutionary biology portal

• Coalescent theory
• Dual inheritance theory
• Ecological genetics
• Evolutionarily Significant Unit
• Ewens's sampling formula
• Fitness landscape
• Founder effect
• Genetic diversity
• Genetic drift
• Genetic erosion
• Genetic hitchhiking
• Genetic pollution
• Gene pool
• Genotype-phenotype distinction
• Habitat fragmentation
• Haldane's dilemma
• Hardy-Weinberg principle
• Hill-Robertson effect
• Linkage disequilibrium
• Microevolution
• Molecular evolution
• Muller's ratchet
• Mutational meltdown
• Neutral theory of molecular evolution
• Population bottleneck
• Quantitative genetics
• Reproductive compensation
• Selection
• Selective sweep
• Small population size
• Viral quasispecies
[edit] References
1. ^ Hartl, Daniel (2007). Principles of Population Genetics. Sinauer Associates. p. 95.
ISBN 978-0-87893-308-2.
2. ^ O'Neil, Dennis (2008). "Hardy-Weinberg Equilibrium Model". The synthetic theory of
evolution: An introduction to modern evolutionary concepts and theories . Behavioral Sciences
Department, Palomar College. http://anthro.palomar.edu/synthetic/synth_2.htm. Retrieved 2008-01-
06.
3. ^ Bright, Kerry (2006). "Causes of evolution". Teach Evolution and Make It Relevant. National
Science Foundation. http://evoled.dbs.umt.edu/lessons/causes.htm#hardy. Retrieved 2007-12-30.
4. ^ Darwin C (1859) On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life John Murray, London; modern reprint
Charles Darwin, Julian Huxley (2003). The Origin of Species. Signet Classics. ISBN 0-451-52906-5.
Published online at The complete work of Charles Darwin online: On the origin of species by means
of natural selection, or the preservation of favoured races in the struggle for life.
5. ^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. Glossary. ISBN 0-
87893-189-9.
6. ^ Avers, Charlotte (1989). Process and Pattern in Evolution. Oxford University Press
7. ^ Futuyma, Douglas (1998). Evolutionary Biology. Sinauer Associates. p. 320. ISBN 0-
87893-189-9.
8. ^ a b Bertram J (2000). "The molecular biology of cancer". Mol. Aspects Med. 21 (6): 167–223.
doi:10.1016/S0098-2997(00)00007-8. PMID 11173079.
 9. ^ Aminetzach YT, Macpherson JM, Petrov DA (2005). "Pesticide resistance via transposition-
mediated adaptive gene truncation in Drosophila". Science 309 (5735): 764–7.
doi:10.1126/science.1112699. PMID 16051794.
10.^ Burrus V, Waldor M (2004). "Shaping bacterial genomes with integrative and conjugative
elements". Res. Microbiol. 155 (5): 376–86. doi:10.1016/j.resmic.2004.01.012. PMID 15207870.
11.^ Griffiths, William M.; Miller, Jeffrey H.; Suzuki, David T. et al., eds (2000). "Spontaneous
mutations". An Introduction to Genetic Analysis (7th ed.). New York: W. H. Freeman. ISBN 0-7167-
3520-2. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=iga.section.2706.
12.^ Freisinger, E; Grollman, AP; Miller, H; Kisker, C (2004). "Lesion (in)tolerance reveals
insights into DNA replication fidelity.". The EMBO journal 23 (7): 1494–505.
doi:10.1038/sj.emboj.7600158. PMID 15057282.
13.^ Griffiths, William M.; Miller, Jeffrey H.; Suzuki, David T. et al., eds (2000). "Chromosome
Mutation I: Changes in Chromosome Structure: Introduction". An Introduction to Genetic Analysis
(7th ed.). New York: W. H. Freeman. ISBN 0-7167-3520-2.
http://www.ncbi.nlm.nih.gov/books/bv.fcgi?rid=iga.section.2844.
14.^ Sawyer SA, Parsch J, Zhang Z, Hartl DL (2007). "Prevalence of positive selection among
nearly neutral amino acid replacements in Drosophila". Proc. Natl. Acad. Sci. U.S.A. 104 (16): 6504–
10. doi:10.1073/pnas.0701572104. PMID 17409186.
15.^ Sniegowski P, Gerrish P, Johnson T, Shaver A (2000). "The evolution of mutation rates:
separating causes from consequences". Bioessays 22 (12): 1057–66. doi:10.1002/1521-
1878(200012)22:12<1057::AID-BIES3>3.0.CO;2-W. PMID 11084621.
16.^ Drake JW, Holland JJ (1999). "Mutation rates among RNA viruses". Proc. Natl. Acad. Sci.
U.S.A. 96 (24): 13910–3. doi:10.1073/pnas.96.24.13910. PMID 10570172. PMC 24164.
http://www.pnas.org/content/96/24/13910.long.
 17.^ Holland J, Spindler K, Horodyski F, Grabau E, Nichol S, VandePol S (1982). "Rapid
evolution of RNA genomes". Science 215 (4540): 1577–85. doi:10.1126/science.7041255.
PMID 7041255.
18.^ Hastings, P J; Lupski, JR; Rosenberg, SM; Ira, G (2009). "Mechanisms of change in gene
copy number". Nature Reviews. Genetics 10 (8): 551–564. doi:10.1038/nrg2593. PMID 19597530.
19.^ Carroll SB, Grenier J, Weatherbee SD (2005). From DNA to Diversity: Molecular Genetics
and the Evolution of Animal Design. Second Edition . Oxford: Blackwell Publishing. ISBN 1-4051-
1950-0.
20.^ Harrison P, Gerstein M (2002). "Studying genomes through the aeons: protein families,
pseudogenes and proteome evolution". J Mol Biol 318 (5): 1155–74. doi:10.1016/S0022-
2836(02)00109-2. PMID 12083509.
21.^ Orengo CA, Thornton JM (2005). "Protein families and their evolution-a structural
perspective". Annu. Rev. Biochem. 74: 867–900. doi:10.1146/annurev.biochem.74.082803.133029.
PMID 15954844.
22.^ Long M, Betrán E, Thornton K, Wang W (November 2003). "The origin of new genes:
glimpses from the young and old". Nat. Rev. Genet. 4 (11): 865–75. doi:10.1038/nrg1204.
PMID 14634634.
23.^ Wang M, Caetano-Anollés G (2009). "The evolutionary mechanics of domain organization
in proteomes and the rise of modularity in the protein world". Structure 17 (1): 66–78.
doi:10.1016/j.str.2008.11.008. PMID 19141283.
24.^ Bowmaker JK (1998). "Evolution of colour vision in vertebrates". Eye (London, England) 12
(Pt 3b): 541–7. PMID 9775215.
 25.^ Gregory TR, Hebert PD (1999). "The modulation of DNA content: proximate causes and
ultimate consequences". Genome Res. 9 (4): 317–24. doi:10.1101/gr.9.4.317 (inactive 2009-11-14).
PMID 10207154. http://genome.cshlp.org/content/9/4/317.full.
26.^ Hurles M (July 2004). "Gene duplication: the genomic trade in spare parts". PLoS Biol. 2
(7): E206. doi:10.1371/journal.pbio.0020206. PMID 15252449.
27.^ Liu N, Okamura K, Tyler DM (2008). "The evolution and functional diversification of animal
microRNA genes". Cell Res. 18 (10): 985–96. doi:10.1038/cr.2008.278. PMID 18711447.
PMC 2712117. http://www.nature.com/cr/journal/v18/n10/full/cr2008278a.html.
28.^ Siepel A (October 2009). "Darwinian alchemy: Human genes from noncoding DNA".
Genome Res. 19 (10): 1693–5. doi:10.1101/gr.098376.109. PMID 19797681. PMC 2765273.
http://genome.cshlp.org/content/19/10/1693.full.
29.^ Morjan C, Rieseberg L (2004). "How species evolve collectively: implications of gene flow
and selection for the spread of advantageous alleles". Mol. Ecol. 13 (6): 1341–56. doi:10.1111/j.1365-
294X.2004.02164.x. PMID 15140081.
30.^ Su H, Qu L, He K, Zhang Z, Wang J, Chen Z, Gu H (2003). "The Great Wall of China: a
physical barrier to gene flow?". Heredity 90 (3): 212–9. doi:10.1038/sj.hdy.6800237.
PMID 12634804.
31.^ Short RV (1975). "The contribution of the mule to scientific thought". J. Reprod. Fertil.
Suppl. (23): 359–64. PMID 1107543.
32.^ Gross B, Rieseberg L (2005). "The ecological genetics of homoploid hybrid speciation". J.
Hered. 96 (3): 241–52. doi:10.1093/jhered/esi026. PMID 15618301.
33.^ Burke JM, Arnold ML (2001). "Genetics and the fitness of hybrids". Annu. Rev. Genet. 35:
31–52. doi:10.1146/annurev.genet.35.102401.085719. PMID 11700276.
 34.^ Vrijenhoek RC (2006). "Polyploid hybrids: multiple origins of a treefrog species". Curr. Biol.
16 (7): R245. doi:10.1016/j.cub.2006.03.005. PMID 16581499.
35.^ Wendel J (2000). "Genome evolution in polyploids". Plant Mol. Biol. 42 (1): 225–49.
doi:10.1023/A:1006392424384. PMID 10688139.
36.^ Sémon M, Wolfe KH (2007). "Consequences of genome duplication". Curr Opin Genet Dev
17 (6): 505–12. doi:10.1016/j.gde.2007.09.007. PMID 18006297.
37.^ Comai L (2005). "The advantages and disadvantages of being polyploid". Nat. Rev. Genet.
6 (11): 836–46. doi:10.1038/nrg1711. PMID 16304599.
38.^ Soltis P, Soltis D (June 2000). "The role of genetic and genomic attributes in the success of
polyploids". Proc. Natl. Acad. Sci. U.S.A. 97 (13): 7051–7. doi:10.1073/pnas.97.13.7051.
PMID 10860970.
39.^ Boucher Y, Douady CJ, Papke RT, Walsh DA, Boudreau ME, Nesbo CL, Case RJ, Doolittle
WF (2003). "Lateral gene transfer and the origins of prokaryotic groups". Annu Rev Genet 37: 283–
328. doi:10.1146/annurev.genet.37.050503.084247. PMID 14616063.
40.^ Walsh T (2006). "Combinatorial genetic evolution of multiresistance". Curr. Opin. Microbiol.
9 (5): 476–82. doi:10.1016/j.mib.2006.08.009. PMID 16942901.
41.^ Kondo N, Nikoh N, Ijichi N, Shimada M, Fukatsu T (2002). "Genome fragment of Wolbachia
endosymbiont transferred to X chromosome of host insect". Proc. Natl. Acad. Sci. U.S.A. 99 (22):
14280–5. doi:10.1073/pnas.222228199. PMID 12386340.
42.^ Sprague G (1991). "Genetic exchange between kingdoms". Curr. Opin. Genet. Dev. 1 (4):
530–3. doi:10.1016/S0959-437X(05)80203-5. PMID 1822285.
43.^ Gladyshev EA, Meselson M, Arkhipova IR (May 2008). "Massive horizontal gene transfer in
bdelloid rotifers". Science 320 (5880): 1210–3. doi:10.1126/science.1156407. PMID 18511688.
 44.^ Baldo A, McClure M (1 September 1999). "Evolution and horizontal transfer of dUTPase-
encoding genes in viruses and their hosts". J. Virol. 73 (9): 7710–21. PMID 10438861.
45.^ Poole A, Penny D (2007). "Evaluating hypotheses for the origin of eukaryotes". Bioessays
29 (1): 74–84. doi:10.1002/bies.20516. PMID 17187354.
46.^ a b c Bowler 2003, pp. 325–339
47.^ a b c Larson 2004, pp. 221–243
48.^ Mayr & Provine 1998, pp. 295–298, 416
49.^ Mayr, E§year=1988. Towards a new philosophy of biology: observations of an evolutionist .
Harvard University Press. pp. 402.
50.^ Mayr & Provine 1998, pp. 338–341
• J. Beatty. "The synthesis and the synthetic theory" in Integrating Scientific Disciplines, edited
by W. Bechtel and Nijhoff. Dordrecht, 1986.
• Buston, PM; et al. (2007). "Are clownfish groups composed of close relatives? An analysis of
microsatellite DNA vraiation in Amphiprion percula". Molecular Ecology 12 (3): 733–742.
PMID 12675828.
• Luigi Luca Cavalli-Sforza. Genes, Peoples, and Languages. North Point Press, 2000.
• Luigi Luca Cavalli-Sforza et al. The History and Geography of Human Genes. Princeton
University Press, 1994.
• James F. Crow and Motoo Kimura. Introduction to Population Genetics Theory. Harper &
Row, 1972.
• Warren J Ewens. Mathematical Population Genetics. Springer-Verlag New York, Inc., 2004.
ISBN 0-387-20191-2
 • John H. Gillespie Population Genetics: A Concise Guide, Johns Hopkins Press, 1998. ISBN
0-8018-5755-4.
• Richard Halliburton. Introduction to Population Genetics. Prentice Hall, 2004
• Daniel Hartl. Primer of Population Genetics, 3rd edition. Sinauer, 2000. ISBN 0-87893-304-2
• Daniel Hartl and Andrew Clark. Principles of Population Genetics, 3rd edition. Sinauer, 1997.
ISBN 0-87893-306-9.
• Richard C. Lewontin. The Genetic Basis of Evolutionary Change. Columbia University Press,
1974.
• William B. Provine. The Origins of Theoretical Population Genetics. University of Chicago
Press. 1971. ISBN 0-226-68464-4.
• Repaci, V; Stow AJ, Briscoe DA (2007). "Fine-scale genetic structure, co-founding and
multiple mating in the Australian allodapine bee ( Ramphocinclus brachyurus". Journal of Zoology
270: 687–691. doi:10.1111/j.1469-7998.2006.00191.x.
• Spencer Wells. The Journey of Man. Random House, 2002.
• Spencer Wells. Deep Ancestry: Inside the Genographic Project. National Geographic
Society, 2006.
• Cheung, KH; Osier MV, Kidd JR, Pakstis AJ, Miller PL, Kidd KK (2000). "ALFRED: an allele
frequency database for diverse populations and DNA polymorphisms". Nucleic Acids Research 28
(1): 361–3. doi:10.1093/nar/28.1.361. PMID 10592274.
• Xu, J. Microbial Population Genetics. Caister Academic Press, 2010. ISBN 978-1-904455-
59-2
 [edit] External links
• Yale University
• EHSTRAFD.org - Earth Human STR Allele Frequencies Database
• History of population genetics
• National Geographic: Atlas of the Human Journey (Haplogroup-based human migration
maps)
• Monash Virtual Laboratory - Simulations of habitat fragmentation and population genetics
online at Monash University's Virtual Laboratory.

[show]
v•d•e
Topics in population genetics

Key concepts Hardy-Weinberg law · Genetic linkage · Linkage disequilibrium · Fisher's

fundamental theorem · Neutral theory · Price equation

Selection Natural · Sexual · Artificial · Ecological

Effects of
selection Genetic hitchhiking · Background selection
on genomic variation
 Genetic drift Small population size · Population bottleneck · Founder effect ·
Coalescence · Balding–Nichols model

Founders R. A. Fisher · J. B. S. Haldane · Sewall Wright

Related topics Evolution · Microevolution · Evolutionary game theory · Fitness landscape ·

Genetic genealogy

List of evolutionary biology topics

[show]
v•d•e
Genetics

Introduction · History · Related topics · List of organizations

Key
components Chromosome · DNA · Nucleotide · RNA · Genome

Fields of Classical genetics · Conservation genetics · Ecological genetics ·

genetics Immunogenetics · Molecular genetics · Population genetics · Quantitative genetics

Related topics Geneticist · Genomics · Genetic code · Medical genetics · Molecular

evolution · Reverse genetics · Genetic engineering · Genetic diversity · Heredity

[show]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental Canalisation · Inversion · Modularity · Phenotypic plasticity
biology (Evo-devo)
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Population_genetics"

Categories: Genetics | Population genetics | Evolutionary biology | Statistical genetics

• This page was last modified on 23 August 2010 at 03:32.

 W000

Génétique des populations

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

La génétique des populations traite des fluctuations des fréquences des différentes versions d'un
gène (allèles) au cours du temps dans les populations d'êtres vivants, sous l'influence de la sélection
naturelle, de la dérive génétique, des mutations et des migrations. Ces différents facteurs sont appelés des
« pressions évolutives ». Les fluctuations de fréquence des allèles sont la première étape de l'évolution. En
effet, la fixation de certains allèles peut conduire à des adaptations. Par la suite, l'accumulation d'adaptations
différentes dans différentes populations peut conduire au processus de spéciation.
La génétique des populations est une application, commencée dans les années 1920 à 1940 par
Ronald Fisher, J. B. S. Haldane et Sewall Wright, des principes fondamentaux de la génétique mendélienne
à l'échelle des populations. Cette application a permis de faire la synthèse entre la génétique mendélienne et
la théorie de l'évolution, donnant ainsi naissance au néo-darwinisme (théorie synthétique de l'évolution).
La génétique des populations a des applications en épidémiologie où elle permet de comprendre la
transmission des maladies génétiques, mais aussi en agronomie, où des programmes de sélection modifient
le patrimoine génétique de certains organismes pour créer des races ou variétés plus performantes, ou plus
résistantes à des maladies. Elle permet également de comprendre les mécanismes de conservation et de
disparition des populations et des espèces (Génétique de la conservation).
C'est une discipline des sciences de la vie faisant un fort usage d'outils mathématiques.
Sommaire
[masquer]
• 1 Introduction simplifiée à la génétique des populations
humaines
• 1.1 Les ancêtres de l'humanité
• 1.2 Les marqueurs génétiques
• 1.3 Les haplogroupes et leur classification
• 1.4 Les haplotypes
• 1.5 Les autres systèmes de classification
• 1.6 Signature génétique des chromosomes
homologues
• 2 Définition de la population
• 3 Mutation, dérive, sélection et migration
• 3.1 Mutations
• 3.2 Dérive et sélection
• 3.3 Migrations
• 3.4 Régimes de reproduction
• 4 Méthodes d'étude de la variabilité
• 4.1 Notion de polymorphisme
• 4.2 Mesure de la diversité génétique
• 4.3 Notion de population théorique idéale et loi de
Introduction simplifiée à la génétique des populations humaines [modifier]
Les êtres vivants et les êtres humains en particulier possèdent tous de l'ADN. L'étude de cet ADN
pour une population et la comparaison de cet ADN avec l'ADN d'autres populations constituent la base de la
génétique des populations.
Nous possédons 22 paires de chromosomes dit homologues (ou autosomes) et deux chromosomes
dit sexuels (ou gonosomes). Par ailleurs, nous possédons aussi de l'ADN dit mitochondrial (ADN-mt ou mt-
DNA en anglais) qui n'est pas à proprement parler un chromosome. Cet ADN-mt se transmet intégralement
de la mère aux enfants. En revanche, seuls les hommes possèdent le chromosome sexuel appelé Y (ADN-Y
ou Y-DNA en anglais) qui se transmet donc intégralement du père aux fils.
Notre ADN peut parfois muter, c'est-à-dire qu'un des éléments de base (58 millions de paires de
bases pour l'ADN-Y et 16000 paires de bases pour l'ADN-mt) qui le constitue se transforme lors de la recopie
de cet ADN. Le résultat de cette mutation s'appelle polymorphisme nucléotidique simple (SNP en anglais).
Cette mutation arrive très approximativement une fois toutes les 25 à 500 générations pour l'ADN-Y pour
l'ADN-mt (il n'y a pas de consensus à ce sujet).
Comme nous allons le voir ci-dessous, les mutations de l'ADN-Y et de l'ADN-mt sont utilisées pour
caractériser des groupes de populations. Par ailleurs, ces deux ADN sont peu sujets à la sélection naturelle.
Ils sont par conséquent particulièrement bien adaptés pour suivre l'évolution des populations.

Les ancêtres de l'humanité [modifier]

Au début des années 2000, Spencer Wells(en) a conclu que tous les êtres humains vivants avaient
au moins un ancêtre mâle[1] et un ancêtre femelle en commun. Le plus récent de ces ancêtres mâle est
appelé Adam Y-chromosomique (Y-MRCA pour Most Recent Common Ancestor) et le plus récent ancêtre
femelle est appelé Eve mitochondrial (mt-MRCA). Il faut souligner que contrairement à l'Eve et l'Adam de la
Bible, l'Adam Y-chromosomique et l'Eve mitochondriale ne sont pas les premiers ancêtres de l'humanité
actuelle. De plus, Y-MRCA n'est pas forcément le seul ancêtre de l'humanité puisqu'il peut exister un ancêtre
mâle n'ayant eut que des filles. De même, il peut exister un ancêtre femelle commun à l'humanité actuelle
n'ayant donné que des fils.
Enfin, Y-MRCA et mt-MRCA ne se sont jamais rencontrés.
Dans la suite du paragraphe il n'est fait référence qu'aux lignées paternelles mais les explications
sont les mêmes pour les lignées maternelles.
Le chromosome Y de cet Adam s'est transmis à ses descendants mâles. Certains des chromosomes
Y de ses descendants ont subi une mutation. Cette mutation définit une nouvelle branche à laquelle on peut
associer un nouvel ancêtre commun. Si le chromosome Y d'un des descendants de cette branche subit une
nouvelle mutation, cela crée une nouvelle sous-branche et ainsi de suite. On peut ainsi définir l'arbre de la
filiation paternelle de l'humanité.

Les marqueurs génétiques [modifier]

Pour caractériser un chromosome on utilise des marqueurs génétiques. Il existe différents types de
marqueurs, les plus utilisés sont
• les marqueurs SNP (qui définissent la mutation d'une seule base), ils sont utilisés entre
autres pour définir les arbres des filiations de l'humanité. Pour le chromosome Y, ils prennent le nom
de XN où X est un indice définissant le laboratoire ou l'entreprise ayant découvert le marqueur et N
 le nième marqueur découvert dans ce laboratoire.[2] Par exemple M35 est le 35e marqueur SNP
découvert par l'Université de Stanford.
• et les marqueurs STR (Short Tandem Notice ou encore microsatellites). Un chromosome
contient des séquences répétées de nucléotides (de paires de bases). Le nombre de répétition varie
d'une personne à l'autre. Un STR du chromosome Y est désigné par un nombre DYS (DNA Y-
chromosome Segment number). Lorsqu'on « test » une personne, on associe au marqueur DYS le
nombre de répétitions de la séquence STR du chromosome la personne concernée. Ils sont utilisés
pour définir les haplotypes (voir ci-dessous), la résolution de l'haplotype croît avec le nombre de
marqueurs STR.
Pour mettre en évidence ces marqueurs génétiques on extrait l'ADN et on lui fait subir différents
processus physico-chimiques.

Les haplogroupes et leur classification [modifier]

En génétique des populations, chaque branche majeure s'appelle haplogroupe et chaque sous-
branche majeure sous-haplogroupe. Le terme haplogroupe ou sous-haplogroupe n'est pas absolu, il est
relatif à l'endroit étudié de l'arbre. La définition de cet arbre est loin d'être achevée si bien que la
dénomination des haplogroupes change régulièrement. Une branche prend parfois le terme biologique de
clade.
La plupart des études utilisent maintenant cet arbre généalogique de l'ADN-Y avec sa nomenclature
associée. Cette nomenclature a été définie une première fois en 2002 par le Y Chromosome Consortium
(YCC). Cet arbre comprend 15 haplogoupes majeurs (A, B, C, D, E, G, H, I, J, L, M, N, O, Q et R). Chaque
sous-haplogroupe associé à son haplogroupe est nommé par le nom de son haplogroupe plus un numéro de
sous-branche (exemple R1). Puis les sous-haplogroupes des sous-haplogroupes sont nommés avec la
dénomination de son haplogroupe parent plus une lettre minuscule (exemple R1b) et ainsi de suite en
alternant lettres et chiffres.
Une cartographie des haplogroupes des ADN-Y de l'ensemble des populations est en train d'être
réalisée. Elle permet de mieux comprendre les migrations et les affinités des patrimoines génétiques
paternels des populations humaines. Nous donnons ci-dessous la version française de l'arbre sur wikipedia
mais nous recommandons au lecteur intéressé de consulter la version anglaise qui est mise à jour
régulièrement.
Haplogroupes du chromosome Y (Y-ADN)

|
|
C | F
D E G H I J K
L M
N
 Puisque les marqueurs SNP définissent la mutation d'une base, ils sont particulièrement bien
adaptés pour définir les haplogroupes. Afin d'illustrer ceci revenons à l'exemple du marqueur SNP M35
correspond à l'haplogroupe E1b1b1b (pour le savoir il faut consulter l'arbre des filiations paternels de
l'humanité). Cet haplogroupe est particulièrement fréquent dans les populations berbères. Il possède des
sous-haplogroupes définis par d'autres marqueurs SNP.
Cette nomenclature évoluant encore, on associe presque systématiquement le marqueur SNP
caractérisant l'haplogroupe à l'haplogroupe correspondant.
Les lignées paternelles d'une population sont caractérisées par la distribution d'haplogroupes de
l'ADN-Y c'est-à-dire par l'ensemble et la proportion des haplogroupes que l'on trouve en son sein et par les
haplotypes les plus fréquents de cette population.

Les haplotypes [modifier]

La signature complète de l'ADN-Y d'un homme s'appelle en théorie haplotype. Cependant, ce terme
est souvent employé abusivement et ne se référe généralement qu'à la signature partielle de l'ADN-Y.
Il existe plusieurs façons de caractériser un haplotype mais la façon largement la plus répandue est
l'utilisation des marqueurs STR.
On définit parfois des haplotypes modèles. Un des plus fameux est le CMH (Cohen Modal
Haplotype). Celui-ci est obsolète mais nous l'utilisons pour illustrer le concept. Il est défini par 6 marqueurs
DYS. Si l'on teste l'ADN-Y d'un homme avec ces 6 marqueurs et que le nombre de répétitions de séquences
pour chacun des marqueurs est la suivante alors on dit que cet homme réagit positivement au CMH.
Cohen Modal Haplotype
 DYS393 DYS390 DYS19 DYS391 DYS388 DYS392
12 23 14 10 16 11
Il était sensé définir l'haplotype de tous les Cohen et uniquement des Cohen. Mais on s'est aperçu
que sa résolution n'était pas assez grande et de ce fait un très grand nombre d'être humains répondaient
positivement au test. Un CMH étendu a été redéfini, il correspond bien uniquement à des Cohen mais à une
partie seulement des Cohen.
Il existe d'autres haplotypes modèles comme l'Atlantic Modal Haplotype (AMH) ou haplotype 15 qui
est porté par une très grande majorité d'hommes habitant l'ouest de l'Europe. Il y a parfois un accord entre
un haplotype et un haplogroupe. C'est le cas du AMH qui n'est porté que par l'haplogroupe R1b et
particulièrement par le sous-haplogroupe R1b1b2.

Les autres systèmes de classification [modifier]

Il existe encore des auteurs utilisant d'autres techniques pour classifier l'ADN-Y des populations. On
peut citer le système p49a,f qui est un RFLP (Restriction Fragment Lengh Polymorphism) utilisant l'enzyme
TaqI pour couper ("restreindre") l'ADN. Il est encore pratiqué par le Professeur Lucotte à Paris. Cette sonde
permet de définir un certain nombre d'haplotypes mais il est souvent difficile de faire le lien entre ce système
et le système du YCC.

Signature génétique des chromosomes homologues [modifier]

Enfin, en plus des études sur l'ADN-mt et l'ADN-Y, il existe de nombreuses études sur les
chromosomes homologues. Dans ce cas, on ne regarde que la signature génétique des populations. Il ne
peut y avoir d'arbre généalogique des chromosomes homologues puisque ces chromosomes se mélangent
lors de la meiose. Par ailleurs, les chromosomes homologues sont sujets à la sélection naturelle ce qui pose
des problèmes lorsqu'on compare des populations vivant dans des milieux différents.

Définition de la population [modifier]

La population étudiée par la génétique des populations est un ensemble d'individus qui montrent une
unité de reproduction : les individus d'une population peuvent se croiser entre eux, ils se reproduisent moins
avec les individus des populations voisines, desquelles ils sont géographiquement isolés. Une population
n'est donc pas une espèce, mais est déterminée par des critères d'ordres spatiaux, temporels et par un
patrimoine génétique, qui est un génome collectif, somme de génotypes individuels (pools de gènes).
L'évolution du patrimoine génétique au cours des générations est étudiée par la génétique des populations.
Cette population idéale reste un modèle d'étude, et correspond très rarement à la réalité. Dans la
mesure où des critères spatio-temporels rentrent en ligne de compte, les limites d'une population sont la
plupart du temps très incertaines. Ces limites dépendent ainsi de la répartition spatiale et temporelle des
individus, de leur mobilité, de leur mode de reproduction, de leur durée de vie, de leur sociabilité, etc.

Mutation, dérive, sélection et migration [modifier]

Des mutations, l'effet fondateur, la dérive génétique et les pressions de sélection variables sont à la
source de l'évolution. Elles conduisent à des différences génétiques entre populations de plus en plus
importantes, différences desquelles peut résulter la spéciation.
Mutations [modifier]
La variabilité génétique est le résultat des mutations qui font apparaître de nouveaux allèles. Une
même mutation peut avoir des effets phénotypiques différents.
• Les mutations créent de nouveaux allèles :
• mutations ponctuelles;
• remaniement chromosomique : il s'agit de modifications de la structure
chromosomique. Ces modifications ne sont guère favorables. Il en existe plusieurs types
(translocation, délétion, duplication, inversion);
• changement du nombre de chromosomes :
• aneuploïdie : perte ou ajout de chromosomes (voir, par exemple, trisomie 21)
• polyploïdie
• mutation neutre : on appelle neutres les mutations qui n'ont pas d'effet sur
l'organisme où elles se produisent;
• mutation létale : on appelle létales les mutations qui diminuent l'espérance de vie.

Dérive et sélection [modifier]

• La dérive génétique et la sélection provoquent des variations de fréquence des allèles de
façon interne aux populations. La première est l'effet du hasard (particulièrement efficace quand la
population est d'effectif réduit). La seconde est due aux effets des allèles sur les chances de
reproduction des organismes qui les possèdent.
Article détaillé : valeur sélective.
Migrations [modifier]
Article détaillé : Flux génétique.
• L'effet fondateur : la fréquence allélique d'un groupe migrant peut ne pas être représentative
de la population dont il est issu. Par exemple, un allèle peu fréquent peut être sur-représenté.
• Les migrations sont l'occasion de transmission d'allèles d'une population à l'autre. Elles
modifient bien évidemment la fréquence des allèles dans les populations concernées.

Régimes de reproduction [modifier]

L'efficacité de la sélection dépend du régime de reproduction. Les modèles de génétique des
populations prennent donc en compte ce paramètre.
Au sein d'une population, tous les individus peuvent se reproduire entre eux avec la même
probabilité (on dit alors que la population est panmictique). Dans le cas contraire, ils peuvent se reproduire
davantage avec eux-mêmes (possible chez les espèces hermaphrodites) ou avec des apparentés - plus
proches géographiquement - qu'avec les autres individus de la population. On parle alors de régime fermé,
ou consanguin. Enfin, ils peuvent se reproduire moins souvent avec eux-mêmes ou leurs proches qu'avec le
reste de la population (par exemple s'il existe des systèmes d'auto-incompatibilité ou des règles sociales
d'évitement), et on parle alors de régime ouvert.
Lorsqu'un individu se reproduit avec lui-même, on parle d'autofécondation. Lorsqu'il se reproduit
avec d'autres individus (apparentés ou non), on parle d'allofécondation.
Méthodes d'étude de la variabilité [modifier]

Notion de polymorphisme [modifier]

La génétique des populations permet d'étudier les variabilités d'origine génétique des populations.
Cette variabilité est appelée « polymorphisme ». Une population est dite polymorphe si dans cette population
une portion d'ADN a une variation de séquence correspondant à plusieurs formes alléliques dont la plus
fréquente ne dépasse pas plus d'une certaine fraction de la population totale, entre 95 ou 99 pour cent.
Dans une population, on dit qu'un gène est polymorphe, s'il possède au moins deux allèles ayant
une fréquence supérieure ou égale a 1 %. S'il ne possède pas deux allèles avec une fréquence supérieure
ou égale à 1 % mais que le gène existe quand même en plusieurs exemplaires, il est polyallélique (un gène
polymorphe est donc obligatoirement polyallélique).
• monomorphisme : les gènes monomorphes n'ont pas de variabilité.
• cryptopolymorphisme : moins de 1 % ; les maladies génétiques humaines sont en général
dans ce cas.

Mesure de la diversité génétique [modifier]

On peut calculer la fréquence des phénotypes observés quand une population est polymorphe pour
un caractère donné. Dans une population de N individus dont Nx ont tel caractère x et Ny tel autre caractère
y:
• fréquence du phénotype x : f[x] = Nx/N
• fréquence du phénotype y : f[y] = Ny/N
 Dans le cas d'un gène à deux allèles A et a, a étant récessif, seule la fréquence phénotypique de aa
peut être calculée, puisque l'on ne peut distinguer Aa et AA au niveau du phénotype.
En revanche, s'il y a trois caractères (x, y, z) gouvernés par deux allèles co-dominants (A1, A2), les
phénotypes permettent de distinguer les trois génotypes possibles, et il sera cette fois possible de calculer la
fréquence génotypique :
• f(A1,A1) = Nx/N
• f(A1,A2) = Ny/N
• f(A2,A2) = Nz/N
La fréquence génotypique permet ensuite de calculer la fréquence allélique (la mesure de
l'abondance d'un allèle dans une population).

Notion de population théorique idéale et loi de Hardy-Weinberg [modifier]

La prévision de la variabilité génétique d'une population est très difficile à réaliser du fait des
mutations, de la transmission simultanée de plusieurs gènes qui peut entraîner des interactions, etc. Pour
éviter ces problèmes, on peut tenter de formuler des hypothèses par la définition d'un modèle où les
croisements seraient réellement aléatoires (croisements panmictiques), sans migration ni mutation, en
faisant abstraction de la sélection (tous les individus ont les mêmes chances de se reproduire). L'étude doit
se faire au sein d'une population suffisamment grande pour être considérée comme infinie, afin de pouvoir
identifier la probabilité d'obtention de chaque génotype à sa fréquence effective d'apparition.
 Dans ce modèle, les fréquences des allèles et des génotypes suivent une loi, la loi de Hardy-
Weinberg, qui est le modèle de référence de la génétique des populations. Cette loi énonce que les
fréquences alléliques et les fréquences génotypiques restent stables de génération en génération.
Cette loi n'est jamais parfaitement observée dans la nature, et, en réalité ce sont les écarts au
modèle qui sont les plus informatifs. Sur un grand nombre de générations, un écart à la panmixie calculée
selon Hardy-Weinberg peut en effet suggérer l'existence d'un processus évolutif sous-jacent, une pression
de sélection, un phénomène d'autofécondation, ou un choix du conjoint (génotypes modulant le potentiel
reproductif).
Voir aussi : principe de Hardy-Weinberg

Applications [modifier]
Rubriques à améliorer
• Étude du polymorphisme phénotypique (morphologique).
• Étude du polymorphisme des protéines :
• polymorphisme enzymatique par électrophorèse
• polymorphisme immunologique
• polymorphisme de l'ADN
• Étude de l'ADN fossile.
• Le génome mitochondrial humain, dont la transmission est uniparentale (par la mère)
possède une importance privilégiée pour étudier l'évolution, car son taux de mutation est élevé, il n'a
pas de recombinaison méiotique et ses variations ne sont donc dues qu'à des mutations cumulées
 (pas de métissage). Sa variation est donc lente et se prête par ailleurs très bien au calcul de distance
génétique sur des périodes relativement brèves. Néanmoins, il ne donne pas d'informations sur
l'ADN nucléaire qui évolue indépendamment. L'étude de l'ADN mitochondrial a ainsi montré que tous
les ADNmt actuels dérivaient d’une ancêtre commune appelée ève vivant en Afrique, il y a environ
150 000 ans[3].
• Un autre marqueur génétique uniparental qui permet le même genre d'études est le
chromosome Y. Ainsi, le plus récent ancêtre patrilinéaire commun est l'homme de qui tous les
chromosomes Y des hommes vivants descendent. En analysant l'ADN de gens dans plusieurs
régions du monde, le généticien Spencer Wells a conclu que tous les humains vivant aujourd'hui
sont les descendants d'un homme qui a vécu en Afrique il y a environ 60 000 ans[4].
• déterminisme monogénique
• déterminisme polygénique

Notes et références [modifier]

1. ↑ Spencer Wells, The Journey of Man: A Genetic Odyssey, p. 55. Random House, ISBN 0-
8129-7146-9
2. ↑
IMS-JST : Institute of Medical Science-Japan Science and Technology Agency, Japan
L : The Family Tree DNA Genomic Research Center, Houston, Texas, United States of
America
M : Stanford University, California, United States of America
 P : University of Arizona, Arizona, United States of America
PK : Biomedical and Genetic Engineering Laboratories, Islamabad, Pakistan
U : University of Central Florida, Florida, United States of America
V : La Sapienza, Rome, Italy
3. ↑ The genographic project du National geographic est mené en partenariat avec IBM depuis
2007.
4. ↑ (en) Documentary Redraws Humans' Family Tree [archive]

Liens internes [modifier]

Génétique au sens large
• Génétique
• Génétique quantitative
• Génétique de la conservation
• Théorie synthétique de l'évolution
• O.G.M.
Génétique des populations humaine
Un tableau synthétique, donnant le pourcentage de chaque haplogroupe du chromosome Y en
fonction des différentes populations, est fourni sur la page du Wikipedia en anglais Y-
DNA_haplogroups_by_ethnic_groups.
 Différentes études ont été menées sur d'autres groupes que les Juifs afin de mieux comprendre leur
origine. Certaines sont abordées sous Wikipedia :
• Origine multirégionale de l'homme moderne
• Origine africaine de l'homme moderne
• Projet génographique
• Études génétiques sur les Juifs
• Les Amérindiens
• Les Azéris
• Les Basques
• Les Étrusques
• Les Finnois
• Les Iraniens
• Les Maghrébins
• Les Peuls
• Les Roms

Lien externe [modifier]

• Cours de génétique des populations de l'université de Lyon
Le site Eupedia dédié principalement aux haplogroupes des populations européennes constitue une
excellente introduction à la génétique des populations d'autant plus que celui-ci est en français et il est, de
plus, régulièrement mis à jour.
 Trois projets privés tentent de cartographier le génome de l'ensemble des populations du globe et de
tracer leurs migrations. Il s'agit de Genebase, du projet Genographic de national Geographic et du DNA
Ancestry project. Seul le premier site donne accès à une base complète des haplogroupes par populations.

• Portail de la biologie cellulaire et moléculaire

Ce document provient de « http://fr.wikipedia.org/wiki/G%C3%A9n%C3%A9tique_des_populations
».
 W000

Systematics
From Wikipedia, the free encyclopedia

Jump to: navigation, search

"Systematic" redirects here. For the band, see Systematic (band).
For other uses, see Systematics (disambiguation).
 A comparison of phylogenetic and phenetic concepts
Biological systematics is the study of the diversification of life on the planet Earth, both past and
present, and the relationships among living things through time. Relationships are visualized as evolutionary
trees (synonyms: cladograms, phylogenetic trees, phylogenies). Phylogenies have two components,
branching order (showing group relationships) and branch length (showing amount of evolution).
Phylogenetic trees of species and higher taxa are used to study the evolution of traits (e.g., anatomical or
molecular characteristics) and the distribution of organisms (biogeography). Systematics, in other words, is
used to understand the evolutionary history of life on Earth.
"Systematic biology" and "taxonomy" (terms that are often confused and used interchangeably) were
defined in relationship to one another as follows:[1]
Systematic biology (hereafter called simply systematics) is the field that (a) provides scientific
names for organisms, (b) describes them, (c) preserves collections of them, (d) provides
classifications for the organisms, keys for their identification, and data on their distributions, (e)
investigates their evolutionary histories, and (f) considers their environmental adaptations. This
is a field with a long history that in recent years has experienced a notable renaissance,
principally with respect to theoretical content. Part of the theoretical material has to do with
evolutionary areas (topics e and f above), the rest relates especially to the problem of
classification. Taxonomy is that part of systematics concerned with topics (a) to (d) above.

The term "systematics" is sometimes used synonymously with "taxonomy" and may be confused with
"scientific classification". However, taxonomy is more specifically the identification, description and naming
(i.e. nomenclature) of organisms, while "classification" is focused on placing organisms within hierarchical
groups that show their relationships to other organisms. All of these biological disciplines can be involved
with extinct and extant organisms. However, systematics alone deals specifically with relationships through
time, and can be synonymous with phylogenetics, broadly dealing with the inferred hierarchy of organisms.
Systematics uses taxonomy as a primary tool in understanding organisms, as nothing about an
organism's relationships with other living things can be understood without it first being properly studied and
described in sufficient detail to identify and classify it correctly. Scientific classifications are aids in recording
and reporting information to other scientists and to laymen. The systematist, a scientist who specializes in
systematics, must, therefore, be able to use existing classification systems, or at least know them well
enough to skillfully justify not using them.
Phenetic systematics was an attempt to determine the relationships of organisms through a measure
of similarity, considering plesiomorphies (ancestral traits) and apomorphies (derived traits) to be equally
informative. From the 20th century onwards, it was superseded by cladistics, which considers plesiomorphies
to be uninformative for an attempt to resolve the phylogeny of Earth's various organisms through time.
Today's systematists generally make extensive use of molecular biology and computer programs to study
organisms.
Systematics is fundamental to biology because it is the foundation for all studies of organisms, by
showing how any organism relates to other living things (ancestor-descendant relationships).
Systematics is also of major importance in understanding conservation issues because it attempts to
explain the Earth's biodiversity and could be used to assist in allocating limited means to preserve and
protect endangered species, by looking at, for example, the genetic diversity among various taxa of plants or
animals and deciding how much of that to preserve.

[edit] See also

• Cladistics - a popular methodology in systematics
• Phenetics - an obsolete methodology, predecessor to cladistics
• Phylogeny - what is analyzed in systematics and taxonomy
• 16S ribosomal RNA - the most intensively studied nucleic acid in phylogenetics
 • Phylogenetic comparative methods - use of evolutionary trees to study biodiversity and
comparative biology
• Scientific classification - the result of research in systematics and taxonomy
• Taxonomy - a branch of the biological sciences related to systematics

[edit] References
[edit] Notes
1. ^ Michener, Charles D., John O. Corliss, Richard S. Cowan, Peter H. Raven, Curtis W.
Sabrosky, Donald S. Squires, and G. W. Wharton (1970). Systematics In Support of Biological
Research. Division of Biology and Agriculture, National Research Council. Washington, D.C. 25 pp.

[edit] Further reading

• Schuh, Randall T. and Andrew V. Z. Brower. 2009. Biological Systematics: Principles and
Applications, 2nd edn. ISBN 978-0-8014-4799-0
• Simpson, Michael G. 2005. Plant Systematics. ISBN 978-0-12-644460-5

[edit] External links

• Society of Australian Systematic Biologists
• Society of Systematic Biologists
• Willi Hennig Society
 [hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Systematics"

Categories: Evolutionary biology | Scientific classification

 w000

Systématique
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Pour les articles homonymes, voir Systématique (homonymie).

En sciences de la vie et en histoire naturelle, la systématique est la science qui a pour objet de
dénombrer et de classer les taxons dans un certain ordre, basé sur des principes divers. Elle ne doit pas être
confondue avec la systémique, qui est une méthode qui aborde une question en prenant en compte
l'ensemble de ses composants et de leurs interactions, l'ensemble du système.
Sommaire
[masquer]
• 1 Définition
• 2 Évolutions récentes
• 3 Notes et références
• 4 Voir aussi
• 4.1 Articles connexes
• 4.2 Liens externes

Définition [modifier]
La systématique n'est pas synonyme de taxinomie, qui s'attache à décrire et définir les taxons, mais
plutôt son prolongement.
La confusion entre les deux termes vient du fait que les taxinomistes ont de tout temps été également
nommés systématiciens car, après avoir étudié et décrit des organismes, ils ont tout naturellement essayé de
les classer à partir du bas niveau des espèces (alpha taxonomy ou « taxinomie primaire »).
Dans la pratique, le terme « systématique » désigne aussi bien la méthode utilisée (on dira par
exemple la « systématique phylogénétique ») que le résultat obtenu avec cette méthode (Ex : la
« systématique des Agaricales »).
 Dans le sens concret de résultat, les deux sciences sont peu distinctes et souvent confondues, car
pratiquées simultanément par les mêmes personnes.
Ceux qui utilisent surtout le sens de méthode, les phylogénéticiens notamment, nomment souvent le
résultat classification ou encore « taxinomie ».

Évolutions récentes [modifier]

• Les progrès fait dans la génétique conduisent à revoir le classement de certains taxons dans
l'arbre phylogénétique du Vivant. Les approches systémiques et fonctionnelles de la biodiversité
tendent à prendre de l'importance, mais la systématique reste fondamentale pour inventorier et pour
évaluer les évolutions du Vivant.
• Les progrès de l'informatique, qui ont permis de développer la bio-informatique, et du travail
collaboratif (à l'origine du réseau Tela-botanica qui associe environ 11 000 botanises francophones
dans 35 pays (fin 2009)
À titre d'exemple, un logiciel d'identification (de plantes) assistée par ordinateur (IDAO [1]) permet à
des non-spécialistes d'identifier plus rapidement une plante sans passer par une complexe clé de
détermination hiérarchique, sur la base d'une méthode de type "portrait-robot" de plantes. Cela
facilite l'identification des plantes par des "amateurs" (ou parataxonomistes). Il est aussi utilisé par le
projet « Pl@ntnet » avec l'Université de Montpellier et Tela-botanica).
• De nombreuses études basées sur des protocoles de type « Sciences citoyennes » sont en
développement dans le monde.
 Notes et références [modifier]
1. ↑ Logiciel développé par Pierre Grad, agronome du CIRAD

Voir aussi [modifier]

Articles connexes [modifier] Voir
• Taxinomie « systématique » sur le
• Taxon Wiktionnaire.
• Cladistique
• Classification scientifique
• Classification classique
• Classification phylogénétique
• Francisation des termes en latin scientifique
• Biologie
• Bioinformatique

Liens externes [modifier]

• Sélection de sites web sur la systématique et l’évolution dans le répertoire encyclopédique :
Les Signets de la Bibliothèque nationale de France
 Taxinomie

Règne - Sous-règne - Infra-règne -

Superembranchement - Embranchement / Division - Sous-embranchement -
Superclasse - Classe - Sous-classe - Infraclasse -
Superordre - Ordre - Sous-ordre - Infraordre
Superfamille - Famille - Sous-famille - Tribu -
Genre - Sous-genre - Section - Série -
Espèce - Sous-espèce -
Variété ou Race - Forme ou Type

Biologie - Nomenclature - Nom binominal

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Génomique · Géobiologie · Microbiologie · Origine de la vie · Paléontologie · Parasitologie · Physiologie ·
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 • Portail de l’origine et de l’évolution du vivant
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Objections to evolution
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Biology portal · v • d • e
Objections to evolution have been raised since evolutionary ideas came to prominence in the 19th
century.[1] When Charles Darwin published his 1859 book On the Origin of Species, his theory of evolution
by natural selection initially met opposition from alternate scientific theories, but came to be universally
accepted by the scientific community.[2] The observation of evolutionary processes occurring, as well as the
current theory explaining that evidence, have been uncontroversial among mainstream biologists for nearly a
century.[3]
Since then, nearly all criticisms of evolution have come from religious sources, rather than from the
scientific community.[4] Although most religions have accepted the occurrence of evolution, such as those
advocating theistic evolution, there still exist religious beliefs which reject evolutionary explanations in favor
of creationism, the belief that a deity supernaturally created the world largely in its current form.[5] The
resultant U.S. centric creation-evolution controversy has been a focal point of recent conflict between religion
and science.
In contrast to earlier objections to evolution that were either strictly scientific or explicitly religious,
recent objections to evolution have frequently blurred the distinction between the two. Such objections have
often centered on undermining evolution's scientific basis, with the intent of combating the teaching of
evolution as fact and opposing the spread of "atheistic materialism".[6] Modern creationism is characterized
by movements such as Creation Science, neocreationism, and Intelligent Design which argue that the idea of
life being directly "designed" by a God or intelligence is at least as scientific as evolutionary theory, and
should therefore be taught in public schools. Their arguments against evolution have become widespread,
and include objections to evolution's evidence, methodology, plausibility, morality, and scientific acceptance.
The scientific community, however, does not accept such objections as having any validity, citing detractors'
misinterpretations of scientific method, evidence, and basic physical laws.[7]
While objections primarily originate from the United States, there is widespread belief in creationism
in the Muslim world,[8] South Africa and India[9] with smaller followings in Australia, New Zealand, the United
Kingdom and Canada.[10] However, in India 77% of respondents who had heard of Charles Darwin and
knew something about the theory of evolution agreed it was backed by scientific evidence.[11]
Contents
[hide]
• 1 Defining evolution
• 2 History
• 3 Scientific acceptance
• 3.1 Status as a theory
• 3.2 Degree of acceptance
• 4 Scientific status
• 4.1 Religious nature
• 4.2 Unfalsifiability
• 5 Evidence
• 5.1 Lack of observation
• 5.2 Instability of evidence
• 5.3 Unreliable or inconsistent evidence
• 6 Plausibility
• 6.1 Improbability
• 6.2 Unexplained aspects of the natural world
• 7 Implausibility
• 7.1 Creation of complex structures
• 7.2 Creation of information
• 7.3 Violation of the second law of thermodynamics
[edit] Defining evolution
Further information: Evolution (term)
One of the main sources of confusion and ambiguity in the creation-evolution debate is the definition
of evolution itself. In the context of biology, evolution is genetic changes in populations of organisms over
successive generations. However, the word has a number of different meanings in different fields, from
evolutionary computation to molecular evolution to sociocultural evolution to stellar and galactic evolution. It
can even refer to metaphysical evolution, spiritual evolution, or any of a number of evolutionist philosophies.
When biological evolution is conflated with other evolutionary processes, this can cause errors such as the
claim that modern evolutionary theory says anything about abiogenesis or the Big Bang.[12]
In colloquial contexts, evolution can refer to any sort of progressive development, and often bears a
connotation of gradual improvement: here evolution is understood as a process that results in greater quality
or complexity. This common definition, when misapplied to biological evolution, leads to frequent
misunderstandings. For example, the idea of devolution ("backwards" evolution) is a result of erroneously
assuming that evolution is directional or has a specific goal in mind (cf. orthogenesis). In reality, the evolution
of an organism has no "objective" other than increasing the organism's ability to survive and reproduce in its
environment; and its suitability is only defined in relation to this environment. Biologists do not consider any
one species, such as humans, to be more highly evolved or advanced than another. Certain sources have
been criticized for indicating otherwise due to a tendency to evaluate nonhuman organisms according to
anthropocentric standards rather than more objective ones.[13]
Evolution also does not require that organisms become more complex. Although the history of life
shows an apparent trend towards the evolution of complexity; there is a question if this appearance of
increased complexity is real, or if this conclusion comes from neglecting the fact that the majority of life on
earth has always consisted of prokaryotes.[14] In this view, complexity is not a necessary consequence of
evolution; rather, it is a consequence of the specific circumstances of evolution on Earth, which frequently
made greater complexity advantageous, and thus naturally selected for. Depending on the situation,
organisms' complexity can either increase, decrease, or stay the same, and all three of these trends have
been observed in evolution.[13]
Creationist sources frequently define evolution according to a colloquial, rather than scientific,
meaning.[15] As a result, many attempts to rebut evolution do not address the findings of evolutionary
biology (see straw man argument). This also means that advocates of creationism and evolutionary biologists
often simply speak past each other.[12][16]

[edit] History
Further information: History of evolutionary thought, History of creationism and Creation-evolution
controversy
 Darwin's theory of evolution through common descent gained widespread acceptance, but there was
continued resistance to his views on the significance of natural selection.
Various evolutionary ideas came to prominence around the start of the nineteenth century, in
particular the transmutation of species theory put forward by Jean-Baptiste Lamarck. These were opposed
on scientific grounds, most notably by Georges Cuvier, as well as meeting political and religious objections.
[1] These ideas that natural laws controlled the development of nature and society gained vast popular
audiences with George Combe's The Constitution of Man of 1828 and the anonymous Vestiges of the
Natural History of Creation of 1844. When Charles Darwin published his 1859 book On the Origin of Species,
within fifteen to twenty years he convinced most of the scientific community that evolution through common
descent was true, but while most accepted that natural selection was a valid and empirically testable
hypothesis, his view that it was the primary mechanism of evolution was generally rejected.[2]
The earliest objections to Darwinian evolution were both scientific and religious. Darwin's
contemporaries eventually came to accept the transmutation of species based upon fossil evidence; forming
the X Club to defend it against the church and wealthy amateurs,[17] although the specific evolutionary
mechanism which Darwin provided – natural selection – was actively disputed by alternative theories such as
Lamarckism and orthogenesis. Darwin's gradualistic account was also opposed by saltationism and
catastrophism. Lord Kelvin led scientific opposition to gradualism on the basis of his thermodynamic
calculations that the Age of the Earth was between 24 and 400 million years old, an estimate strongly
disputed by geologists. These figures were corrected in 1907 when radioactive dating of rocks showed that
the Earth was billions of years old.[18][19] Kelvin's own views favoured a version of theistic evolution
speeded up by divine guidance.[20] The specific hereditary mechanism Darwin provided, pangenesis, lacked
any supporting evidence. In the early 20th century, pangenesis was replaced by Mendelian inheritance,
leading to the rise of the modern evolutionary synthesis. The modern synthesis rose to universal acceptance
among biologists with the help of new evidence, such as genetics, which confirmed Darwin's predictions and
refuted the competing theories.[21]
Protestantism, especially in America, broke out in "acrid polemics" and argument about evolution
from 1860 to the 1870s – with the turning point possibly marked with the death of Louis Agassiz in 1873 – and
by 1880 a form of "Christian evolution" was becoming the consensus.[22] In Britain while publication of The
Descent of Man by Darwin in 1871 reinvigorated debate from the previous decade, Sir Henry Chadwick notes
a steady acceptance of evolution "among more educated Christians" between 1860 and 1885. This led
evolutionary theory to be "both permissible and respectable" by 1876.[22] Frederick Temple's lectures on
Relations between Religion and Science (1884) speaking on how evolution was not "antagonistic" to religion
highlighted this trend.[23] Temple's appointment to Archbishop of Canterbury in 1896 showed the broad
acceptance of evolution within the church hierarchy.[22]
For decades Catholicism avoided official refutation of evolution. However, it would rein in Catholics
who proposed that evolution could be reconciled with the Bible, as this conflicted with First Vatican Council's
(1869–70) finding that everything was created out of nothing by God, and to question that could lead to
excommunication. In 1950, the encyclical Humani Generis of Pope Pius XII first mentioned evolution directly
and officially.[24] It allowed for inquiry into humans coming from pre-existing living matter, but to not question
Adam and Eve or the creation of the soul. In 1996, Pope John Paul II stated evolution was "more than a
hypothesis" and acknowledged the large body of work accumulated in its support, but reiterated that any
attempt to give a material explanation of the human soul was "incompatible with the truth about man."[25]
Muslim reaction covered the gamut with those believing in literal creation from the Qur'an while many
educated Muslims subscribed to a version of theistic or guided evolution where the Qur'an reinforced rather
than contradicted mainstream science. This occurred relatively quickly as medieval Madrasah's taught ideas
of Al-Jahiz, a Muslim scholar from the 9th century, who proposed concepts similar to natural selection.[8]
However, acceptance of evolution remains low in the Muslim world as prominent figures reject evolution's
underpinning philosophy of materialism as unsound to human origins and a denial of Allah.[8] Further
objections by Muslim scholars and writers largely reflect those put forward in the Western world.[26]
Regardless of acceptance from major religious hierarchies, early religious objections to Darwin's
theory are still used in opposition to evolution. The ideas that species change over time through natural
processes and that different species share common ancestors seemed to contradict the Genesis account of
Creation. Believers in Biblical infallibility attacked Darwinism as heretical. The natural theology of the early
19th century was typified by Paley's watchmaker analogy, an argument from design still used by the
creationist movement. Natural theology included a range of ideas and arguments from the outset, and when
Darwin's theory was published, ideas of theistic evolution were presented in which evolution is accepted as a
secondary cause open to scientific investigation, while still holding belief in God as a first cause with a non-
specified role in guiding evolution and creating humans.[27] This position has been adopted by
denominations of Christianity and Judaism in line with modernist theology which views the Bible and Torah
as allegorical thus removing the conflict between evolution and religion.
However, in the 1920s Christian fundamentalists in the United States developed their literalist
arguments against Modernist theology into opposition to the teaching of evolution due to fears that
‘‘Darwinism’’ had led to German militarism and was a threat to religion and morality. This opposition
developed into the creation-evolution controversy involving Christian literalists in the United States objecting
to the teaching of evolution in public schools. Although early objections dismissed evolution for contradicting
their interpretation of the Bible, this argument was invalidated when the Supreme Court ruled in Epperson v.
Arkansas that forbidding the teaching of evolution on religious grounds violated the Establishment Clause.
[28]
 Since then creationists have developed more nuanced objections to evolution, alleging variously that
it is unscientific, infringes on creationists' religious freedoms or that the acceptance of evolution is a religious
stance.[29] Creationists have appealed to democratic principles of fairness, arguing that evolution is
controversial, and that science classrooms should therefore "Teach the Controversy".[30] These objections
to evolution culminated in the intelligent design movement in the early 2000s that unsuccessfully attempted
to present itself as a scientific alternative to evolution.[31][32]

[edit] Scientific acceptance

Recent objections to evolutionary theory have focused on its scientific validity,[33][34] or attempting
to come up with alternative ideas such as Creationism to debate its findings.[35] Creationists often argue, for
example, that evolution is unproven, non-factual, or controversial.[36]

[edit] Status as a theory

Further information: Evolution as theory and fact
Critics of evolution frequently assert that evolution is "just a theory", with the intent of emphasizing
evolution's unproven nature, or of characterizing it as a matter of opinion rather than of fact or evidence. This
reflects a misunderstanding of the meaning of theory in a scientific context: whereas in colloquial speech a
theory is a conjecture or guess, in science a theory is simply an explanation or model of the world that makes
testable predictions. When evolution is used to describe a theory, it refers to an explanation for the diversity
of species and their ancestry. An example of evolution as theory is the modern synthesis of Darwinian natural
selection and Mendelian inheritance. As with any scientific theory, the modern synthesis is constantly
debated, tested, and refined by scientists. There is an overwhelming consensus in the scientific community
that it remains the only robust model that accounts for the known facts concerning evolution.[37]
Critics state that evolution is not a fact.[38] In science, a fact is a verified empirical observation; in
colloquial contexts, however, a fact can simply refer to anything for which there is overwhelming evidence.
For example, in common usage theories such as "the Earth revolves around the Sun" and "objects fall due to
gravity" may be referred to as "facts", even though they are purely theoretical. From a scientific standpoint,
therefore, evolution may be called a "fact" for the same reason that gravity can: under the scientific definition,
evolution is an observable process that occurs whenever a population of organisms genetically changes over
time. Under the colloquial definition, the theory of evolution can also be called a fact, referring to this theory's
well-established nature. Thus, evolution is widely considered both a theory and a fact by scientists.[39][40]
[41]
Similar confusion is involved in objections that evolution is "unproven";[42] strict proof is possible
only in formal sciences such as logic and mathematics, not natural sciences (where the proper term is
"validated"), so this is trivially true, and no more an indictment of evolution than calling it a "theory". The
confusion arises, however, in that the colloquial meaning of proof is simply "compelling evidence", in which
case scientists would indeed consider evolution "proven". The distinction is an important one in philosophy of
science, as it relates to the lack of absolute certainty in all empirical claims, not just evolution.[43]

[edit] Degree of acceptance

Further information: Level of support for evolution
In 1999 a new major objection to evolution appeared in Kansas, and is in a sense self-fulfilling: it
argues that evolution is controversial or contentious.[44] Unlike past creationist arguments which sought to
abolish the teaching of evolution altogether, this argument makes the weaker claim that evolution, being
controversial, should be presented alongside other, alternative views, and students should be allowed to
evaluate and choose between the options on their own.[45] This appeal to "fairness" and a more democratic,
"balanced" approach in which conflicting views are given "equal time" appeals to American creationists, and
was endorsed by U.S. President George W. Bush.[32][42][46]
This objection forms the basis of the "Teach the Controversy" campaign, an attempt by the Discovery
Institute to promote the teaching of intelligent design in public schools. This in turn forms a major part of the
Institute's "wedge strategy", an attempt to gradually undermine evolution and ultimately to "reverse the stifling
materialist world view and replace it with a science consonant with Christian and theistic convictions".[30]
Scientists and U.S. courts have rejected this objection on the grounds that science is not based on
appeals to popularity, but on evidence. The scientific consensus of biologists, not popular opinion or fairness,
determines what is considered acceptable science, and it is argued that although evolution is clearly
controversial in the public arena, it is entirely uncontroversial among experts in the field.[47]
In response, creationists have disputed the level of scientific support for evolution. The Discovery
Institute has gathered over 600 scientists since 2001 to sign "A Scientific Dissent From Darwinism" in order
to show that there are a number of scientists who dispute what they refer to as "Darwinian evolution". This
statement did not profess outright disbelief in evolution, but expressed skepticism as to the ability of "random
mutation and natural selection to account for the complexity of life." Several counter-petitions have been
launched in turn, including A Scientific Support for Darwinism, which gathered over 7,000 signatures in four
days, and Project Steve, a tongue-in-cheek petition that has gathered 1,140 (as of August 12, 2010)
evolution-supporting scientists named "Steve".
 Creationists have argued for over a century that evolution is "a theory in crisis" that will soon be
overturned,[48] based on objections that it lacks reliable evidence or violates natural laws. These objections
have been rejected by most scientists, as have claims that intelligent design, or any other creationist
explanation, meets the basic scientific standards that would be required to make them scientific alternatives
to evolution. It is also argued that even if evidence against evolution exists, it is a false dilemma to
characterize this as evidence for intelligent design.[46][49]
A similar objection to evolution is that certain scientific authorities – mainly pre-modern ones – have
doubted or rejected evolution.[50] Most commonly, it is argued that Darwin "recanted" on his deathbed, a
false anecdote originating from the Lady Hope Story.[51] These objections are generally rejected as appeals
to authority. Even if this myth were true, it would hold no bearing on the merit of the theory itself.

[edit] Scientific status

A common neo-creationist objection to evolution is that evolution does not adhere to normal scientific
standards—that it is not genuinely scientific. It is argued that evolutionary biology does not follow the scientific
method, and therefore should not be taught in science classes, or at least should be taught alongside other
views (i.e., creationism). These objections often deal with the very nature of evolutionary theory and the
scientific method.

[edit] Religious nature

Further information: Relationship between religion and science, Scientism
Creationists commonly argue against evolution on the grounds that "evolution is a religion; it is not a
science".[52] The purpose of this criticism is to undermine the higher ground biologists claim in debating
creationists, and to reframe the debate from being between science (evolution) and religion (creationism) to
being between two equally religious beliefs – or even to argue that evolution is religious while intelligent
design is not.[53][54] Those that oppose evolution frequently refer to supporters of evolution as
"Evolutionists" or "Darwinists".[52]
The arguments for evolution being a religion generally amount to arguments by analogy: it is argued
that evolution and religion have one or more things in common, and that therefore evolution is a religion.
Examples of claims made in such arguments are statements that evolution is based on faith, that supporters
of evolution revere Darwin as a prophet, and that supporters of evolution dogmatically reject alternative
suggestions out-of-hand.[42][55] These claims have become more popular in recent years as the
neocreationist movement has sought to distance itself from religion, thus giving it more reason to make use
of a seemingly anti-religious analogy.[47]
In response, supporters of evolution have argued that no scientist's claims, including Darwin's, are
treated as sacrosanct, as shown by the aspects of Darwin's theory that have been rejected or revised by
scientists over the years, to form first Neo-Darwinism and later the modern evolutionary synthesis.[56][57]
The claim that evolution relies on faith, often based on the creationist belief that evolution has never been
observed, is likewise rejected on the grounds that evolution has strong supporting evidence, and therefore
does not require faith.
In general, the argument that evolution is religious has been rejected on the grounds that religion is
not defined by how dogmatic or zealous its adherents are, but by its spiritual or supernatural beliefs.
Evolutionary supporters point out evolution is neither dogmatic nor based on faith, and they accuse
creationists of equivocating between the strict definition of religion and its colloquial usage to refer to
anything that is enthusiastically or dogmatically engaged in. U.S. courts have also rejected this objection:
 Assuming for the purposes of argument, however, that evolution is a religion or religious tenet,
the remedy is to stop the teaching of evolution, not establish another religion in opposition to it.
Yet it is clearly established in the case law, and perhaps also in common sense, that evolution is
not a religion and that teaching evolution does not violate the Establishment Clause.[58]

A related claim is that evolution is atheistic; creationists sometimes merge the two claims and
describe evolution as an "atheistic religion" (cf. humanism).[54] This argument against evolution is also
frequently generalized into a criticism of all science; it is argued that "science is an atheistic religion", on the
grounds that its methodological naturalism is as unproven, and thus as "faith-based", as the supernatural and
theistic beliefs of creationism.[59]

[edit] Unfalsifiability
A statement is considered falsifiable if there is an observation or a test that could be made that would
demonstrate that the statement is false. Statements that are not falsifiable cannot be examined by scientific
investigation since they permit no tests that evaluate their accuracy. Creationists such as Henry M. Morris
have claimed that evolution is unfalsifiable. They claim that any observation can be fitted into the
evolutionary framework, and that therefore it is impossible to demonstrate that evolution is wrong. As such,
they claim that evolution is non-scientific.[60][61]
 Darwin and an ape, circa 1874 from Julkaistussa picture source.
However, evolution is considered falsifiable by scientists because it can make predictions that, were
they contradicted by the evidence, would falsify evolution. Several kinds of evidence have been proposed
that could falsify evolution, such as the fossil record showing no change over time, confirmation that
mutations are prevented from accumulating, or observations showing organisms being created
supernaturally or spontaneously.[60] Many of Darwin's ideas and assertions of fact have been falsified as
evolutionary science has developed and has continued to confirm his central concepts.[62] In contrast,
creationism consists largely of unsubstantiated claims that evolution has been falsified.[60] Creationist
explanations involving the direct intervention of the supernatural in the physical world are not falsifiable,
because any result of an experiment or investigation could be the unpredictable action of an omnipotent
deity.[63]
In 1976, philosopher Karl Popper said that "Darwinism is not a testable scientific theory but a
metaphysical research programme".[64] However, Popper later recanted and offered a more nuanced view of
its status:
However, Darwin's own most important contribution to the theory of evolution, his theory of
natural selection, is difficult to test. There are some tests, even some experimental tests; and in
some cases, such as the famous phenomenon known as 'industrial melanism', we can observe
natural selection happening under our very eyes, as it were. Nevertheless, really severe tests of
the theory of natural selection are hard to come by, much more so than tests of otherwise
comparable theories in physics or chemistry.[65][66]

The most direct evidence that evolutionary theory is falsifiable may be the original words of Charles
Darwin who, in chapter 6 of On the Origin of Species wrote: "If it could be demonstrated that any complex
organ existed, which could not possibly have been formed by numerous, successive, slight modifications, my
theory would absolutely break down."[67]
In response to the unfalsifiability criticism of evolutionary theory, numerous examples of potential
ways to falsify evolution have been proposed. J.B.S. Haldane, when asked what hypothetical evidence could
disprove evolution, replied "fossil rabbits in the Precambrian era".[68][69] Numerous other potential ways to
falsify evolution have also been proposed.[43] For example, the fact that humans have one fewer pair of
chromosomes than the great apes offered a testable hypotheses involving the fusion or splitting of
chromosomes from a common ancestor. The fusion hypothesis was confirmed in 2005 by discovery that
human chromosome 2 is homologous with a fusion of two chromosomes that remain separate in other
primates. Extra, inactive telomeres and centromeres remain on human chromosome 2 as a result of the
fusion.[70] The assertion of common descent could also have been disproven with the invention of DNA
analysis. If true, human DNA should be far more similar to chimpanzees and other great apes, than to other
mammals. If not, then common descent is falsified. DNA analysis has shown that humans and chimpanzees
share a large percentage of their DNA (between 95% to 99.4% depending on the measure).[71] Also, the
evolution of chimpanzees and humans from a common ancestor predicts a (geologically) recent common
ancestor. Numerous transitional fossils have since been found.[72] Hence, human evolution has passed
several falsifiable tests.
A related claim, also once used, but then abandoned, by Popper, is that natural selection is
tautological.[65] Specifically, it is often argued that the phrase "survival of the fittest" is a tautology, in that
fitness is defined as ability to survive and reproduce. However, this phrase, first used by Herbert Spencer in
1864, is rarely used by biologists. Additionally, fitness is more accurately defined as the state of possessing
traits that make survival more likely; this definition, unlike simple "survivability", avoids being trivially true.[73]
[74][75]
Similarly, it is argued that evolutionary theory is circular reasoning, in that evidence is interpreted as
supporting evolution, but evolution is required to interpret the evidence. An example of this is the claim that
geological strata are dated through the fossils they hold, but that fossils are in turn dated by the strata they
are in.[42] However, in most cases strata are not dated by their fossils, but by their position relative to other
strata and by radiometric dating, and most strata were dated before the theory of evolution was formulated.
[76]
In his book, Abusing Science: The Case Against Creationism, philosopher of science Philip Kitcher
specifically addresses the "falsifiability" question by taking into account notable philosophical critiques of
Popper by Carl Gustav Hempel and Willard Van Orman Quine that reject his definition of theory as a set of
falsifiable statements.[77] As Kitcher points out, if one took a strictly Popperian view of “theory,” observations
of Uranus when first discovered in 1781 would have “falsified” Newton’s celestial mechanics. Rather, people
suggested that another planet influenced Uranus’ orbit – and this prediction was indeed eventually confirmed.
Kitcher agrees with Popper that “there is surely something right in the idea that a science can succeed only if
it can fail.”[78] But he insists that we view scientific theories as consisting of an “elaborate collection of
statements,” some of which are not falsifiable, and others – what he calls “auxiliary hypotheses,” which are.
According to Kitcher, good scientific theories must have three features – unity, fecundity, and
independent testability of auxiliary hypotheses:
Unity
“A science should be unified .... Good theories consist of just one problem-solving strategy, or
a small family of problem-solving strategies, that can be applied to a wide range of problems” (1982:
47).
Fecundity
A great scientific theory, like Newton’s, opens up new areas of research... Because a
theory presents a new way of looking at the world, it can lead us to ask new questions,
and so to embark on new and fruitful lines of inquiry... Typically, a flourishing science is
 incomplete. At any time, it raises more questions than it can currently answer. But
incompleteness is no vice. On the contrary, incompleteness is the mother of fecundity... A
good theory should be productive; it should raise new questions and presume that those
questions can be answered without giving up its problem-solving strategies (1982: 47–48).

Auxiliary hypothesis that are independently testable

“An auxiliary hypothesis ought to be testable independently of the particular problem it is
introduced to solve, independently of the theory it is designed to save” (1982: 46) (e.g. the evidence for
the existence of Neptune is independent of the anomalies in Uranus’s orbit).
Like other definitions of theories, including Popper’s, Kitcher makes it clear that a good theory
includes statements that have (in his terms) “observational consequences.” But, like the observation of
irregularities in Uranus’s orbit, falsification is only one possible consequence of an observation. The
production of new hypotheses is another possible – and equally important – observational consequence.
Kitcher’s account of a good theory is based not only on his understanding of how physical sciences work. He
is also taking into account the way the life sciences work.
From Kitcher’s point of view, Darwinian theory not only meets the three conditions for a good
scientific theory; it is without question an extraordinarily successful theory:
The heart of Darwinian evolutionary theory is a family of problem-solving strategies, related by
their common employment of a particular style of historical narrative. A Darwinian history is a
piece of reasoning of the following general form. The first step consists in a description of an
ancestral population of organisms. The reasoning proceeds by tracing the modification of the
population through subsequent generations, showing how characteristics were selected,
 inherited, and became prevalent. Reasoning like this can be used to answer a host of biological
questions.[79]
The same kind of story can be told again and again to answer all sorts of questions about all
sorts of living things. Evolutionary theory is unified because so many diverse questions ... can
be addressed by advancing Darwinian histories. Moreover, these narratives constantly make
claims that are subject to independent check.[80]
Darwin not only provided a scheme for unifying the diversity of life. He also gave a structure to
our ignorance. After Darwin, it was important to resolve general issues about the
presuppositions of Darwinian histories. The way in which biology should proceed had been
made admirably plain, and it was clear that biologists had to tackle questions for which they had,
as yet, no answers.[81]

[edit] Evidence
Further information: Evidence of evolution
Objections to the evidence that evolution occurs tend to be more concrete and specific, often
involving direct analysis of evolutionary biology's methods and claims.
[edit] Lack of observation

Transitional species such as the Archaeopteryx have been a fixture of the creation-evolution debate
for almost 150 years.
A common claim of creationists is that evolution has never been observed.[82] Challenges to such
objections often come down to debates over how evolution is defined (see above). Under the conventional
biological definition of evolution, it is a simple matter to observe evolution occurring. Evolutionary processes,
in the form of populations changing their genetic composition from generation to generation, have been
observed in different scientific contexts, including the evolution of fruit flies, mice and bacteria in the
laboratory,[83] and of tilapia in the field. Such studies on experimental evolution, particularly those using
microorganisms, are now providing important insights into how evolution occurs.[83][84]
In response to such examples, creationists specify that they are objecting only to macroevolution, not
microevolution:[85][86] most creationist organizations do not dispute the occurrence of short-term, relatively
minor evolutionary changes, such as that observed even in dog breeding. Rather, they dispute the
occurrence of major evolutionary changes over long periods of time, which by definition cannot be directly
observed, only inferred from microevolutionary processes and the traces of macroevolutionary ones.
However, as biologists define macroevolution, both microevolution and macroevolution have been
observed. Speciations, for example, have been directly observed many times, despite popular
misconceptions to the contrary.[87] Additionally, the modern evolutionary synthesis draws no distinction
between macroevolution and microevolution, considering the former to simply be the latter on a larger scale.
[43][88] An example of this is ring species.
Additionally, past macroevolution can be inferred from historical traces. Transitional fossils, for
example, provide plausible links between several different groups of organisms, such as Archaeopteryx
linking birds and dinosaurs,[89] or the recently-discovered Tiktaalik linking fish and limbed amphibians.[90]
Creationists dispute such examples, from asserting that such fossils are hoaxes or that they belong
exclusively to one group or the other, to asserting that there should be far more evidence of obvious
transitional species.[91] Darwin himself found the paucity of transitional species to be one of the greatest
weaknesses of his theory: "Why then is not every geological formation and every stratum full of such
intermediate links? Geology assuredly does not reveal any such finely graduated organic chain; and this,
perhaps, is the most obvious and gravest objection which can be urged against my theory. The explanation
lies, as I believe, in the extreme imperfection of the geological record." Darwin appealed to the limited
collections then available, the extreme lengths of time involved, and different rates of change with some living
species differing very little from fossils of the Silurian period. In later editions he added "that the periods
during which species have been undergoing modification, though very long as measured by years, have
probably been short in comparison with the periods during which these same species remained without
undergoing any change."[92] The number of clear transitional fossils has increased enormously since
Darwin's day, and this problem has been largely resolved with the advent of the theory of punctuated
equilibrium, which predicts a primarily stable fossil record broken up by occasional major speciations.[93]
Creationists counter that even observed speciations and transitional fossils are insufficient evidence
for the vast changes summarized by such phrases as "fish to philosophers" or "particles to people".[94] As
more and more compelling direct evidence for inter-species and species-to-species evolution has been
gathered, creationists have redefined their understanding of what amounts to a "created kind", and have
continued to insist that more dramatic demonstrations of evolution be experimentally produced.[95] One
version of this objection is "Were you there?", popularized by Ken Ham. It argues that because no one except
God could directly observe events in the distant past, scientific claims are just speculation or "story-telling".
[96][97] DNA sequences of the genomes of organisms allow an independent test of their predicted
relationships, since species which diverged more recently will be more closely related genetically than
species which are more distantly related; such phylogenetic trees show a hierarchical organization within the
tree of life, as predicted by common descent.[98][99]
In fields such as astrophysics or meteorology, where direct observation or laboratory experiments are
difficult or impossible, the scientific method instead relies on observation and logical inference. In such fields,
the test of falsifiability is satisfied when a theory is used to predict the results of new observations. When
such observations contradict a theory's predictions, it may be revised or discarded if an alternative better
explains the observed facts. For example, Newton's theory of gravitation was replaced by Einstein's theory of
General Relativity when the latter was observed to more precisely predict the orbit of Mercury.[100]

[edit] Instability of evidence

A related objection is that evolution is based on unreliable evidence. This objection goes further than
the less substantial "evolution isn't proven" arguments, claiming that evolution isn't even well-evidenced.
Typically, this is either based on the argument that evolution's evidence is full of frauds and hoaxes, that
current evidence for evolution is likely to be overturned because past evidence has been, or that certain
types of evidence are inconsistent and dubious.
Arguments against evolution's reliability are thus often based on analyzing the history of evolutionary
thought or the history of science in general. Creationists point out that in the past, major scientific revolutions
have overturned theories that were at the time considered near-certain. They thus claim that current
evolutionary theory is likely to undergo such a revolution in the future, on the basis that it is a "theory in crisis"
for one reason or another.[101]
 Romanes's 1892 copy of Ernst Haeckel's embryo drawings, often attributed incorrectly to Haeckel.
[102]
Critics of evolution commonly appeal to past scientific hoaxes such as the Piltdown Man forgery. It is
argued that because scientists have been mistaken and deceived in the past about evidence for various
aspects of evolution the current evidence for evolution is likely to also be based on fraud and error. Much of
the evidence for evolution has been accused of being fraudulent at various times, including Archaeopteryx,
peppered moth melanism, and Darwin's finches; these claims have been subsequently refuted.[103][104]
[105]
It has also been claimed that certain former pieces of evidence for evolution which are now
considered out-of-date and erroneous, such as Ernst Haeckel's 19th-century comparative drawings of
embryos, used to illustrate his Recapitulation theory ("Ontogeny recapitulates Phylogeny"), were not merely
errors but frauds.[106] Jonathan Wells criticizes biology textbooks by alleging that they continue to reproduce
such evidence after it has been debunked.[104] In response, the National Center for Science Education
notes that none of the textbooks reviewed by Wells makes the claimed error, as Haeckel's drawings are
shown in a historical context with discussion about why they are wrong, and the accurate modern drawings
and photos used in the textbooks are misrepresented by Wells.[107]

[edit] Unreliable or inconsistent evidence

Creationists claim that evolution relies on certain types of evidence that do not give reliable
information about the past. It is argued, for example, that radiometric dating, the technique of evaluating a
material's age based on the radioactive decay rates of certain isotopes, generates inconsistent, and thus
unreliable, results. Radiocarbon dating, based on the Carbon 14 isotope, has been particularly criticized. It is
argued that radiometric decay relies on a number of unwarranted assumptions, such as the principle of
uniformitarianism, consistent decay rates, or rocks acting as closed systems. Such arguments have been
dismissed by scientists on the grounds that independent methods have confirmed the reliability of radiometric
dating as a whole; additionally, different radiometric dating methods and techniques have independently
confirmed each other's results.[108]
Another form of this objection is that fossil evidence is not reliable. This is based on a much wider
range of claims. These include that there are too many "gaps" in the fossil record,[109][110] that fossil-dating
is circular (see evolution is unfalsifiable), or that certain fossils, such as polystrate fossils, are seemingly "out
of place". Examination by geologists have found polystrate fossils to be consistent with in situ formation.[111]
It is argued that certain features of evolution support creationism's catastrophism (cf. Great Flood), rather
than evolution's gradualistic punctuated equilibrium,[112] which some assert is an "ad-hoc" theory to explain
the fossil gaps.[113]
[edit] Plausibility
Some of the oldest and most common objections to evolution dispute whether evolution can truly
account for all the apparent complexity and order in the natural world. It is argued that evolution is too
unlikely or otherwise lacking to account for various aspects of life, and therefore that an intelligence, God,
must at the very least be appealed to for those specific features.

[edit] Improbability
Further information: Teleological argument, Watchmaker analogy, Evolutionary argument against
naturalism
 Because the theory of evolution is often thought of as the idea that life arose "by chance", design
arguments such as William Paley's watchmaker analogy have been popular objections to the theory since
Darwin's day.[114]
A common objection to evolution is that it is simply too unlikely for life, in its complexity and apparent
"design", to have arisen "by chance". It is argued that the odds of life having arisen without a deliberate
intelligence guiding it are so astronomically low that it is unreasonable not to infer an intelligent designer from
the natural world, and specifically from the diversity of life.[115] A more extreme version of this argument is
that evolution cannot create complex structures. The idea that it is simply too implausible for life to have
evolved is often encapsulated with a quotation that the "probability of life originating on earth is no greater
than the chance that a hurricane sweeping through a scrap-yard would have the luck to assemble a Boeing
747" (a claim attributed to astrophysicist Fred Hoyle and known as Hoyle's fallacy).[116]
This view is thus invariably justified with arguments from analogy. The basic idea of this argument for
a designer is the teleological argument, an argument for the existence of God based on the perceived order
or purposefulness of the universe. A common way of using this as an objection to evolution is by appealing to
the 18th-century philosopher William Paley's watchmaker analogy, which argues that certain natural
phenomena are analogical to a watch (in that they are ordered, or complex, or purposeful), which means that,
like a watch, they must have been designed by a "watchmaker"—an intelligent agent. This argument forms the
core of intelligent design, a neocreationist movement seeking to establish certain variants of the design
argument as legitimate science, rather than as philosophy or theology, and have them be taught alongside
evolution.[28][47]
This objection is fundamentally an argument by lack of imagination, or argument from incredulity: a
certain explanation is seen as being counter-intuitive, and therefore an alternate, more intuitive explanation is
appealed to instead. Supporters of evolution generally respond by arguing that evolution is not based on
"chance", but on predictable chemical interactions: natural processes, rather than supernatural beings, are
the "designer". Although the process involves some random elements, it is the non-random selection of
survival-enhancing genes that drives evolution along an ordered trajectory. The fact that the results are
ordered and seem "designed" is no more evidence for a supernatural intelligence than the appearance of
complex natural phenomenon (e.g. snowflakes).[117] It is also argued that there is insufficient evidence to
make statements about the plausibility or implausibility of abiogenesis, that certain structures demonstrate
poor design, and that the implausibility of life evolving exactly as it did is no more evidence for an intelligence
than the implausibility of a deck of cards being shuffled and dealt in a certain random order.[47][114]
 It has also been noted that arguments against some form of life arising "by chance" are really
objections to nontheistic abiogenesis, not to evolution. Indeed, arguments against "evolution" are based on
the misconception that abiogenesis is a component of, or necessary precursor to, evolution. Similar
objections sometimes conflate the Big Bang with evolution.[12]
Christian apologist and philosopher Alvin Plantinga, a supporter of intelligent design, has formalized
and revised the improbability argument as the evolutionary argument against naturalism, which asserts that it
is irrational to reject a supernatural, intelligent creator because the apparent probability of certain faculties
evolving is so low. Specifically, Plantinga claims that evolution cannot account for the rise of reliable
reasoning faculties. Plantinga argues that whereas a God would be expected to create beings with reliable
reasoning faculties, evolution would be just as likely to lead to unreliable ones, meaning that if evolution is
true, it is irrational to trust whatever reasoning one relies on to conclude that it is true.[118] This novel
epistemological argument has been criticized similarly to other probabilistic design arguments. It has also
been argued that rationality, if conducive to survival, is more likely to be selected for than irrationality, making
the natural development of reliable cognitive faculties more likely than unreliable ones.[119][120]

[edit] Unexplained aspects of the natural world

It is frequently argued that a great weakness of evolutionary theory is that it does not, or cannot,
explain a certain aspect of the natural world. Although there is broad agreement that certain aspects of life
remain unexplained, creationists argue that evolution should be abandoned altogether because of the
phenomena it does not explain. It is argued that an alternative explanation, such as intelligent design, can
explain the things which evolution cannot. For example, Michael Behe has argued that current evolutionary
theory cannot account for certain complex structures, particularly in microbiology. On this basis, Behe argues
that such structures were "purposely arranged by an intelligent agent" (see argument from incredulity).[121]
 In addition to complex structures and systems, among the phenomena that critics variously claim
evolution cannot explain are consciousness, free will, hominid intelligence, instincts, emotions,
metamorphosis, photosynthesis, homosexuality, music, language, religion, morality, and altruism (see
altruism in animals).[122] Most of these, such as homosexuality, hominid intelligence, instinct, emotion,
photosynthesis, language, and altruism, have been well-explained by evolution, while others remain
mysterious, or only have preliminary explanations. However, supporters of evolution contend that no
alternative explanation has been able to adequately explain the biological origin of these phenomena either.
[123][124]
Creationists argue against evolution on the grounds that it cannot explain certain non-evolutionary
processes, such as abiogenesis, the Big Bang, or the meaning of life. In such instances, evolution is being
redefined to refer to the entire history of the universe, and it is argued that if one aspect of the universe is
seemingly inexplicable, the entire body of scientific theories must be baseless. At this point, objections leave
the arena of evolutionary biology and become general scientific or philosophical disputes.[125]

[edit] Implausibility
This class of objections is more radical than the above, claiming that a major aspect of evolution is
not merely unscientific or implausible, but rather impossible, because it contradicts some other law of nature
or is constrained in such a way that it cannot produce the biological diversity of the world.

[edit] Creation of complex structures

Further information: Irreducible complexity
 The bacterial flagellum is often invoked to illustrate the concept of irreducible complexity. Careful
analysis shows that there are no major obstacles to a gradual evolution of flagella.
"Living things have fantastically intricate features-at the anatomical, cellular, and molecular
levels-that could not function if they were any less complicated or sophisticated. The only
prudent conclusion is that they are products of Intelligent Design, not Evolution" - from Refuting
Evolution 2, Jonathan Sarfati[126]

Modern evolutionary theory posits that all biological systems must have evolved incrementally,
through a combination of natural selection and genetic drift. Both Darwin and his early detractors recognized
the potential problems that could arise for his theory of natural selection if the lineage of organs and other
biological features could not be accounted for by merely gradual, step-by-step changes over successive
generations; if all the intermediary stages between an initial organ and the organ it will become are not all
improvements upon the original, it will be impossible for the later organ to develop by the process of natural
selection alone. Anticipating early criticisms that the evolution of the eye and other complex organs seemed
impossible, Darwin noted that:
[R]eason tells me, that if numerous gradations from a perfect and complex eye to one very
imperfect and simple, each grade being useful to its possessor, can be shown to exist; if further,
the eye does vary ever so slightly, and the variations be inherited, which is certainly the case;
and if any variation or modification in the organ be ever useful to an animal under changing
conditions of life, then the difficulty of believing that a perfect and complex eye could be formed
by natural selection, though insuperable by our imagination, can hardly be considered real.[92]

Similarly, Richard Dawkins said on the topic of the evolution of the feather in an interview for the
television program The Atheism Tapes:
There's got to be a series of advantages all the way in the feather. If you can't think of one, then
that's your problem not natural selection's problem... It's perfectly possible feathers began as
fluffy extensions of reptilian scales to act as insulators... The earliest feathers might have been a
different approach to hairiness among reptiles keeping warm.

Creationist arguments have been made such as "What use is half an eye?" and "What use is half a
wing?".[127] Research has confirmed that the natural evolution of the eye and other intricate organs is
entirely feasible.[128][129] Creationist claims have persisted that such complexity evolving without a
designer is inconceivable, however, and this objection to evolution has been refined in recent years as the
more sophisticated irreducible complexity argument of the intelligent design movement, formulated by
biochemist Michael Behe.[28]
Irreducible complexity is the idea that certain biological systems cannot be broken down into their
constituent parts and remain functional, and therefore that they could not have evolved naturally from less
complex or complete systems. Whereas past arguments of this nature generally relied on macroscopic
organs, Behe's primary examples of irreducible complexity has been cellular and biochemical in nature. He
has argued that the components of systems such as the blood clotting cascade, the immune system, and the
bacterial flagellum are so complex and interdependent that they could not have evolved from simpler
systems.[130]
"In fact, my argument for intelligent design is open to direct experimental rebuttal. Here is a
thought experiment that makes the point clear. In Darwin’s Black Box (Behe 1996) I claimed that
the bacterial flagellum was irreducibly complex and so required deliberate intelligent design.
The flip side of this claim is that the flagellum can’t be produced by natural selection acting on
random mutation, or any other unintelligent process. To falsify such a claim, a scientist could go
into the laboratory, place a bacterial species lacking a flagellum under some selective pressure
(for mobility, say), grow it for ten thousand generations, and see if a flagellum—or any equally
complex system—was produced. If that happened, my claims would be neatly disproven." -
Michael Behe[131]

In the years since Behe proposed irreducible complexity, new developments and advances in
biology, such as an improved understanding of the evolution of flagella,[132] have already undermined these
arguments.[133][134] The idea that seemingly irreducibly complex systems cannot evolve has been refuted
through evolutionary mechanisms, such as exaptation (the adaptation of organs for entirely new functions)
[135] and the use of "scaffolding", which are initially necessary features of a system that later degenerate
when they are no longer required. Potential evolutionary pathways have been provided for all of the systems
Behe used as examples of irreducible complexity.[133][136][137]

[edit] Creation of information

Further information: Physical information, Biosemiotics
Another new, and increasingly common, objection of creationists to evolution is that evolutionary
mechanisms such as mutation cannot generate new information. Creationists such as William A. Dembski,
Werner Gitt, and Lee Spetner have attempted to use information theory to dispute evolution. Dembski has
argued that life demonstrates specified complexity, and that evolution without an intelligent agent cannot
account for the generation of information that would be required to produce specified complexity. The
Christian apologetics site Answers in Genesis, for example, makes frequent appeals to concepts from
information theory in its objections to evolution and affirmations of the Genesis account of Creation; "[I]t
should be clear that a rigorous application of the science of information is devastating to materialistic
philosophy in the guise of evolution, and strongly supportive of Genesis creation."[138]
However, these claims have been widely rejected by the scientific community; new information is
regularly generated in evolution, whenever a novel mutation or gene duplication arises. Dramatic examples
of entirely new, unique traits arising through mutation have been observed in recent years, such as the
evolution of nylon-eating bacteria, which developed new enzymes to efficiently digest a material that never
existed before the modern era.[139][140] In fact, when an organism is considered together with the
environment it evolved in, there is no need to account for the creation of information. The information in the
genome forms a record of how it was possible to survive in a particular environment. It is not created, but
rather gathered from the environment through research—by trial and error, as mutating organisms either
reproduce or fail.[141]
A related argument against evolution is that most mutations are harmful.[142] However, in reality the
vast majority of mutations are neutral, and the minority of mutations which are beneficial or harmful are often
situational; a mutation that is harmful in one environment may be helpful in another.[143]

[edit] Violation of the second law of thermodynamics

Further information: Entropy and life
Another objection is that evolution violates the second law of thermodynamics. Though the law
applies to all systems, in the case of a closed one it states, "the entropy of an isolated system not in
equilibrium will tend to increase over time, approaching a maximum value at equilibrium". In other words, an
ideal isolated system's entropy (a measure of the dispersal of energy in a physical system so that it is not
available to do mechanical work) will tend to increase or stay the same, not decrease. Creationists argue that
evolution violates this physical law by requiring a decrease in entropy, or disorder, over time.[144]
However, this claim is based on a manifestation of the law only applicable to isolated systems, which
do not exchange matter or energy with their surroundings. Organisms, in contrast, are open systems, as they
constantly exchange energy and matter with their environment: for example animals eat food and excrete
waste, and radiate and absorb heat. Similarly, the Earth absorbs energy from the Sun and emits energy back
into space. The Sun-Earth-space system does not violate the second law, because the enormous increase in
entropy due to the Sun and Earth radiating into space dwarfs the local decrease in entropy caused by the
existence and evolution of self-organizing life.[39][145]
 Since the second law of thermodynamics has a precise mathematical definition, this argument can
be analyzed quantitatively.[146][147] This was done by physicist Daniel Styer, who concluded: "Quantitative
estimates of the entropy involved in biological evolution demonstrate that there is no conflict between
evolution and the second law of thermodynamics."[146]
In a published letter to the editor of The Mathematical Intelligencer titled "How anti-evolutionists
abuse mathematics", mathematician Jason Rosenhouse stated:
The fact is that natural forces routinely lead to decreases in entropy. Water freezes into ice and
fertilised eggs turn into babies. Plants use sunlight to convert carbon dioxide and water into
sugar and oxygen, but [we do] not invoke divine intervention to explain the process [...]
thermodynamics offers nothing to dampen our confidence in Darwinism.[148]

[edit] Moral implications

Other common objections to evolution allege that evolution leads to objectionable results, including
bad beliefs, behaviors, and events. It is argued that the teaching of evolution degrades values, undermines
morals, and fosters irreligion or atheism. These may be considered appeals to consequences (a form of
logical fallacy), as the potential ramifications of belief in evolutionary theory have nothing to do with its
objective empirical reality.
[edit] Humans as animals

1871 caricature of Charles Darwin as an ape, accompanied by a satirical disclaimer claiming a fait
accompli for its inclusion[149]
 Huxley's book Man's Place in Nature was the first devoted to human evolution and an early example
of comparative biology.
Even though biology has long shown that humans are animals, some people feel that humans should
be considered separate from, and/or superior to, animals. The mediaeval concept of a great chain of being
set out a static hierarchy in which humans are "above" animals, but below angels and God. As evolutionary
thought developed it was feared that evolution implied that there was no real separation between man and
brute, and that it undermined human social hierarchy. Assertions of ape ancestry in the 1844 publication
Vestiges of Creation gained much public acceptance but were scornfully attacked by establishment
scientists, and though Darwin avoided the subject when announcing his theory in On the Origin of Species in
1859, the issue was quickly raised. Anatomists claimed that humans had unique physical features; however,
English biologist Thomas Henry Huxley showed that they were wrong and overcame their opposition to
Darwin's ideas. Evolutionary common descent does not imply that human beings should behave like other
animals, but instead shows that animals behave in different ways, and teaches that humans behave like
humans.[150]
Traditionalists still object to the idea that diversity in life, including human beings, arose through
natural processes without a need for supernatural intervention, and they argue against evolution on the basis
that it contradicts their literal interpretation of creation myths about separate "kinds". However, most religions,
such as Catholicism, have reconciled their beliefs with evolution through theistic evolution.[151][152]

[edit] Social effects

Further information: Social effect of evolutionary theory
 Creationists claim that perceived social ills like crime, teen pregnancies, homosexuality, abortion,
immorality, wars and genocide are caused by a belief in evolution.[153] R. Albert Mohler, Jr., President of
Southern Baptist Theological Seminary in Louisville, Kentucky, wrote August 8, 2005, in National Public
Radio's forum, "Taking Issue", that "Debates over education, abortion, environmentalism, homosexuality and
a host of other issues are really debates about the origin—and thus the meaning—of human life.... Evolutionary
theory stands at the base of moral relativism and the rejection of traditional morality".[154][155] Creationist
Ken Ham likens evolution to a horde of termites, weakening society's foundation. In Why Won't They Listen?,
Ham suggests that "evolutionary termites" are responsible for pornography, homosexual behavior and
lawlessness. He also writes, "I'm not saying that evolution is the cause of abortion or school violence. What
I'm saying is that the more a culture abandons God's word as the absolute authority, and the more a culture
accepts an evolutionary philosophy, then the way people think, and their attitudes, will also change."[156]
Former Texas Republican Representative Tom DeLay claimed that the Columbine school shootings
were caused by the teaching of evolution. DeLay is quoted as stating that "Our school systems teach the
children that they are nothing but glorified apes who are evolutionized [sic] out of some primordial soup."[157]
Henry M. Morris, engineering professor and founder of the Creation Research Society and the Institute of
Creation Research, claims that evolution was part of a pagan religion that emerged after the Tower of Babel,
was part of Plato's and Aristotle's philosophies, and was responsible for everything from war to pornography
to the breakup of the nuclear family.[158]
Rev. D. James Kennedy of The Center for Reclaiming America for Christ and Coral Ridge Ministries
claims that Darwin was responsible for Adolf Hitler's atrocities. In D. James Kennedy's documentary, and the
accompanying pamphlet with the same title, Darwin’s Deadly Legacy, Kennedy states that "To put it simply,
no Darwin, no Hitler." In his efforts to expose the "harmful effects that evolution is still having on our nation,
our children, and our world", Kennedy also states that, "We have had 150 years of the theory of Darwinian
evolution, and what has it brought us? Whether Darwin intended it or not, millions of deaths, the destruction
of those deemed inferior, the devaluing of human life, increasing hopelessness."[159][160][161] Discovery
Institute fellow Richard Weikart has made similar claims,[162][163] as have other creationists.[164] The claim
was central to Ben Stein's film Expelled: No Intelligence Allowed promoting intelligent design creationism.
The Anti-Defamation League describes such claims as outrageous misuse of the the Holocaust and its
imagery, and as trivializing "the many complex factors that led to the mass extermination of European Jewry.
Hitler did not need Darwin or evolution to devise his heinous plan to exterminate the Jewish people, and
Darwin and evolutionary theory cannot explain Hitler’s genocidal madness. Moreover, anti-Semitism existed
long before Darwin ever wrote a word."[161][165]
Kent Hovind of Creation Science Evangelism blames communism, socialism, World War I, World
War II, racism, the Holocaust, Stalin's war crimes, the Vietnam War and Pol Pot's Cambodian killing fields on
evolution, as well as the increase in crime, unwed mothers, and other social ills.[82] Kent Hovind's son Eric
Hovind has now taken over the family business while his father is in prison, and claims that evolution is
responsible for tattoos, body piercing, premarital sex, unwed births, sexually transmitted diseases (STDs),
divorce and child abuse.[166]
Supporters of evolution dismiss such criticisms as counterfactual, and some argue that the opposite
seems to be the case. A study published by the author and illustrator Gregory S. Paul found that religious
beliefs, including belief in creationism and disbelief in evolution, are positively correlated with social ills like
crime.[167] The Barna Group surveys find that Christians and non-Christians in the US have similar divorce
rates, and the highest divorce rates in the US are among Baptists and Pentecostals, both sects which reject
evolution and embrace creationism.[168]
Michael Shermer argued in Scientific American in October 2006 that evolution supports concepts like
family values, avoiding lies, fidelity, moral codes and the rule of law.[169] He goes on to suggest that
evolution gives more support to the notion of an omnipotent creator, rather than a tinkerer with limitations
based on a human model, the more common image subscribed to by creationists. Careful analyses of the
creationist charges that evolution has led to moral relativism and the Holocaust yields the conclusion that
these charges appear to be highly suspect.[170] Such analyses conclude that the origins of the Holocaust
are more likely to be found in historical Christian anti-semitism than in evolution.[171][172]

[edit] Atheism
Another charge leveled at evolutionary theory by creationists is that belief in evolution is either
tantamount to atheism, or conducive to atheism.[173] It is commonly claimed that all proponents of
evolutionary theory are "materialistic atheists".[174] On the other hand, Davis Young argues that Creation
Science itself is harmful to Christianity because its bad science will turn more away than it recruits. Young
asks, "Can we seriously expect non-Christians to develop a respect for Christianity if we insist on teaching
the brand of science that creationism brings with it?"[175] However, evolution neither requires nor rules out
the existence of a supernatural being. Philosopher Robert Pennock makes the comparison that evolution is
no more atheistic than plumbing.[176] H. Allen Orr, professor of biology at University of Rochester, notes
that:
Of the five founding fathers of twentieth-century evolutionary biology — Ronald Fisher, Sewall
Wright, J. B. S. Haldane, Ernst Mayr, and Theodosius Dobzhansky — one was a devout Anglican
who preached sermons and published articles in church magazines, one a practicing Unitarian,
one a dabbler in Eastern mysticism, one an apparent atheist, and one a member of the Russian
Orthodox Church and the author of a book on religion and science.[177]
 In addition, a wide range of religions have reconciled a belief in a supernatural being with evolution.
[178] Molleen Matsumura of the National Center for Science Education found that "of Americans in the twelve
largest Christian denominations, 89.6% belong to churches that support evolution education". These
churches include the United Methodist Church, National Baptist Convention USA, Evangelical Lutheran
Church in America, Presbyterian Church (USA), National Baptist Convention of America, African Methodist
Episcopal Church, the Roman Catholic Church, the Episcopal Church, and others.[179] A poll in 2000 done
for People for the American Way found that 70% of the American public felt that evolution was compatible
with a belief in God. Only 48% of the people polled could choose the correct definition of evolution from a list,
however.[180]
One poll reported in the journal Nature showed that among American scientists (across various
disciplines), about 40 percent believe in both evolution and an active deity (theistic evolution).[181] This is
similar to the results reported for surveys of the general American public. Also, about 40 percent of the
scientists polled believe in a God that answers prayers, and believe in immortality.[182] While about 55% of
scientists surveyed were atheists, agnostics, or nonreligious theists, atheism is far from universal among
scientists who support evolution, or among the general public that supports evolution. Very similar results
were reported from a 1997 Gallup survey of the American public and scientists.[183]
Belief in evolution
Belief in Young Belief in God-
Group[183] without God guiding the
Earth Creationism guided evolution
process

American
44% 39% 10%
public
 American
5% 40% 55%
scientists

[edit] See also

Book:Evolution

Books are collections of articles that can be downloaded or ordered in print.

• Faith and rationality

[edit] Notes
1. ^ a b Johnston, Ian C. (1999). "Section Three: The Origins of Evolutionary Theory". ... And
Still We Evolve. Liberal Studies Department, Malaspina University College.
http://records.viu.ca/~johnstoi/darwin/title.htm. Retrieved 2007-07-25.
2. ^ a b van Wyhe, John (2002-7). "Charles Darwin: gentleman naturalist: A biographical
sketch". The Complete Work of Charles Darwin Online. University of Cambridge. http://darwin-
online.org.uk/darwin.html. Retrieved 2007-07-25.
3. ^ IAP Statement on the Teaching of Evolution, Interacademy Panel
 4. ^ In his comprehensive treatise on Creationism, The Creationists, historian Ronald Numbers
traces the religious motivations and scientific pretensions, of prominent creationists from George
Frederick Wright through George McCready Price, Harry Rimmer, John C. Whitcomb, Henry M.
Morris and his Institute for Creation Research (and lesser figures) to Phillip E. Johnson and the
Intelligent design movement.
Numbers, Ronald (November 30, 2006). The Creationists: From Scientific Creationism to Intelligent
Design, Expanded Edition. Harvard University Press. p. 624 pages. ISBN 0674023390.
5. ^ Godfrey, Laurie R. Scientists Confront Creationism. Pg 8. W. W. Norton & Company
(1984). ISBN 0-393-30154-0.
6. ^ Sarfati, Jonathan & Matthews, Michael (2000). Argument: Creationism is religion, not
science. Answers in Genesis.
7. ^ "Statement on the Teaching of Evolution" (PDF). American Association for the
Advancement of Science. 2006.
http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf. Retrieved 2007-03-20.
8. ^ a b c Abdul Majid (2002). "The Muslim Responses To Evolution" (html). Islamic Research
Foundation International, Inc.. http://diberri.dyndns.org/cgi-bin/templatefiller/?type=&id=. Retrieved
2010-05-24.
9. ^ "Worldwide creationism, Shotgun stunner, and more" (html). New Scientist.
http://www.newscientist.com/article/mg20327153.900-worldwide-creationism-shotgun-stunner-and-
more.html. Retrieved 2010-05-24.
10.^ "About Us" (html). http://creation.com/about-us#who_we_are. Retrieved 2010-05-24.
11.^ "Opinions on evolution from ten countries | NCSE" (html). National Center for Science
Education. http://ncse.com/news/2009/07/opinions-evolution-from-ten-countries-004885. Retrieved
2010-06-14.
 12.^ a b c Moran, Laurence (1993). "What is Evolution?". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/evolution-definition.html. Retrieved 2007-03-24.
13.^ a b "Ask the experts:Biology-Is the human race evolving or devolving?". Scientific American.
1998. http://www.scientificamerican.com/article.cfm?id=is-the-human-race-evolvin. Retrieved 2007-
03-24.
14.^ Carroll SB (2001). "Chance and necessity: the evolution of morphological complexity and
diversity". Nature 409 (6823): 1102–9. doi:10.1038/35059227. PMID 11234024.
15.^ "CA212: Definition of evolution" (HTML). talkorigins.org.
http://www.talkorigins.org/indexcc/CA/CA212.html. Retrieved 2010-06-07.
16.^ Doolan, Robert (1996). "Oh! My aching wisdom teeth!". Answers in Genesis.
http://www.answersingenesis.org/creation/v18/i3/wisdom_teeth.asp. Retrieved 2007-03-24.
17.^ AboutDarwin.com
18.^ England, P.; Molnar, P.; Righter, F. (January 2007). "John Perry's neglected critique of
Kelvin's age for the Earth: A missed opportunity in geodynamics". GSA Today 17 (1): 4–9.
doi:10.1130/GSAT01701A.1.
19.^ Boltwood, B. B. (1907). "On the ultimate disintegration products of the radio-active
elements. Part II. The disintegration products of uranium". American Journal of Science 23: 77–88.
20.^ Bowler, Peter J. (1983). The eclipse of Darwinism: anti-Darwinian evolution theories in the
decades around 1900 (paperback ed.). Baltimore: Johns Hopkins University Press. pp. 23–24.
ISBN 0-8018-4391-X.
21.^ Bowler, PJ (2003). Evolution: The History of an Idea, Third Edition, Completely Revised
and Expanded. University of California Press. ISBN 978-0520236936.
 22.^ a b c Moore, James R. (1981). The Post-Darwinian Controversies: A Study of the Protestant
Struggle to Come to Terms with Darwin in Great Britain and America . Cambridge University Press.
p. 10. ISBN 9780521285179.
23.^ The Relations Between Religion and Science by Frederick Temple Lecture IV of Eight
Lectures Preached Before the University of Oxford in 1884
24.^ Pius XII, encyclical Humani Generis
25.^ John Paul II, Message to the Pontifical Academy of Sciences on Evolution
26.^ Adnan Oktar (1999). "The Evolution Deceit" (html).
http://www.harunyahya.com/evolution_specialpreface.php. Retrieved 2010-05-24.
27.^ "Darwin Correspondence Project – Darwin and design: historical essay".
http://www.darwinproject.ac.uk/content/view/110/104/. Retrieved 2008-09-02.
28.^ a b c Scott EC, Matzke NJ (May 2007). "Biological design in science classrooms". Proc.
Natl. Acad. Sci. U.S.A. 104 Suppl 1: 8669–76. doi:10.1073/pnas.0701505104. PMID 17494747.
PMC 1876445. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=17494747.
29.^ "Evolution is Religion". http://www.creationists.org/evolutionism-is-a-religion.html.
Retrieved 2010-05-01.
30.^ a b A copy of the Discovery Institutes Wedge Strategy document can be found here:
"Wedge Strategy" (PDF). Discovery Institute. 1999. http://www.antievolution.org/features/wedge.pdf.
Retrieved 2007-03-24.
31.^ "Polls Apart on Human Origins". Harris Interactive.
http://www.publicopinionpros.norc.org/features/2006/aug/bishop3.asp. Retrieved 2008-10-27.
32.^ a b "National Science Teachers Association Disappointed About Intelligent Design
Comments Made by President Bush". National Science Teachers Association Press. 2005.
http://www.nsta.org/about/pressroom.aspx?id=50794. Retrieved 2007-03-24.
 33.^ "A Scientific Dissent from Darwinism" (pdf). September 2001.
http://www.discovery.org/articleFiles/PDFs/100ScientistsAd.pdf. Retrieved 2007-10-30. ; original
"100 Scientists" advertisement.
34.^ Edwards, Mark (2001-09-24). "100 Scientists, National Poll Challenge Darwinism" (php).
Discovery Institute. http://www.reviewevolution.com/press/pressRelease_100Scientists.php.
Retrieved 2007-10-30.
35.^ Dembski, William (2001). "Is Intelligent Design Testable?".
http://www.arn.org/docs/dembski/wd_isidtestable.htm. Retrieved 2010-05-23.
36.^ Ham, Ken (2010). "Separation of Christianity and State".
http://www.answersingenesis.org/articles/au/separation-of-christianity-and-state. Retrieved 2010-05-
23.
37.^ Moran, Laurence (1993). "Evolution is a Fact and a Theory". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/evolution-fact.html. Retrieved 2007-03-24.
38.^ Dr. David N. Menton. "Is Evolution a Theory, a Fact, or a Law?" (html). Missouri
Association for Creation. http://mall.turnpike.net/C/cs/theory.htm. Retrieved 2010-06-16.
39.^ a b Isaak, Mark (2003). "Five Major Misconceptions about Evolution". The TalkOrigins
Archive. http://www.talkorigins.org/faqs/faq-misconceptions.html. Retrieved 2007-03-24.
40.^ Gould, SJ (1994). Hen's Teeth and Horse's Toes. W. W. Norton & Company. pp. 253–262.
ISBN 0393017168. http://www.stephenjaygould.org/library/gould_fact-and-theory.html. Retrieved
2007-03-24.
41.^ Lenski, RE (2000). "Evolution: Fact and Theory". ActionBioscience.org.
http://www.actionbioscience.org/evolution/lenski.html. Retrieved 2007-03-24.
42.^ a b c d Morris, HM (1985). Scientific Creationism. Master Books. ISBN 978-0890510025.
 43.^ a b c Theobald, Douglas (2004). "29+ Evidences for Macroevolution: Scientific "Proof",
scientific evidence, and the scientific method". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/comdesc/sciproof.html. Retrieved 2007-03-24.
44.^ The Crusade Against Evolution, Evan Ratliff, October 2004, Wired magazine
45.^ Meyer, SC (2002). "Teach the controversy". Cincinnati Enquirer.
http://www.arn.org/docs/meyer/sm_teachthecontroversy.htm. Retrieved 2007-03-24.
46.^ a b Isaak, M (2004). "Index to Creationist Claims, Claim CA040: Equal time". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CA/CA040.html. Retrieved 2007-03-24.
47.^ a b c d Scott, EC (2004). Evolution vs. Creationism: An Introduction. University of California
Press. ISBN 0520246500.
48.^ "Blown away by design". http://www.answersingenesis.org/creation/v21/i4/design.asp.
Retrieved 2010-05-01.
49.^ Morton, GR (2002). "The Imminent Demise of Evolution: The Longest Running Falsehood
in Creationism". http://home.entouch.net/dmd/moreandmore.htm. Retrieved 2007-03-24.
50.^ "World's Greatest Creation Scientists from Y1K to Y2K".
http://www.creationsafaris.com/wgcs_4.htm. Retrieved 2010-05-04.
51.^ "Was Darwin a Christian? Did he believe in God? Did he recant evolutionism when he
died? - ChristianAnswers.Net". http://www.christiananswers.net/q-aig/darwin.html. Retrieved 2010-
05-04.
52.^ a b Ham, K (1987). The Lie: Evolution. Master Books. ISBN 0-89051-158-6.
http://www.creationists.org/evolutionisreligion.html. Retrieved 2007-03-24. See Evolution is Religion,
Chapter 2.
 53.^ Dembski, WA (2006). The Design Inference: Eliminating Chance through Small
Probabilities (Cambridge Studies in Probability, Induction and Decision Theory) . Cambridge
University Press. ISBN 978-0521678674.
54.^ a b Morris, HM (2001). "Evolution Is Religion—Not Science" (PDF). Impact: Vital Articles on
Science/Creation 332. http://www.icr.org/pdf/imp/imp-332.pdf.
55.^ Wiker, BD (2003). "Does Science Point to God? Part II: The Christian Critics". Crisis
Magazine. http://www.crisismagazine.com/julaug2003/feature1.htm. Retrieved 2007-03-25.
56.^ Isaak, Mark (2004). "Index to Creationist Claims, Claim CA611: Evolution Sacrosanct?".
TalkOrigins Archive.
57.^ Kutschera U, Niklas KJ (June 2004). "The modern theory of biological evolution: an
expanded synthesis". Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y.
PMID 15241603.
58.^ McLean v Arkansas Board of Education, 529 F.Supp. 1255 .
59.^ Cline, A (2006). "Myth: Science is a Religion for Atheists that Requires Faith". about.com.
http://atheism.about.com/od/atheismscienceevolution/a/ScienceFaith.htm. Retrieved 2007-03-25.
60.^ a b c TalkOrigins Claim CA211, Mark Isaak, editor, Index to Creationist Claims, TalkOrigins,
2006. Retrieved on 2008-04-20.
61.^ Scientific Creationism, Henry M. Morris, 1974 Master Books, Arkansas, p. 6-7
62.^ Wilkins, JS (1997). "Evolution and Philosophy: Is Evolution Science, and What Does
'Science' Mean?". The TalkOrigins Archive. http://www.talkorigins.org/faqs/evolphil/falsify.html.
Retrieved 2007-03-25.
63.^ "Expelled Exposed: Why Expelled Flunks » Science & Religion". National Center for
Science Education. http://www.expelledexposed.com/index.php/the-truth/science-religion. Retrieved
2010-03-07.
 64.^ Popper, K (1985). Unended Quest: An Intellectual Autobiography. Open Court. ISBN 978-
0087583436.
65.^ a b Popper, K (1978). "Natural selection and the emergence of mind". Dialectica 32 (32):
339. doi:10.1111/j.1746-8361.1978.tb01321.x.
66.^ Misquoted Scientists Respond National Center for Science Education 1981 by John R.
Cole quoting Popper: "I have changed my mind about the testability and logical status of the theory of
natural selection, and I am glad to have the opportunity to make a recantation."
67.^ Darwin, Charles (1859). On the Origin of Species by Means of Natural Selection, or the
Preservation of Favoured Races in the Struggle for Life . p. 189.
68.^ Ridley, M (2003). Evolution, Third Edition. Blackwell Publishing Limited. ISBN 978-
1405103459.
69.^ Wallis, C (2005-08-07). "The Evolution Wars". Time Magazine. pp. 32.
http://www.time.com/time/archive/preview/0,10987,1090909,00.html. Retrieved 2007-03-24.
70.^ "Human Chromosome 2". 2002–2008 WGBH Educational Foundation.
http://www.teachersdomain.org/resource/evol07.sci.life.evo.genconnect/. Retrieved 2008-12-05.
71.^ Jeff Hecht (2003-05-19). "Chimps are human, gene study implies". NewScientist.
http://www.newscientist.com/article/dn3744-chimps-are-human-gene-study-implies.html. Retrieved
2008-05-10.
72.^ Jim Foley. "Fossil Hominids: The Evidence for Human Evolution".
http://www.talkorigins.org/faqs/homs/. Retrieved 2008-05-10.
73.^ Wilkins, JS (1997). "Evolution and Philosophy: A Good Tautology is Hard to Find". The
TalkOrigins Archive. http://www.talkorigins.org/faqs/evolphil/tautology.html. Retrieved 2007-03-24.
74.^ "Arguments we think creationists should NOT use". Answers in Genesis. 2006.
http://www.answersingenesis.org/Home/Area/faq/dont_use.asp. Retrieved 2007-03-24.
 75.^ See Survival of the fittest for a more thorough discussion.
76.^ MacRae, A (1998). "Radiometric dating and the geological time scale: Circular reasoning or
reliable tools". The TalkOrigins Archive. http://www.talkorigins.org/faqs/dating.html. Retrieved 2007-
03-24.
77.^ Hempel. C.G. 1951 “Problems and Changes in the Empiricist Criterion of Meaning” in
Aspects of Scientific Explanation. Glencoe: the Free Press. Quine, W.V.O 1952 “Two Dogmas of
Empiricism” reprinted in From a Logical Point of View. Cambridge: Harvard University Press
78.^ Kitcher, Philip (1982). Abusing science: the case against creationism . Cambridge, Mass:
MIT Press. pp. 45. ISBN 0-262-61037-X.
79.^ Kitcher 1982 p. 50
80.^ Kitcher 1982 p. 52
81.^ Kitcher 1982 p. 52-53
82.^ a b Kent Hovind. (2006). The Dangers of Evolution. [DVD]. USA: Creation Science
Evangelism.
83.^ a b Buckling A, Craig Maclean R, Brockhurst MA, Colegrave N (February 2009). "The
Beagle in a bottle". Nature 457 (7231): 824–9. doi:10.1038/nature07892. PMID 19212400.
84.^ Elena SF, Lenski RE (June 2003). "Evolution experiments with microorganisms: the
dynamics and genetic bases of adaptation". Nat. Rev. Genet. 4 (6): 457–69. doi:10.1038/nrg1088.
PMID 12776215.
85.^ Questions frequently asked about the TBSEF, Texans for Better Science Education
Foundation
86.^ Kansas Evolution Hearings Part 10, Kansas evolution hearings
87.^ Boxhorn, Joseph (1995). "Observed Instances of Speciation". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/faq-speciation.html#part5. Retrieved 2007-03-24.
 88.^ Wilkins, J (2006). "Macroevolution: Its Definition, Philosophy and History". The TalkOrigins
Archive. http://www.talkorigins.org/faqs/macroevolution.html. Retrieved 2007-03-24.
89.^ Mayr G, Pohl B, Peters DS (December 2005). "A well-preserved Archaeopteryx specimen
with theropod features". Science (journal) 310 (5753): 1483–6. doi:10.1126/science.1120331.
PMID 16322455.
90.^ Shubin NH, Daeschler EB, Jenkins FA (April 2006). "The pectoral fin of Tiktaalik roseae
and the origin of the tetrapod limb". Nature 440 (7085): 764–71. doi:10.1038/nature04637.
PMID 16598250.
91.^ Hunt (1997). "Transitional Vertebrate Fossils FAQ". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/faq-transitional.html. Retrieved 2007-03-24.
92.^ a b Darwin, C (1859). On the Origin of Species. John Murray. p. 280–313, 4th edition of
1866, p. 359–360
93.^ Elsberry, WR (1998). "Missing links still missing!". The TalkOrigins Archive.
http://www.talkorigins.org/origins/postmonth/feb98.html. Retrieved 2007-03-24.
94.^ "A faithful man takes on faith-based teaching". Answers in Genesis.
http://www.answersingenesis.org/docs2004/0806faithful.asp. Retrieved 2007-03-24.
95.^ Wieland, C (1991). "Variation, information and the created kind" ([dead link] – Scholar search).
Journal of Creation 5 (1): 42–47. http://creation.com/variation-information-and-the-created-kind.
Retrieved 2007-03-24.
96.^ Ham, Ken (1989). "Were You There?". Institute for Creation Research.
http://www.icr.org/index.php?module=articles&action=view&ID=670. Retrieved 2007-03-24.
97.^ Isaak, M (2005). "Index to Creationist Claims, Claim CA221: Were you there?". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CA/CA221.html. Retrieved 2007-03-24.
 98.^ Huelsenbeck JP, Rannala B (April 1997). "Phylogenetic methods come of age: testing
hypotheses in an evolutionary context". Science 276 (5310): 227–32.
doi:10.1126/science.276.5310.227. PMID 9092465.
99.^ Delsuc F, Brinkmann H, Philippe H (May 2005). "Phylogenomics and the reconstruction of
the tree of life". Nat. Rev. Genet. 6 (5): 361–75. doi:10.1038/nrg1603. PMID 15861208.
100.^ Einstein, Albert (1916). "The Foundation of the General Theory of Relativity" (PDF).
Annalen der Physik 49: 769–822. http://www.alberteinstein.info/gallery/gtext3.html. Retrieved 2006-
09-03.
101.^ Isaak, M (2004). "Index to Creationist Claims, Claim CA110: Evolution will soon be widely
rejected.". The TalkOrigins Archive. http://www.talkorigins.org/indexcc/CA/CA110.html. Retrieved
2007-03-24.
102.^ Richardson and Keuck, "Haeckel’s ABC of evolution and development," p. 516
103.^ Nedin, C (1997). "On Archaeopteryx, Astronomers, and Forgery". The TalkOrigins
Archive. http://www.talkorigins.org/faqs/archaeopteryx/forgery.html. Retrieved 2007-03-24.
104.^ a b Wells, J (2002). Icons of Evolution. Regnery Publishing, Inc.. ISBN 978-0895262004.
105.^ "Icons of Evolution FAQs". The TalkOrigins Archive. 2006.
http://www.talkorigins.org/faqs/wells/. Retrieved 2007-03-24.
106.^ "CB701: Haeckel's embryo pictures." (HTML). talkorigins.org.
http://www.talkorigins.org/indexcc/CB/CB701.html. Retrieved 2010-06-07.
107.^ "Icon 4 — Haeckel's Embryos". National Center for Science Education. November 23,
2006. http://ncse.com/creationism/analysis/icon-4-haeckels-embryos. Retrieved 2008-12-17.
108.^ Isaak, M (2004). "Index to Creationist Claims, Claim CD010: Radiometric Dating". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CD/CD010.html. Retrieved 2007-03-24.
 109.^ Isaak, M (2004). "Index to Creationist Claims, CC200: Transitional fossils". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CC/CC200.html. Retrieved 2008-07-13.
110.^ Isaak, M (2004). "Index to Creationist Claims, CC200.1: Transitional fossil abundance".
The TalkOrigins Archive. http://www.talkorigins.org/indexcc/CC/CC200_1.html. Retrieved 2008-07-
13.
111.^ Isaak, M (2004). "Index to Creationist Claims, CC340: Out-of-place fossils". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CC/CC340.html. Retrieved 2008-07-13.
112.^ Isaak, M (2004). "Index to Creationist Claims, Claim CC363: Requirements for
fossilization". The TalkOrigins Archive. http://www.talkorigins.org/indexcc/CC/CC363.html. Retrieved
2007-03-24.
113.^ Isaak, M (2004). "Index to Creationist Claims, CC201: Phyletic gradualism". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CC/CC201.html. Retrieved 2008-07-13.
114.^ a b Wilkins, J (1997). "Evolution and Chance". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/chance/chance.html. Retrieved 2007-03-24.
115.^ Battern, D (1995). "Cheating with chance". Answers in Genesis.
http://creation.com/cheating-with-chance. Retrieved 2009-12-06.
116.^ Lies, Damned Lies, Statistics, and Probability of Abiogenesis Calculations
117.^ "CI100: Intelligent Design". http://www.talkorigins.org/indexcc/CI/CI100.html.
118.^ Plantinga, A (1993). Warrant and Proper Function. Oxford University Press.
doi:10.1093/0195078640.001.0001. ISBN 0-19-507864-0.
119.^ Fitelson, B; Sober, E (1997). "Plantinga's Probability Arguments Against Evolutionary
Naturalism" (PDF). http://fitelson.org/plant.pdf. Retrieved 2007-03-24.
120.^ Isaak, M (2005). "Index to Creationist Claims, Claim CA120: Mind's fallibility". The
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CA/CA120.html. Retrieved 2007-03-24.
 121.^ Behe, MJ (1996-10-29). "Darwin under the microscope". New York Times. p. 25.
http://www.arn.org/docs/behe/mb_dm11496.htm. Retrieved 2007-03-24.
122.^ Johnson, P (1990). "Evolution as dogma: The establishment of naturalism".
http://www.arn.org/docs/johnson/pjdogma1.htm. Retrieved 2007-03-24.
123.^ "CB401: Inconceivable instinct". http://www.talkorigins.org/indexcc/CB/CB401.html.
Retrieved 2009-12-05.
124.^ "CB400: Evolution of consciousness". http://www.talkorigins.org/indexcc/CB/CB400.html.
Retrieved 2009-12-05.
125.^ Isaak, M (2004). "Index to Creationist Claims, Claim CE440: The origin of it all".
theTalkOrigins Archive. http://www.talkorigins.org/indexcc/CE/CE440.html. Retrieved 2007-03-24.
126.^ Jonathan Sarfati, Ph.D; 2003 Refuting Evolution 2. Master Books ISBN 0-89051-387-2
127.^ "CB921.2: Half a wing" (HTML). talkorigins.org.
http://www.talkorigins.org/indexcc/CB/CB921_2.html. Retrieved 2010-06-07.
128.^ Gehring, W.J. (2005). "New Perspectives on Eye Development and the Evolution of Eyes
and Photoreceptors". Journal of Heredity 96 (3): 171–184. doi:10.1093/jhered/esi027.
PMID 15653558.
129.^ "Eyes, Part One: Opening Up the Russian Doll. The Loom: A blog about life, past and
future".
http://www.corante.com/loom/archives/2005/02/15/eyes_part_one_opening_up_the_russian_doll.php
. Retrieved 2007-09-22.
130.^ Behe, MJ (1996). Darwin's Black Box: The Biochemical Challenge to Evolution. Free
Press. ISBN 978-0743290319.
 131.^ Behe, Michael (2000-07-31). "Philosophical Objections to Intelligent Design: Response to
Critics". http://www.arn.org/docs/behe/mb_philosophicalobjectionsresponse.htm. Retrieved 2010-01-
02.
132.^ Liu, R.Y.; Ochman, H. (2007). "Stepwise formation of the bacterial flagellar system".
PNAS 104: 7116-7121.
133.^ a b Isaak, M (2005). "Index to Creationist Claims, Claim CB200: Irreducible complexity".
The TalkOrigins Archive. http://www.talkorigins.org/indexcc/CB/CB200.html. Retrieved 2007-03-24. ;
and references therein
134.^ Ussery, D. (1999). "A biochemist's response to "The biochemical challenge to evolution"".
Bios 70: 40-45. http://www.cbs.dtu.dk/staff/dave/Behe.html.
135.^ Aharoni, A.; Gaidukov, L.; Khersonsky, O.; Gould, S.M.; Roodveldt, C; Tawfik, D.S.
(2005). "The 'evolvability' of promiscuous protein functions". Nature Genetics 37: 73-76.
136.^ Robison, K (1996). "Darwin's Black Box: Irreducible Complexity or Irreproducible
Irreducibility?". The TalkOrigins Archive. http://www.talkorigins.org/faqs/behe/review.html. Retrieved
2007-03-24.
137.^ Sanz, V.C. (2009). "Darwin's golden flame. Responses of biochemistry to intelligent
design". Teorema 28: 173-188.
138.^ Gitt, Werner (1996). Information, science and biology. Answers in Genesis.
139.^ Musgrave, I; Baldwin, R, et al. (2005). "Information Theory and Creationism". The
TalkOrigins Archive. http://www.talkorigins.org/faqs/information/infotheory.html. Retrieved 2007-03-
24.
140.^ Thomas, D. "Evolution and Information: The Nylon Bug". New Mexicans for Science and
Reason. http://www.nmsr.org/nylon.htm. Retrieved 2007-03-24.
 141.^ Bergstrom, CT; Lachmann, M (2006). "The fitness value of information".
http://arxiv.org/pdf/q-bio.PE/0510007. Retrieved 2007-03-24.
142.^ "CB101: Most mutations harmful?" (html). talkorigins.org.
http://www.talkorigins.org/indexcc/CB/CB101.html. Retrieved 2010-05-30.
143.^ Harter, R (1999). "Are Mutations Harmful?". The TalkOrigins Archive.
http://www.talkorigins.org/faqs/mutations.html. Retrieved 2007-03-24.
144.^ Lambert, F (2002). "Disorder — A Cracked Crutch For Supporting Entropy Discussions".
Journal of Chemical Education 79: 187–192. doi:10.1021/ed079p187.
http://www.entropysite.com/cracked_crutch.html. Retrieved 2007-03-24.
145.^ Oerter, RN (2006). "Does Life On Earth Violate the Second Law of Thermodynamics?".
http://physics.gmu.edu/~roerter/EvolutionEntropy.htm. Retrieved 2007-03-24.
146.^ a b Daniel F. Styer, "Entropy and evolution," American Journal of Physics, Vol. 76, No. 11,
November 2008, p. 1031
147.^ Emory F. Bunn, "Evolution and the Second Lqw of Thermodynamics," American Journal
of Physics, Vol. 77, No. 10, October 2009, p. 922
148.^ Rosenhouse, J (2001). "How Anti-Evolutionists Abuse Mathematics" (PDF). The
Mathematical Intelligencer 23 (4): 3–8. http://www.math.jmu.edu/~rosenhjd/sewell.pdf. Retrieved
2007-03-26.
149.^ A venerable Orang-utang "I have to apologize once more for the wild flights of my
incorrigible artist. I told him most clearly and positively to draw me a life-like portrait of that profound
philosopher, Mr. Darwin..." – The Hornet, 1871, from the collection of Darwin Online
150.^ Isaak, M (2004). "Claim CA009: Evolution teaches that people are animals. We should not
be surprised when people who are taught evolution start behaving like animals". The TalkOrigins
Archive. http://www.talkorigins.org/indexcc/CA/CA009.html. Retrieved 2007-03-24.
 151.^ Churches urged to challenge Intelligent Design
152.^ Pope John Paul II, "Truth Cannot Contradict Truth", New Advent, ed. Kevin Knight, 2009-
Feb-15 <http://www.newadvent.org/library/docs_jp02tc.htm>
153.^ Morris, H (1982). The Troubled Waters of Evolution. Master Books. ISBN 978-
0890510872.
154.^ Mohler, RA (2005). "The Origins of Life: An Evangelical Baptist View". NPR.
http://www.npr.org/templates/story/story.php?storyId=4760816. Retrieved 2007-03-24.
155.^ "The Result of Believing Evolution". Living Word Bible Church.
http://www.lwbc.co.uk/Genesis/results%20of%20believing%20evolution.htm. Retrieved 2007-03-24.
156.^ Ham, K (2002). Why Won't They Listen? A Radical New Approach to Evangelism. Master
Books. ISBN 0890513783. http://www.answersingenesis.org/home/area/WWTL/index.asp.
157.^ Raymo, C (1999-09-06). "Darwin's Dangerous De-evolution". Boston Globe.
http://www.boston.com/globe/search/stories/reprints/darwin100199.htm. Retrieved 2007-03-24.
158.^ Morris, HM (1989). The Long War Against God: The History and Impact of the
Creation/Evolution Conflict. Baker Book House. ISBN 0-89051-291-4.
159.^ "Kennedy: Evolution to Blame for Death, Hopelessness in World". Right Wing Watch.
http://www.rightwingwatch.org/2006/08/kennedy_evoluti.html. Retrieved 2007-03-24.
160.^ Martin, A; Parker, J (2006). "TV Producer Defends Documentary Exposing Darwin-Hitler
Link". Agape Press. http://headlines.agapepress.org/archive/8/252006b.asp. Retrieved 2007-03-24.
161.^ a b "ADL Blasts Christian Supremacist TV Special & Book Blaming Darwin For Hitler".
Anti-Defamation League. August 22, 2006. http://www.adl.org/PresRele/HolNa_52/4877_52.htm.
Retrieved 2009-12-07.
162.^ Weikart, R (2004). From Darwin to Hitler: Evolutionary Ethics, Eugenics, and Racism in
Germany. Palgrave Macmillan. ISBN 978-1403972019.
 163.^ From Darwin to Hitler: A Pathway to Horror (Updated) , Jonathan Witt, Evolution News and
Views, Discovery Institute, December 15, 2006.
164.^ This creationist claim that is part of a Discovery Institute campaign (New book by
Discovery Institute Fellow shows influence of Darwinian principles on Hitler's Nazi regime , Discovery
Institute) and is amply repeated in creationist literature. For example:
• Darwinism and the Nazi race Holocaust, Jerry Bergman, CEN Technical Journal,
13(2):101–111, 1999.
• The Holocaust and evolution, Jonathan Sarfati, Creation 22(1):4, December 1999.
• From Darwin to Hitler: Evolutionary Ethics, Eugenics, and Racism in Germany ,
Richard Weikart, Palgrave MacMillan, 2004.
165.^ "Anti-Evolution Film Misappropriates the Holocaust". Anti-Defamation League. April 29,
2008. http://www.adl.org/PresRele/HolNa_52/5277_52.htm. Retrieved 2009-12-07.
"Intelligent Design: It's Not Science – Religious Freedom Resources". Anti-Defamation League. 2009.
http://www.adl.org/religious_freedom/resource_kit/intelligent_design.asp. Retrieved 2009-12-07.
166.^ Creationist Links Origins to Faith, Everyday Life: Says outlook on Genesis account affects
every aspect of life , Bob Ellis, Dakota Voice, 5/7/2006
167.^ Paul, GS (2005). "Cross-National Correlations of Quantifiable Societal Health with Popular
Religiosity and Secularism in the Prosperous Democracies: A First Look". Journal of Religion &
Society 7. http://moses.creighton.edu/JRS/2005/2005-11.html. Retrieved 2007-03-24.
The paper was criticized by Moreno-Riaño, Smith, and Mach in a published article in the same
journal because "[Paul's] methodological problems do not allow for any conclusive statement to be
advanced regarding the various hypotheses Paul seeks to demonstrate or falsify." Of course,
correlation does not imply causality, and Paul does not produce any speculations about the cause of
these correlations.
 168.^ "Born Again Christians Just As Likely to Divorce As Are Non-Christians". The Barna
Group. 2004. http://www.barna.org/barna-update/article/5-barna-update/194-born-again-christians-
just-as-likely-to-divorce-as-are-non-christians. Retrieved 2004-03-24.
169.^ Shermer, M (2006). "Darwin on the Right: Why Christians and conservatives should
accept evolution". Scientific American. http://www.sciam.com/article.cfm?
chanID=sa006&articleID=00068F43-E189-150E-A18983414B7F0000&colID=13. Retrieved 2007-04-
26.
170.^ Darwin and Hitler: a not-very-intelligent link, Michael Ruse, My View, Tallahassee
Democrat, February 6, 2008
171.^ Talkorigins Claim CA006.1: Adolf Hitler exploited the racist ideas of Darwinism to justify
genocide, Mark Isaak, Index to Creationist Claims, Talkorigins, created 2001-4-29, modified 2005-7-
1, © 2006
172.^ Creationists for Genocide, Hector Avalos, Talkreason
173.^ Strobel, Lee (2004). The Case for a Creator. Zondervan. p. 32. ISBN 0310241448. "In my
quest to determine if contemporary science points toward or away from God, I knew I had to first
examine the claims of evolution in order to conclude once and for all whether Darwinism creates a
reasonable foundation for atheism. That's because if the materialism of Darwinian evolution is a fact,
then the atheist conclusions I reached as a student might still be valid."
174.^ Johnson, Phillip (1999). "The Church of Darwin". Wall Street Journal.
http://www.arn.org/docs/johnson/chofdarwin.htm. Retrieved 2010-05-23.
175.^ Young, D (1988). Christianity and the Age of the Earth. Artisan Publishers.
ISBN 093466627X.
176.^ Pennock, RT (2000). Tower of Babel: The Evidence Against the New Creationism. MIT
Press. ISBN 978-0262661652. http://ncse.com/store/title/tower-babel. Retrieved 2007-03-24.
 177.^ Devolution: Why intelligent design isn’t, H. Allen Orr, Annals of Science, The New Yorker,
May 30, 2005.
178.^ "Statements from Religious Organizations". National Center for Science Education. 2002.
http://ncse.com/media/voices/religion. Retrieved 2007-03-24.
179.^ Schrock, JR (2005-05-17). "Christianity, Evolution Not in Conflict" (PDF). Wichita Eagle.
pp. 17A. http://www.emporia.edu/biosci/schrock/docs/Eagle-25.pdf. Retrieved 2007-03-24.
180.^ "Evolution and Creationism In Public Education: An In-depth Reading Of Public Opinion"
(PDF). People for the American Way. 2002. http://docs.google.com/viewer?
a=v&q=cache:UDwdl6q3IgwJ:67.192.238.59/multimedia/pdf/Reports/evolutionandcreationisminpubli
ceducation.pdf+evolutionandcreationisminpubliceducation.pdf&hl=en&gl=ca&pid=bl&srcid=ADGEES
j-27vCwtszKiottg8mjbOKovsUvtJqRXXlLit-
R3VitD5tlX4OfMPkfEEzNwQjvUSioQSua6y3LN1SR72877bW3dc6wFzFlG2pFdWl-
21CVU68FWs00NDtWslzvTBvY-TLInH7&sig=AHIEtbQjqyUehD_pbin4wiuZlHKkn--pTA. Retrieved
2007-03-24.
181.^ Larson, EJ; Witham, L (1997). "Scientists are still keeping the faith". Nature 386: 435–436.
doi:10.1038/386435a0.
182.^ Witham, L (1997). "Many scientists see God's hand in evolution". Reports of the National
Center for Science Education 17 (6): 33. http://ncse.com/rncse/17/6/many-scientists-see-gods-hand-
evolution. Retrieved 2007-03-24.
183.^ a b Robinson, BA (1995). "Public beliefs about evolution and creation".
http://www.religioustolerance.org/ev_publi.htm. Retrieved 2007-03-24.
[edit] Further reading
• Philip Kitcher 1982 Abusing Science: The Case Against Creationism. Cambridge: The MIT
Press
• Jonathan Sarfati, Ph.D; 2003 Refuting Evolution 2. Master Books ISBN 0-89051-387-2

Look up evolution in Wiktionary, the free dictionary.

• Coleman, Simon; Leslie Carlin (2004). The Cultures of Creationism: Antievolution in English-
speaking Countries. Burlington, Vermont: Ashgate. p. 195. ISBN 075460912X.
• Isaak, Mark (2005). The Counter-Creationism Handbook. Westport, Connecticut: Greenwood
Press. p. 330. ISBN 031333305X.
• Rennie J (July 2002). "15 answers to creationist nonsense" (PDF). Sci. Am. 287 (1): 78–85.
doi:10.1038/scientificamerican0702-78. PMID 12085506.
http://www.swarthmore.edu/NatSci/cpurrin1/textbookdisclaimers/wackononsense.pdf.

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 W000

Historique des critiques des théories de

l'évolution
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En tant que théorie scientifique, la théorie darwinienne de l'évolution fait l'objet de diverses critiques.
Certains milieux religieux rejettent l'idée d'évolution biologique car elle s'oppose d'après eux à certains textes
sacrés, comme la Bible, et lui préfèrent le créationnisme. D'autres acceptent l'idée d'évolution mais refusent
l'idée, inspirée de Darwin, que les mécanismes principaux en soient les mutations et la sélection naturelle,
théorie désignée sous le nom de « darwinisme », et avancent que l'évolution serait pour certains dirigée
(orthogenèse), ou pour d'autres orientée vers une complexification croissante notamment par les contraintes
et facteurs internes des organismes (conformément aux vues de Jean-Baptiste Lamarck sur la question).
D'autres auteurs encore critiquent les conséquences idéologiques d'une « utilisation » (ou d'un
détournement) de la théorie darwinienne de l'évolution, qui n'est selon eux, qu'une projection sur le monde
naturel du fonctionnement de l'économie capitaliste, servant en retour à présenter celle-ci comme naturelle.
La majorité de la communauté scientifique considère cependant ces critiques comme injustifiées et
le créationnisme et l'évolution dirigée comme des idées obsolètes, qui continuent d'être soutenues pour des
raisons religieuses.
Sommaire
[masquer]
• 1 Origines profondes du débat
• 2 Critiques à caractère idéologique
• 2.1 Karl Marx et Friedrich Engels
• 2.2 Friedrich Nietzsche
• 2.3 Autres auteurs
• 3 Critiques à caractère scientifique
• 3.1 Les chainons manquants
• 3.2 Néo-catastrophisme
• 3.3 La complexité irréductible
• 3.4 Le second principe de la thermodynamique
• 4 Critiques à caractère religieux
• 4.1 Créationnisme
• 4.2 Le Dessein Intelligent
• 4.3 Articles connexes
• 4.4 Liens externes
• 4.5 Ouvrages
• 5 Notes et références
Origines profondes du débat [modifier]
Ces débats ont des racines antiques[1] : La cosmogonie mythologique des anciens Grecs fait surgir
l'ordre du chaos primordial. Cette énigme est à l'origine de nombre de supputations métaphysiques. Les
débats engagés chez les pré-socratiques se sont poursuivis durant vingt siècles donnant lieu à des courants
de pensées opposés : monisme, dualisme, matérialisme, spiritualisme, réalisme, idéalisme, mécanisme,
vitalisme...
Les monistes voient l'univers formé d'une seule réalité fondamentale (les milésiens et les atomistes
grecs : Démocrite, Épicure), alors que les dualistes séparent le monde matériel et le monde spirituel, l'au-
delà (Platon[2], Aristote[3]).
Issu du monisme, le matérialisme considère qu'il n'y a pas d'autre substance que la matière. La
pensée, la conscience dérivent par transformation de la matière. Le matérialisme rejette l'existence de tout
principe non réductible à une substance fondamentale (pas d'esprit transcendant la matière, d'âme, de dieux
et d'au-delà).
Au contraire pour les spiritualistes, l'univers possède une nature spirituelle, supérieure à la matière.
L'homme, par sa conscience, son esprit ou son âme, ne se réduit pas à la seule matière.
L'opposition idéalisme/réalisme, quant à elle, porte sur l'origine de la connaissance (celle du
spiritualisme et du matérialisme porte sur la nature de l'Être). Pour les tenants de l'idéalisme, il n'y a pas de
réalité indépendamment de la pensée. Le monde réel n'existe qu'à travers les idées et les états de
conscience. Le monde et même l'être se réduisent donc aux représentations que nous en avons. À l'inverse
pour un réaliste, l'homme connait les choses telles qu'elles sont réellement en elles-mêmes et elles
n'existent pas en dehors de leur matérialisation.
 Issus de ces débats, deux courants (et les tentatives de synthèses attenantes) ont traversé les
siècles. D'un côté une vision du monde « moniste et causale » où les phénomènes naturels sont produits par
le jeu des forces physico-chimiques immanentes à la matière. De l'autre, une conception « dualiste », où
l'explication mécanique est tenue pour valable dans le domaine inorganique, mais non valable pour le vivant.
Le matérialisme moniste explique tout par les causes efficientes. Au contraire, selon les non-
matérialistes, des principes différents opèrent selon qu'il s'agit du vivant ou de l'inanimé. Pour le vivant, la
doctrine dualistique introduit des causes finales et met à l'origine des organismes soit un plan de
développement, soit la décision d'un créateur qui a disposé chacune des espèces selon un plan : d'un côté
les mécanistes et de l'autre les vitalistes ; d'un côté les déterministes (les formes sont le résultat du jeu des
causes efficientes) et de l'autre les finalistes (il existe un but, une cause finale qui dirige l'évolution des
formes).
Au début du XXe siècle, les progrès des sciences physiques et biochimiques ainsi que la théorie
darwinienne produisent un monde où la frontière entre vivant et inanimé semble devoir disparaitre.
Différentes personnes considèrent l'explication mécaniste comme universellement valide mais restent
toutefois attachées à la présence d'une cause finale. Elles considèrent que les êtres vivants sont des totalités
(wholes) inanalysables qui ne s'expliquent pas par un assemblage de parties ; il y a quelque chose, selon
elles, qui ordonne ces parties et qui n'est pas de l'ordre de la causalité efficiente. Elles postulèrent l'existence
de formes, de types d'organisation qui tendent à se réaliser, de potentiels qui guident l'évolution vers un but
assigné, poursuivant en cela un débat engagé vingt-trois siècles auparavant par Aristote sur la forme,
organisatrice et conservatrice de l'être vivant (« forma est qua ens est id quod est »)
Pour synthétiser les enjeux philosophiques des théories de l'évolution on peut distinguer plusieurs
familles philosophiques :
 « L'objectivité cependant nous oblige à reconnaitre le caractère téléonomique des êtres vivants,
à admettre que dans leurs structures et performances, ils réalisent et poursuivent un projet. Il y
a donc là, au moins en apparence, une contradiction épistémologique profonde. Le problème
central de la biologie, c'est cette contradiction elle-même, qu'il s'agit de résoudre si elle n'est
qu'apparente, ou de prouver radicalement insoluble si en vérité il en est bien ainsi.[4] »

1) Les déterministes :
« La pierre angulaire de la méthode scientifique est le postulat de l'objectivité de la Nature.
C'est-à-dire le refus systématique de considérer comme pouvant conduire à une connaissance
"vraie" toute interprétation des phénomènes donnée en termes de causes finales, c'est-à-dire
de "projet". [...] Postulat pur, à jamais indémontrable, car il est évidemment impossible
d'imaginer une expérience qui pourrait prouver la non-existence d'un projet, d'un but poursuivi,
où que ce soit dans la nature. Mais le postulat d'objectivité est consubstantiel à la science, il a
guidé tout son prodigieux développement depuis trois siècles. Il est impossible de s'en défaire,
fût-ce provisoirement, ou dans un domaine limité, sans sortir de celui de la science elle-même.
[4] »

Cette position philosophique qui fonde la science fut d'une fécondité indiscutable. La rationalisation
qui s'en suivit, aboutit pour la première fois dans l'histoire à des guérisons innombrables, applications
pratiques des compréhensions fines et des découvertes biologiques lié à l'hérédité, l'organogenèse et les
mécanismes biomoléculaires de l'évolution. Ce tronc philosophique du déterminisme est l'origine de la
théorie synthétique de l'évolution (déterministe, mécanistes et réductionniste).
2) Les finalistes :
 • Parmi ceux-ci certains rejettent toutes idées d'évolution ( → créationnisme).
• D'autres intègrent les faits scientifiques accumulés en faveur de l'évolutionnisme mais
rajoutent une force organisatrice, qui conduit l'atome vers la conscience. Ici point de créationnisme,
l'évolution est acceptée et se déroule au sein d'un « champ directeur ». Ce principe « directionnel » a
porté entre autres dénominations : entéléchie, force vitale, principe directeur, holisme[5],
organicistes, orthogenèse, dessein intelligent, principe anthropique ou point Ω de Pierre Teilhard de
Chardin.

Note
On peut réfléchir que cette opposition en apparence insoluble provient (du moins en partie) d'un
manque de conceptualisation des catégories de phénomènes. En effet, un principe de causalité est ici
considéré comme « évident », alors que le concept de « projet », qui apparemment serait aussi suivi,
(présenté ici dans le monde biologique) ne puisse être directement analysé...
Pourtant, de la succession (causale) des interactions des divers systèmes physiques amènent à considérer
naturellement la notion de devenir, de « but », même si ce n'est pas directement final.
Exemples : dans le domaine du vivant, mais aussi dans les divers cycles physiques des interactions avec les
autres systèmes, comme les climats, les tectoniques planétaires, la vie des étoiles, l'évolution galactique et
cosmique, ect...
Ne serait-on pas là devant des notions complémentaires ?
Critiques à caractère idéologique [modifier]
Elles sont les premières à intervenir : Darwin a d'abord été jugé selon le contexte économique
politique et social de son temps.

Karl Marx et Friedrich Engels [modifier]

Karl Marx et Friedrich Engels ont remarqué de suite l'origine idéologique du mécanisme de la
sélection naturelle:
Lettre de Marx à Engels du 18 juin 1862
« Il est curieux de voir comment Darwin retrouve chez les bêtes et les végétaux sa société
anglaise avec la division du travail, la concurrence, l’ouverture de nouveaux marchés, les
« inventions » et la « lutte pour la vie » de Malthus. C’est le bellum omnium contre omnes [la guerre de
tous contre tous] de Hobbes, et cela fait penser à la phénoménologie de Hegel, où la société
bourgeoise figure sous le nom de « règne animal intellectuel », tandis que chez Darwin, c’est le règne
animal qui fait figure de société bourgeoise. »
Lettre de Engels à Lavrov du 12 [17] novembre 1875
« Toute doctrine darwiniste de la lutte pour la vie n’est que la transposition pure et simple, du
domaine social dans la nature vivante, de la doctrine de Hobbes : bellum omnium contre omnes et de
la thèse de la concurrence chère aux économistes bourgeois, associée à la théorie malthusienne de la
population. Après avoir réalisé ce tour de passe-passe […], on retranspose les mêmes théories cette
fois de la nature organique dans l’histoire humaine, en prétendant que l’on a fait la preuve de leur
 validité en tant que lois éternelles de la société humaine. Le caractère puéril de cette façon de
procéder saute aux yeux, il n’est pas besoin de perdre son temps à en parler. »
Le fait est que ni Marx ni Engels ne rendront publiques ces analyses. Au contraire, engagés eux-
mêmes dans des luttes idéologiques et politiques, ils verront dans la sélection naturelle, moteur de
l’évolution, la justification de la lutte des classes comme moteur de l’histoire sociale. Ce parallèle s'impose
comme la conception sociale de leur temps et de l'histoire de la société. L'attitude de Marx et Engels à
l'égard des idées de Darwin reste donc très ambigüe. Dans l’éloge funèbre qu’il lira sur sa tombe, Engels ira
même jusqu’à comparer Marx à Darwin en évoquant cette analogie.
Engels s'intéressait à l'histoire de la nature, ce dont témoignent ses ouvrages Dialectique sur la
nature et Le Rôle du travail dans le passage du singe à l'homme qui s'inspirent des travaux de Ernst
Haeckel[6]. Et, c'est lui qui fait découvrir le livre de Darwin à Marx. Ce dernier va d'ailleurs référencer
l'Origine des Espèces dans Le Capital et y note simplement l'analogie et la distinction entre « l'histoire de la
technologie naturelle » et « l'histoire de la formation des organes productifs de l'homme social ». Ainsi,
comme la « spéciation » observée sur les becs des pinsons des différentes iles des Galapagos, « La période
manufacturière simplifie, améliore et multiplie les outils de travail en les adaptant aux fonctions spécifiques
exclusives des travailleurs partiels.[7] »; « C'est ainsi, par exemple, que des couteaux destinés à couper
toutes sortes de choses peuvent, sans inconvénient, avoir une forme commune, tandis qu'un outil destiné à
un seul usage doit posséder pour tout autre usage une toute autre forme (Darwin, On the Origin of
Species..., 1859, p149).[8] » Marx reprend en cela l'idée de Darwin selon laquelle l'adaptation consiste en
une sorte de "technologie des organismes".
Marx fut encore plus enthousiaste de l'Origine et transformations de l'homme et des autres êtres de
Pierre Trémaux en voyant « un progrès par rapport à Darwin ». Mais, Engels lui annonce qu'il est arrivé « à
la Conclusion que sa théorie (celle de Trémaux) ne repose sur rien, pour la simple raison qu'il ne comprend
pas la géologie et qu'il est incapable de la plus ordinaire des critiques historico-littéraires.[9] »
Par la suite les marxistes, tels Paul Lafargue ou Anton Pannekoek, défendront la théorie darwinienne
de l'évolution pour des raisons essentiellement idéologiques, c'est-à-dire en tant qu'elle s'oppose à la
théologie naturelle, au créationnisme ou au dessein intelligent.

Friedrich Nietzsche [modifier]

Dans un fragment posthume intitulé « Contre le Darwinisme », le philosophe Friedrich Nietzsche
exposait dans les années 1880 une position fort intéressante : « L'utilité d'un organe n'en explique pas la
genèse, au contraire ! Pendant la plus longue partie du temps où une qualité se forme, l'individu n'en
bénéficie pas, elle ne lui sert pas, surtout dans la lutte contre les circonstances extérieures et ses ennemis. »
Mais il va encore plus loin en ajoutant : « L'influence des circonstances extérieures a été follement exagérée
par Darwin. L'essentiel du processus vital est justement cette force immense de formation, qui crée des
formes « du dedans », qui utilise, exploite les « circonstances extérieures. »
Ce point de vue semble se rapprocher du concept philosophique abondamment développé (et très
attaché aux sciences naturelles) de Volonté de puissance[10], qui constituerait ici une force motrice
endogène solidaire du vivant, et dont le milieu ne constituerait plus qu'un moyen. De manière très
schématique, on pourrait parler « d'exploitation active des possibilités offertes par l'environnement »,
tributaire chez l'animal du développement de nouveaux organes et chez l'Homme, capable de modeler son
milieu, de nouveaux instincts. Naturellement, si un tel modèle présente le double avantage d'être à la fois
simple et d'une commodité à toute épreuve, la science expérimentale ne peut que lui refuser la qualification
de découverte scientifique, et à plus forte raison s'il sert des motivations d'ordre idéologique.
 En effet, ce fragment est incompréhensible s'il est fait abstraction de son contexte théorique : au
fondement de l'œuvre du philosophe, parfois soupçonné d'eugénisme[11], réside cette idée que l'Homme est
une espèce en développement qui produit, de-ci de-là, des types supérieurs, caractérisés par la richesse
relative de leurs instincts (Platon, César, Napoléon, Goethe...). Helléniste réputé et aristocrate de
sensibilité[12], l'auteur fut témoin bien malgré lui des profonds changements structurels de la société du XIXe
siècle. Il a ainsi cherché à interpréter l'essor de l'art moderne et la domination des instincts bourgeois – jugés
pauvres – comme une réaction qui prouve la résistance absolue de l'environnement à l'évolution des
espèces[13], et donc également, la fausseté principielle du modèle de Darwin[14].
Ainsi, cette idée que « l'utilité d'un organe n'en explique pas la genèse » et que cette dernière
commence même par être handicapante, trouve son fondement véritable dans la situation de ceux que
l'auteur appelle « les hommes supérieurs ». En constatant chez eux l'apparition d'instincts plus riches
indépendamment de toute espèce de milieu d'une part, et en constatant la souffrance résultant de ces
instincts privés de conditions d'existence d'autre part, l'auteur a admis que l'évolution des espèces était un
phénomène absolument endogène et défavorisant. Son ouvrage central, Ainsi parlait Zarathoustra, est ainsi
un poème mystique destiné à favoriser le développement de l'homme supérieur[15] dans sa quête qui vise à
surmonter la souffrance de son milieu[16], pour aboutir à l'établissement de son milieu propre et à la
réalisation de la finalité dernière de l'évolution : le Surhumain[17].

Autres auteurs [modifier]

En 1910, le sociologue Jacques Novicow publie Le darwinisme social (éd. Alcan) qui contient une
critique du darwinisme au plan biologique et une critique de l'usage qui est fait du darwinisme dans la
sociologie.
 Plus récemment, l'historien des sciences André Pichot a consacré un chapitre entier de son Histoire
de la notion de vie (1993) à une analyse très critique des thèses de Darwin. Pichot est l'auteur de La société
pure : de Darwin à Hitler (2001) et de Aux origines des théories raciales, de la Bible à Darwin (2008), qui
retracent l'origine des idées eugénistes et racistes et montre en quoi le darwinisme, en tant que corpus soi-
disant scientifique, a servi à justifier certaines pratiques et idéologies. Pichot est ainsi un des rares critiques
du darwinisme à ne pas se cantonner au seul terrain scientifique, mais à réussir à articuler en un tout
cohérent la critique épistémologique des théories dominantes en biologie (hérédité, génétique, darwinisme)
avec la critique historique, politique et sociale des idéologies qui les ont en partie inspirées et qu'en retour
elles viennent conforter.
La notion d'idéologie scientifique a été avancée par Georges Canguilhem dans son ouvrage
Idéologie et rationalité dans l’histoire des sciences de la vie : Nouvelles études d’histoire et de philosophie
des sciences (éd. Vrin, 1977). Certains auteurs avancent l'idée que le darwinisme répondrait à cette
définition.

Critiques à caractère scientifique [modifier]

De la part de certains zoologistes, éthologistes, paléontologues tels que Rémy Chauvin, Pierre-Paul
Grassé ou Étienne Rabaud.
Pierre-Paul Grassé a présenté ses principaux arguments contre le darwinisme, sans pour autant
proposer une théorie nouvelle, dans son ouvrage L'évolution du vivant, matériaux pour une nouvelle théorie
transformiste (1973). Contre l'idée selon laquelle l'évolution des êtres vivants est le produit de la sélection
naturelle et des changements qui surviennent dans l'environnement, il met en avant les espèces
panchroniques, c'est-à-dire les espèces qui ont arrêté d'évoluer à un moment donné et qui sont restées à
peu près telles quelles jusqu'à nos jours malgré de grandes modifications géologiques, climatiques, etc. (il en
donne de nombreux exemples dans Les formes panchroniques et les arrêts de l'évolution , p. 133). Ainsi,
l'évolution est pour lui un processus qui n'est pas nécessaire, il ne s'effectue pas sous la contrainte des
forces physiques extérieures à l'être vivant (cf. La nécessité-utilité n'est pas le primus movens de l'évolution
biologique, p. 302). Pour l'expliquer, il pense qu'il faut donner la priorité à la dynamique interne propre aux
êtres vivants. À partir de là et de l'examen des archives fossiles, il en conclut que l'évolution est orientée (et
non dirigée comme l'avance l'orthogénèse, qu'il critique) vers un accroissement de la complexité des êtres
vivants. Ainsi, Pierre-Paul Grassé se situait sur le terrain du néo-lamarckisme.
Plus récemment, Gérard Nissim Amzallag a émis des critiques d'ordre épistémologique dans son
ouvrage La raison malmenée, critique des idées reçues en biologie moderne , et d'ordre scientifique dans son
deuxième ouvrage L'homme végétal, pour une autonomie du vivant, qui s'oppose, à l'aide de nombreux
exemples, à l'idée selon laquelle les êtres vivants sont comparables à des machines et met en avant la
notion d'autonomie du vivant.

Les chainons manquants [modifier]

La théorie de l'évolution suppose qu'entre les espèces vivantes, un grand nombre de formes
intermédiaires existèrent autrefois, le processus de l'évolution étant supposé très lent et progressif, ou
irrégulier selon les théories. On peut donc s'attendre à en trouver les restes sous forme de fossiles.
Les biologistes s'accordent sur le fait que les fossiles retrouvés représentent une part minuscule de ce qui a
vécu sur Terre. En effet, la fossilisation ne s'applique quasiment pas aux corps mous, elle n'intervient que
dans des conditions très spécifiques, puis les fossiles produits doivent résister à l'érosion. Ils doivent ensuite
être découverts.
De plus, s'il était effectivement difficile de trouver des fossiles montrant les étapes de l'évolution
conformément à la théorie à l'époque de Darwin, la biologie contemporaine dispose de nombreux exemples
de fossiles transitionnels par exemple dans la lignée humaine. Le domaine de ce qu'on appelle l'EvoDevo
(évolution et développement) fournit même des éléments précis sur les mécanismes d'apparitions de
nouveaux organes ou de nouvelles structures qui engendrent de nouvelles espèces[18].
Certains biologistes comme David Raup affirment que l'apparition de nouvelles espèces est trop
rapide pour laisser des traces géologiques. Ce dernier a déploré l'instrumentalisation de ses travaux par
certains mouvements créationnistes[19]. Or ce débat entre équilibre ponctué et gradualisme est interne à la
biologie de l'évolution ; il n'en remet pas du tout en cause les principes mais cherche seulement à en
caractériser la dynamique à l'échelle des temps géologiques.
Malgré cela, le manque de formes intermédiaires dans des domaines précis a été dès le début et
reste un argument employé par les détracteurs de l'évolution.
Article détaillé : Chaînon manquant (évolution).
Le problème du chainon manquant de la lignée humaine (un être qui serait intermédiaire entre le
singe et l'homme) a longtemps été employé contre la théorie de l'évolution. Depuis, de nombreux hominidés
ont été découverts, au point que le problème est aujourd'hui plus de déterminer lesquels sont des ancêtres
de l'homme et/ou du singe, et lesquels sont des lignées éteintes.

Néo-catastrophisme [modifier]
Le catastrophisme est une théorie scientifique qui a tenté de construire rationnellement les
croyances sur l'origine du monde et sur l'évolution des espèces en mettant en avant l'impact qu'auraient eu
des catastrophes de courte durée, violentes et inhabituelles. Cette théorie a été qualifiée ainsi au XIXe siècle,
lorsqu'est apparu l'uniformitarisme, théorie qui, quant à elle, postule que les processus qui se sont exercés
dans un passé lointain s'exercent encore de nos jours.
Par ailleurs, depuis le début du XXIe siècle le catastrophisme quitte le domaine des sciences
biologiques et fait explicitement l'objet de diverses théorisations tant sur le plan social, philosophique[20],
que politique[21].
• Argumentation : Selon la théorie de Luis Walter Alvarez et son fils, qui se veut expliquer
l'Extinction du Crétacé c'est l'impact d'une météorite géante qui serait à l'origine de cette extinction
massive[22].

La complexité irréductible [modifier]

Article détaillé : Complexité irréductible.
Plusieurs opposants à la théorie de l'évolution darwinienne, notamment Henri Bergson dans
L'évolution créatrice, arguent que certains organes, comme l'œil humain (ou celui du homard), exigent un
agencement très précis et concourant de différents éléments pour fonctionner correctement. Ils ne pourraient
donc être le résultat d'une évolution progressive par sélection naturelle : une ébauche d'œil ne fonctionnerait
pas et ne donnerait pas un avantage sélectif significatif. Arthur Koestler estime même, dans Janus, que cette
mutation inutile aurait de fortes chances de se diluer et de disparaitre bien avant que les suivantes n'arrivent
pour la compléter.
Bergson s'oppose à Darwin sur un point : la réductibilité du phénomène vivant à une explication
mécaniste, surtout pour des raisons épistémologiques liées à la fonction de notre intelligence ; celle-ci, dans
une perspective pragmatique, vise l'action et constitue elle-même un outil d'adaptation. Le modèle établi par
Darwin, selon lui conséquence de notre faculté de connaître, resterait encore insuffisant et exigerait une
explication philosophique et intuitive de l'évolution. L'existence même d'une évolution n'est jamais remise en
cause par Bergson ni par Koestler, mais tous deux insistent sur le fait que si son rôle d'élimination est
indiscutable, son apport explicatif à des successions d'innovations en cascade reste problématique même
sur les durées considérées, essentiellement pour des raisons de dilution de mutations qui, isolées,
resteraient sans effet.
En fait, un organe photosensible même élémentaire, capable par exemple de simplement distinguer
le jour de la nuit ou le passage d'une ombre, d'un prédateur ou d'un proie potentielle, procure un avantage
sélectif suffisant[23] pour qu'on puisse envisager qu'il s'impose rapidement. De là à obtenir un organe
beaucoup plus complexe tel que l'œil des vertébrés, le nombre d'étapes est certes considérable et demande
une coordination des modifications de plusieurs organes adjacents, mais chaque amélioration accentuera
l'avantage sélectif et s'imposera.
Des travaux montrent que l'apparition d'un œil complexe peut être en fait très rapide. En effet, Nilssion et
Pelger montrent en 1994 que, sous des hypothèses particulièrement pessimistes, 400 000 générations
suffisent à former un œil camérulaire (comme celui des Vertébrés) selon les modèles prédits par la théorie
de l'évolution[24].
Le fait que la rétine des céphalopodes et celle des vertébrés ne soit pas « montée » dans le même
sens montre que les yeux de ces deux groupes ne sont pas homologues et se sont développés par des voies
différentes. Ce montage à l'envers, considéré comme une imperfection chez les vertébrés, est présenté par
Richard Dawkins[25] comme une preuve de l'imperfection de l'œil des vertébrés, et donc du fait que ce
dernier ne résulte pas de l'application d'un plan préétabli mais au contraire de d'adaptations successives,
comme un argument contre le créationnisme et le Dessein Intelligent.
Certaines considérations pourraient montrer ce montage inverse de la rétine des vertébrés comme une
imperfection : existence d'un point aveugle, rétine moins attachée au fond de l'œil (?), lumière filtrée par la
rétine avant les récepteurs...
Néanmoins, certaines de ces caractéristiques « néfastes » ont été minimisés : comme le point aveugle dont
l'étendue a été réduite, une liaison plus forte avec le fond de l'œil (?) Ou bien même, peut être utilisée(s)
comme avantage(s) dans certaines circonstances : comme le pouvoir filtrant de la couche rétinienne.
→ C'est un exemple qui montre que des structures des divers organes dont la réponse à une
contrainte externe, ici filtrer des radiations nocives, comme les ultraviolets ou régler l'intensité lumineuse),
présentent souvent des possibilités multiples, ici la rétine servant de filtre, mais l'humeur vitrée ou/et le
cristallin peuvent aussi jouer ce rôle.
Dans ces considérations, il faut en fait dissiper l'ambiguïté des notions d'organes « élémentaires »,
ou « d'ébauches » d'organes. Les yeux, ou les membres (ou d'autres organes) sont plus ou moins
performants selon leur utilisation dans un organisme donné (vers, poissons, reptiles, oiseaux...) mais ils ont
toujours été « formés » et diversement sélectionnés suivant la lignée.

Le second principe de la thermodynamique [modifier]

Se fondant sur une simplification du deuxième principe de la thermodynamique, certains pseudo-
scientifiques[26],[27] avancent que la physique interdirait l'évolution.
L'augmentation globale de l'entropie, variable de la thermodynamique quantifiant le désordre
statistique d'un système, serait censée interdire l'évolution, qui engendre au contraire la complexité.
Cette affirmation doit être recadrée :
 1. Cette « conclusion » s'applique à l'entropie totale d'un système fermé, ce que la Terre n'est
pas. L'entropie peut diminuer localement s'il y a échange d'énergie avec un autre endroit où l'on a
une augmentation supérieure de l'entropie : typiquement au niveau du Soleil dans le cas de la vie sur
Terre[28]. Ou plus précisément dans la transformation du rayonnement solaire reçu par la Terre.
Globalement la Terre reçoit de l'énergie sous une forme plus « ordonnée » qu'elle n'en réémet : les
rayons solaires sont reçus depuis une zone précise du ciel (le disque solaire), principalement dans le
spectre visible ; alors que la Terre réémet cette énergie reçue sous forme de rayonnement
infrarouge, par un nombre bien supérieur de photons (~20 fois plus) dans toutes les directions du
ciel.
2. Or les êtres vivants sont tous des systèmes « ouverts » (thermodynamiquement parlant) ;
tendant à toutes étapes de leur existence à s'autoconserver, c'est-à-dire à maintenir leur
organisation interne au détriment (au prix) d'une production continue « d'entropie », « de désordre »
(rejets[29], chaleur[30]) payant ainsi le « tribut » à la règle d'entropie. Ils ne sont aucunement des
exceptions à ce principe, contrairement à certaines affirmations péremptoires prétendument
définitives[27]. De plus, le second principe est un principe général, qui montre qu'un système donné
(vivant ou inerte) peut (ou non) fonctionner ; son type interne de fonctionnement est (ou peut être)
expliqué par d'autres sciences (chimie, mécanique, électronique...)
La possibilité d'évolution des populations d'êtres vivants est parfaitement conforme au second
principe, tout comme leur fonctionnement, leur développement, leur conservation. Cette possibilité
s'intègre simplement dans cette succession des générations, qui est elle-même parfaitement
conforme au second principe.
L'évolution ne produit pas directement plus « d'informations » ; il est très difficile de préciser
« l'information » contenue dans les êtres vivants. Du moins le scénario principal le suggère-t-il à
l'échelle des temps géologiques (des centaines de millions aux milliards d'années) ; comme on l'a
vu, cette possibilité est conforme au second principe dans la biosphère. Le questionnement se
portant sur la (ou les) manière(s) dont ça c'est produit.
3. Il n'est pas immédiat d'assimiler le désordre statistique d'objets indifférenciés et
indépendants comme les molécules d'un gaz parfait et l'entropie au sein d'un système prébiotique
comportant différents constituants en interaction chimique. L'évaluation statistique effective des
configurations les plus probables est bien moins évidente et rien n'empêche certains constituants de
se « construire » aux dépens d'autres.
Ainsi il est tout à fait faux de prétendre calculer simplement l'apparition de « la vie » en termes de
« probabilités » simplistes, comme présenté ici[31], ou parfois ailleurs ! Car ce type de calcul
simpliste ignore implicitement complètement tous les cadres dans lequel les diverses réactions ont
(ou peuvent avoir) lieu : mares, évents, plages, volcans, argiles, impacts météoritiques/cométaires
produisant simplement des peptides...
Demeurent toujours certaines interrogations techniques concernant les règles de repliement de
protéines ; question centrale dans la théorie probabiliste de l'organisation de la vie[32]. Les maladies
à prion témoignent de l'importance des différentes règles de pliages des protéines.
4. La physique comporte déjà plusieurs phénomènes qui peuvent sembler violer le deuxième
principe de la thermodynamique si on traduit « entropie » par la notion subjective de « désordre » :
avec la formation des étoiles et des galaxies, la nucléosynthèse, l'Univers croît en entropie tout en
paraissant plus ordonné.
Il serait plus précis de dire que la matière se présente bien sous forme de structures plus ordonnées,
mais l'Univers dans son ensemble est en fait globalement plus désordonné, principalement par le
rayonnement émis lors de la formations de ces structures.
 Critiques à caractère religieux [modifier]
Face au consensus scientifique actuel, certains groupes contestent l'existence de l'évolution. Il s'agit
en général de milieux religieux, tenants de diverses formes de créationnisme et de prédéterminisme. Les
raisons de cette opposition sont la contradiction avec une interprétation littérale de leurs textes sacrés (en
particulier la Genèse), la négation de la volonté divine dans la création du monde et de l'homme, ainsi que le
fait que la théorie de l'évolution n'accorde pas de place particulière (du moins dans une première analyse) à
l'être humain dans l'univers et le monde vivant.
Cette opposition n'est nullement unanime parmi les religions. L'Église catholique par exemple ne
partage pas l'interprétation littérale du livre de la Genèse. La théorie de l'évolution a été présentée comme
une hypothèse scientifique compatible avec la doctrine catholique par le pape Pie XII dans l'encyclique
Humani Generis en 1950. Le pape Jean-Paul II a affirmé en 1996 que c'était « plus qu'une hypothèse.[33]»,
et le chef astronome du Vatican, le révérend George Coyne, a affirmé que le Dessein Intelligent « n'est pas
de la science, même s'il en a la prétention.[34]». De même, le bouddhisme ne rejette pas la théorie de
l'évolution dont les notions sont même plutôt en accord avec la pensée bouddhiste[35].
En science, une théorie est une explication cohérente des phénomènes naturels basés sur
l'observation directe et l'expérimentation. Les théories sont logiques, prédictives et testables. Elles sont
ouvertes à la critique et lorsqu'elles apparaissent erronées, elles sont modifiables ou annulables. Sur base
de cette définition, l'évolution est catégorisée avec d'autres théories scientifiques telle que les théories de la
gravité ou la théorie atomique, qui, comme l'évolution, sont universellement acceptées par les scientifiques.
(...) Contrairement à l'évolution, le dessein intelligent et le créationnisme ne sont pas de la science parce
qu'ils échouent à rencontrer les exigences essentielles et nécessaires : ils ne sont pas fondés sur
l'observation directe ou l'expérimentation pas plus qu'ils ne génèrent des prédictions testables. Dès lors, offrir
ces « croyances » comme alternative à l'évolution ou donner un temps égal dans les classes de science
dénature complètement la nature de la science. [36]

Créationnisme [modifier]
Article détaillé : Créationnisme.
Ces critiques du darwinisme font référence aux attaques des tenants d'une vision religieuse ou
spiritualiste généralement inspirée d'une exégèse littérale de la Bible, contre les présentations de thèmes
présentés comme issus de recherches des scientifiques travaillant à la théorie darwinienne. En tant que
telles ces critiques ne sont pas scientifiques, mais s'inscrivent plutôt dans le débat sur raison et foi et en
particulier science et christianisme.
• Le pape Jean-Paul II critiqua cette position sur la science dans le cadre de l'affaire Galilée :
« Ainsi la science nouvelle, avec ses méthodes et la liberté de recherche qu'elle suppose, obligeait
les théologiens à s'interroger sur leurs propres critères d'interprétation de l'Écriture.
La plupart n'ont pas su le faire. »
• En 1996, le pape affirma que :
« L'évolution était plus qu'une hypothèse.[37] »
Dès lors, c'est surtout des milieux protestants que proviennent les critiques du darwinisme.

Le Dessein Intelligent [modifier]

Article détaillé : Dessein Intelligent.
 Vue postérieure de l'os sphénoïde. Selon Anne Dambricourt Malassé et Marie-Joseph Deshayes le
fléchissement de l'os sphénoïde dans le même sens depuis 60 millions d'années a complètement orienté et
continuerait d'orienter l'évolution de l'Homme...
Le dessein intelligent (Intelligent Design en anglais[38]) est la thèse selon laquelle « certaines
observations de l'Univers et du monde du vivant sont mieux expliquées par une cause intelligente que par
des processus aléatoires tels que la sélection naturelle. »[39]. Cette thèse a été développée par le Discovery
Institute, un cercle de réflexion conservateur chrétien américain. Le dessein intelligent est présenté comme
une théorie scientifique par ses promoteurs, mais dans le monde scientifique, il est considéré comme
relevant de la pseudo-science, tant par des arguments aussi bien internes à la biologie (les promoteurs du
dessein intelligent apparaissant aux biologistes comme ne tenant pas compte de nombreuses observations)
qu'épistémologiques (en particulier le critère de réfutabilité de Karl Popper).
La plupart des commentateurs y voient une résurgence du créationnisme, dissimulée sous une
apparence de scientificité, et les Américains la classent désormais dans les théories néo-créationnistes, en
particulier suite à la publication du Wedge document (voir objectifs et stratégie). D'un point de vue
idéologique, les deux thèses sont apparentées (intervention d'une puissance supérieure).
• Argumentation : Deux chercheuses françaises, Anne Dambricourt Malassé et Marie-Joseph
Deshayes défendent, que l'os sphénoïde, un os situé au centre du crâne, jouerait un rôle clef dans
l'évolution de notre espèce, et particulièrement dans notre position redressée et la bipédie[40].

Articles connexes [modifier]

• Créationnisme
• Néo-créationnisme
• Dessein Intelligent
• Obscurantisme
• Scientisme
Liens externes [modifier]
• Dessein Intelligent - Intelligent Design (Site Hominidés.com)
• Le darwinisme est parfois contesté dans les salles de classe françaises (Site du Centre
Roger Ikor)
• États-Unis: darwinisme ou "dessein intelligent" ? (Site de la BBC)
• les fondamentalistes chrétiens américains contre Darwin
• Paul Lafargue, Le darwinisme sur la scène française, Die Neue Zeit, 1890, p. 184-189.
• Anton Pannekoek, Darwinisme et marxisme, 1909 1er Partie 2e Partie.
• Effervesciences (CINAPS Télévision) : Darwin aujourd'hui (avec Guillaume Lecointre)

Ouvrages [modifier]
• Pierre-Paul Grassé, L'évolution du vivant, matériaux pour une théorie transformiste , Albin
Michel, 1973
• Rémy Chauvin, Le Darwinisme ou la fin d’un mythe, du Rocher, 1997
• Anne Dambricourt-Malassé, La Légende maudite du Vingtième siècle : L'Erreur darwinienne,
Nuée Bleue, 2000
• Stephen Jay Gould, Et Dieu dit : Que Darwin soit ! : Science et religion, enfin la paix ? ,
préface de Dominique Lecourt, Seuil, 2000.
• Dominique Lecourt, L’Amérique entre la Bible et Darwin, suivi de Intelligent design : science,
morale et politique'' (1992, 3e réed. Quadrige/PUF, 2007).
Notes et références [modifier]
1. ↑ Résumé de : LARGEAULT J., Réductionnisme et holisme, Encyclopédia universalis,
(2000) vol. 19, p. 523-527.
2. ↑ Platon, Apologie de Socrate, Criton, Phédon, trad. M.-J. Moreau, ed. Gallimard, Folio
Essais, 1985.
3. ↑ Aristote, La Métaphysique, trad. Annick Jaulin, PUF, 1999.
4. ↑ a et b Jacques MONOD Le Hasard et la Nécessité, éd. du Seuil, coll. Points, pp. 37-38.
5. ↑ Smuts, Jan. Holism and Evolution. Londres: Macmillan & Co Ldt, 1926, p. 362.
6. ↑ Stephen Jay Gould, Darwin et les grandes énigmes de la vie, éd Point, 1997, p225-228
7. ↑ Karl Marx, Le Capital, Livre I, éd Puf, Chap XII Division du Travail et Manufacture, §2 Le
travailleur Partiel et son outil, p.384
8. ↑ in Karl Marx, Le Capital, ed Folio essai, Chap XIV La Manufacture, §II Le travailleur
parcellaire et son outil, p.430
9. ↑ Stephen Jay Gould, Cette vision de la vie, éd du Seuil, 2004, p143-144
10.↑ « La vie, étant la forme de l'être qui nous est le plus connue, est spécifiquement une
volonté d'accumuler la force : - Tous les procès de la vie ont là leur levier ; rien ne veut se conserver,
tout doit être additionné et accumulé. La vie, en tant que cas particulier (l'hypothèse qui, en partant
de là, aboutit au caractère général de l'existence) - aspire à un sentiment maximal de puissance; elle
est essentiellement l'aspiration à un surplus de puissance ; aspirer, ce n'est point autre chose que
d'aspirer à la puissance ; cette volonté demeure ce qu'il y a de plus intime et de plus profond : la
mécanique est une simple sémiotique des conséquences. »
11.↑ « Périssent les faibles et les ratés ! Et il faut même les y aider ! »
 12.↑ « Jusque ici toute élévation du type humain a été l'œuvre d'une société aristocratique, et il
en sera toujours ainsi. »
13.↑ « L'humanité n'avance pas d'un seul trait ; souvent le type déjà atteint se perd de nouveau
(malgré les efforts de trois siècles, nous n'avons plus pu atteindre de nouveau l'homme de la
Renaissance, et, d'autre part, l'« homme » de la Renaissance était resté en arrière sur l'homme de
l'Antiquité). »
14.↑ « Ce qui me surprend le plus, lorsque je passe en revue les grandes destinées de
l'humanité, c'est d'avoir toujours devant les yeux le contraire de ce que voient ou veulent voir
aujourd'hui Darwin et son école. Eux constatent la sélection en faveur des êtres plus forts et mieux
venus, le progrès de l'espèce. Mais c'est précisément le contraire qui saute aux yeux : la suppression
des cas heureux, l'inutilité des types mieux venus, la domination inévitable des types moyens et
même de ceux qui sont au-dessous de la moyenne. À moins que l'on nous démontre la raison qui fait
que l'Homme est l'exception parmi les créatures, j'incline à croire que l'école de Darwin s'est partout
trompée. »
15.↑ « Et qui est celui qui m'appelle là-bas ? » « Tu le sais bien, répondit vivement le devin,
pourquoi te caches-tu ? C'est l'homme supérieur qui t'appelle à son secours ! » Ainsi parlait
Zarathoustra, Quatrième partie, Le cri de détresse.
16.↑ « Aujourd'hui les petites gens sont devenus les maitres, ils prêchent tous la résignation, et
la modestie, et la prudence, et l'application, et les égards et le long ainsi-de-suite des petites vertus.
[...] Surmontez, hommes supérieurs, les petites vertus, les petites prudences, les égards pour les
grains de sable, le fourmillement des fourmis, le misérable contentement de soi, le « bonheur du plus
grand nombre ! » — Et désespérez plutôt que de vous rendre. Car, en vérité, je vous aime, parce que
vous ne savez pas vivre aujourd'hui, ô hommes supérieurs ! Car c'est ainsi que vous vivez — le
mieux ! » Ainsi parlait Zarathoustra, Quatrième partie, De l'homme supérieur.
 17.↑ « Qui devra venir un jour et n'aura pas le droit de passer ? Notre grand hasard, c'est-à-dire
le grand et lointain Empire de l'Homme, le règne de Zarathoustra qui dure mille ans. » Ainsi parlait
Zarathoustra, Quatrième partie, L'offrande du miel.
18.↑ Vincent Homer, Évolution et développement : Vers une nouvelle synthèse ? [archive],
Université Paris IV, Sorbonne.
19.↑ Alain Feuerbacher, Désaccords d'Évolution [archive]
20.↑ Pour un catastrophisme éclairé. Quand l'impossible est certain de Jean-Pierre Dupuy,
Seuil, 2002.
21.↑ Catastrophisme, administration du désastre et soumission durable , de René Riesel et
Jaime Semprun, Encyclopédie des Nuisances, 2008.
22.↑ Cette théorie explique à la fois la disparition des dinosaures et l'importante radiation
évolutive des oiseaux et des mammifères qu'a connue la Terre il y a 65 Ma. Selon la paléontologie
moderne, chaque extinction massive est de cette façon suivie d'une radiation évolutive importante.
Tout cela, bien que mettant en cause des catastrophes suivies d'une multiplication systématique de
nouvelles formes d'espèces, ne relève cependant pas forcément de la théorie catastrophiste, car les
fondements du catastrophisme s'opposent par principe à ceux de la théorie de l'évolution.
23.↑ Dans certaines circonstances, comme celles du Cambrien.
24.↑ Nilsson & Pelger, A pessimistic estimate of the time required for an eye to evolve.Proc.
Biol. Sci .1994 Apr 22;256(1345):53-8
25.↑ L'Horloger Aveugle, 1986.
26.↑ Emmanuel Bozzi, La deuxième loi de la thermodynamique contredit la théorie de
l'évolution [archive], Bibliste.com
27.↑ a et b Harun Yahya, La thermodynamique réfute l'évolution [archive], Le Mensonge de
l'évolution.com
 28.↑ Jacques Monod, Le Hasard et la Nécessité, Seuil, 1970 (ISBN 2020028123)
29.↑ Rejets (excréments, dioxyde de carbone, eau...), qui pourront de nouveau être recyclés
par dégradation de l'énergie solaire.
30.↑ Chaleur, qui est dissipée dans le rayonnement infrarouge de la Terre (qui est elle-même
un système ouvert).
31.↑ Harun Yahya, L'impasse moléculaire de l'évolution [archive], Le Mensonge de
l'évolution.com.
32.↑ Folding@Home Interview:Fold a Protein [archive] Simulation vidéo d'un pliage de protéine
selon les lois physico-chimiques actuellement connue sur cette page. Présentée sur DL TV par
Veronica Belmont et Robert Heron.
33.↑ Jean Paul II, Message delivered to the Pontifical Academy of Sciences 22 October
1996 [archive], Eternal Word Television Network.
34.↑ Vatican official: 'Intelligent design' isn't science [archive], USAToday.com, 18 novembre
2005.
35.↑ Jacques Brosse, Écologie, bouddhisme et christianisme [archive], Nouvelles Clés, (page
consultée le 26 avril 2008).
36.↑ « FASEB opposes using science classes to teach intelligent design, creationism, and
other non-scientific beliefs », The FASEB Journal, 2006;20:408-409, p. 408
37.↑ EVOLUTION WATCH [archive] Posted: November 03, 2005 2:25 pm Eastern ; © 2009
WorldNetDaily.com
38.↑ La traduction en français de design par dessein est devenue usuelle dans ce cadre. Une
traduction par conception serait plus exacte et permettrait de traduire intelligent designer par
concepteur intelligent plutôt que par cause intelligente.
 39.↑ Voir site du Discovery Institute, un des promoteurs du Dessein Intelligent, Questions
About Intelligent Design [archive]
40.↑ L'hypothèse est fondée sur l'étude des crânes fossiles des différentes lignées de primates,
ainsi que sur le développement actuel de notre espèce, elles décrivent un processus interne
d'évolution : l'Inside Story ! L'os sphénoïde se fléchirait graduellement dans le même sens depuis
des dizaines de millions d'années et « orienterait » notre évolution. L'os continuerait toujours de se
fléchir dans le même sens expliquant selon les chercheuses la disparition progressive des dents de
sagesse à cause de la réduction de l'espace disponible dans la bouche... (Voir site : [1] [archive]).
Bien que les auteurs de cette thèse prétendent ne pas défendre la théorie du dessein intelligent :
(Voir : La logique de l'évolution [archive]), des néocréationnistes se sont appropriés celle-ci.
Ce document provient de « http://fr.wikipedia.org/wiki/Historique_des_critiques_des_th
%C3%A9ories_de_l%27%C3%A9volution ».

Catégories : Histoire de la biologie de l'évolution | Concept historique en biologie de l'évolution |

Créationnisme
 W000

Creation–evolution controversy
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The creation–evolution controversy (also termed the creation vs. evolution debate or the origins
debate) is a recurring cultural, political, and theological dispute about the origins of the Earth, humanity, life,
and the universe.[1] The dispute is between those who espouse the belief of creationism and the proponents
of evolution, backed by scientific consensus. The dispute particularly involves the field of evolutionary
biology, but also the fields of geology, palaeontology, thermodynamics, nuclear physics and cosmology.[2]
Though also present in Europe and elsewhere,[3] and often portrayed as part of the culture wars,[4] this
debate is most prevalent in the United States.[citation needed] While the controversy has a long history,[5]
today it is mainly over what constitutes good science,[6] with the politics of creationism primarily focusing on
the teaching of creation and evolution in public education.[7]
The debate also focuses on issues such as the definition of science (and of what constitutes
scientific research and evidence), science education (and whether the teaching of the scientific consensus
view should be 'balanced' by also teaching fringe theories), free speech, separation of Church and State, and
theology (particularly how different Christians and Christian denominations interpret the Book of Genesis).
Within the scientific community and academia the level of support for evolution is essentially
universal,[8] while support for biblically-literal accounts or other creationist alternatives is very small among
scientists, and virtually nonexistent among those in the relevant fields.[9]
The debate is sometimes portrayed as being between science and religion. However, as the National
Academy of Sciences states:
Today, many religious denominations accept that biological evolution has produced the diversity
of living things over billions of years of Earth’s history. Many have issued statements observing
that evolution and the tenets of their faiths are compatible. Scientists and theologians have
written eloquently about their awe and wonder at the history of the universe and of life on this
planet, explaining that they see no conflict between their faith in God and the evidence for
evolution. Religious denominations that do not accept the occurrence of evolution tend to be
those that believe in strictly literal interpretations of religious texts.
—National Academy of Sciences, Science, Evolution, and Creationism[10]
Contents
[hide]
• 1 History of the controversy
• 1.1 Controversies in the age of Darwin
• 1.2 Creationism
• 1.2.1 The Butler Act and the Scopes
monkey trial
• 1.2.2 Epperson v. Arkansas
• 1.2.3 Daniel v. Waters
• 1.3 Creation Science
• 1.3.1 Court cases
• 1.3.1.1 McLean v. Arkansas
• 1.3.1.2 Edwards v. Aguillard
• 1.4 Intelligent Design
• 1.4.1 Controversy in recent times
• 1.4.2 Kansas evolution hearings
• 1.4.3 The Dover Trial
• 2 Viewpoints
• 2.1 Young Earth creationism
• 2.2 Old Earth creationism
• 2.3 Neo-Creationism
[edit] History of the controversy
See also: History of evolutionary thought
The creation-evolution controversy originated from Europe and North America in the late eighteenth
century when discoveries in geology led to various theories of an ancient earth, and fossils showing past
extinctions prompted early ideas of evolution, notably Lamarckism. In England these ideas of continuing
change were seen as a threat to the fixed social order, and were harshly repressed.[11] Conditions eased,
and in 1844 the controversial Vestiges popularised transmutation of species. The scientific establishment
dismissed it scornfully and the Church of England reacted with fury, but many Unitarians, Quakers and
Baptists opposed to the privileges of the Established church favoured its ideas of God acting through laws.
[12]

[edit] Controversies in the age of Darwin

See also: Reaction to Darwin's theory
 A satirical 1871 image of Charles Darwin as an ape reflects part of the social controversy over
whether humans and apes share a common lineage.
 Publication of Charles Darwin's On the Origin of Species by Means of Natural Selection in 1859
brought scientific credibility to evolution, and made it a respectable field of study.[13]
There was intense interest in the religious implications of Darwin's book, but the Church of England's
attention was largely diverted by theological controversy over higher criticism set out in Essays and Reviews
by liberal Christian authors, some of whom expressed support for Darwin, as did many nonconformists. The
Reverend Charles Kingsley openly supported the idea of God working through evolution. However, many
Christians were opposed to the idea and even some of Darwin's close friends and supporters including
Charles Lyell and Asa Gray could not accept some of his ideas.[14] Thomas Huxley, who strongly promoted
Darwin's ideas while campaigning to end the dominance of science by the clergy, coined the term agnostic to
describe his position that God’s existence is unknowable, and Darwin also took this position,[14] but
evolution was also taken up by prominent atheists including Edward Aveling and Ludwig Büchner and
criticised, in the words of one reviewer, as "tantamount to atheism."[15] Following the lead of figures such as
St. George Jackson Mivart and John Zahm Catholics in the United States were accepting of evolution itself
while ambivalent towards natural selection and stressing humanity's divinely imbued soul.[16] Though
evolution was never condemned by the church, initially the more conservative leaning Catholic leadership in
Rome held back but gradually adopted a similar position.[16][17]
Creationists during this period were largely premillennialists, whose belief in Christ's return depended
on a quasi-literal reading of the Bible.[18] However, they were not as concerned about geology, freely
granting scientists any time they needed before the Garden of Eden to account for scientific observations,
such as fossils and geological findings.[19] In the immediate post-Darwinian era, few scientists or clerics
rejected the antiquity of the earth or the progressive nature of the fossil record.[20] Likewise, few attached
geological significance to the Biblical flood, unlike subsequent creationists.[20] Evolutionary skeptics,
creationist leaders and skeptical scientists were usually willing either to adopt a figurative reading of the first
chapter of Genesis, or to allow that the six days of creation were not necessarily 24-hour days.[21]

[edit] Creationism
Main article: History of creationism
See also: Creation and evolution in public education
During the 1800s and up until the mid-1900s, Creationism was widely accepted and was considered
a foundational truth, but there was no official resistance to evolution by mainline denominations within the
United States of America.[14] Around the start of the 20th century some evangelical scholars had ideas
accommodating evolution, such as B. B. Warfield who saw it as a natural law expressing God’s will.
However, development of the eugenics movement led many Catholics to reject evolution.[14] In this
enterprise they received little aid from conservative Christians in Britain and Europe. In Britain this has been
attributed to their minority status leading to a more tolerant, less militant theological tradition. The main
British Creationist movement in this period was the Evolution Protest Movement, formed in the 1930s.[22]

[edit] The Butler Act and the Scopes monkey trial

Main article: Scopes Trial
 Clarence Darrow and William Jennings Bryan chat in court during the Scopes trial.
In the aftermath of World War I, the Fundamentalist-Modernist Controversy brought a surge of
opposition to the idea of evolution, and following the campaigning of William Jennings Bryan several states
introduced legislation prohibiting the teaching of evolution. By 1925, such legislation was being considered in
15 states, and passed in some states, such as Tennessee.[23] The American Civil Liberties Union offered to
defend anyone who wanted to bring a test case against one of these laws. John T. Scopes accepted, and he
confessed to teaching his Tennessee class evolution in defiance of the Butler Act. The textbook in question
was Hunter's Civic Biology (1914). The trial was widely publicized by H. L. Mencken among others, and is
commonly referred to as the Scopes Monkey Trial. Scopes was convicted; however, the widespread publicity
galvanized proponents of evolution. When the case was appealed to the Tennessee Supreme Court, the
Court overturned the decision on a technicality (the judge had assessed the minimum $100 fine instead of
allowing the jury to assess the fine).[24][25]
 Although it overturned the conviction, the Court decided that the law was not in violation of the
Religious Preference provisions of the Tennessee Constitution (section 3 of article 1), which stated that "that
no preference shall ever be given, by law, to any religious establishment or mode of worship."[26] The Court,
applying that state Constitutional language, held
We are not able to see how the prohibition of teaching the theory that man has descended from
a lower order of animals gives preference to any religious establishment or mode of worship. So
far as we know, there is no religious establishment or organized body that has in its creed or
confession of faith any article denying or affirming such a theory....Protestants, Catholics, and
Jews are divided among themselves in their beliefs, and that there is no unanimity among the
members of any religious establishment as to this subject. Belief or unbelief in the theory of
evolution is no more a characteristic of any religious establishment or mode of worship than is
belief or unbelief in the wisdom of the prohibition laws. It would appear that members of the
same churches quite generally disagree as to these things.

...Furthermore, [the Butler Act] requires the teaching of nothing. It only forbids the teaching of
evolution of man from a lower order of animals....As the law thus stands, while the theory of
evolution of man may not be taught in the schools of the State, nothing contrary to that theory
[such as Creationism] is required to be taught.

...It is not necessary now to determine the exact scope of the Religious Preference clause of the
Constitution ... Section 3 of article 1 is binding alike on the Legislature and the school
authorities. So far we are clear that the Legislature has not crossed these constitutional
limitations.
 —Scopes v. State, 289 S.W. 363, 367 (Tenn. 1927).[27]

The interpretation of the Establishment clause up to that time was that the government could not
establish a particular religion as the State religion. The Tennessee Supreme Court's decision held in effect
that the Butler Act was constitutional under the state Constitution's Religious Preference Clause, because the
Act did not establish one religion as the "State religion."[28] As a result of the holding, the teaching of
evolution remained illegal in Tennessee, and continued campaigning succeeded in removing evolution from
school textbooks throughout the United States.[29][30]

[edit] Epperson v. Arkansas

Main article: Epperson v. Arkansas
In 1968, the United States Supreme Court invalidated a forty year old Arkansas statute that
prohibited the teaching of evolution in the public schools. A Little Rock high school biology teacher, Susan
Epperson, filed suit charging the law violated the federal constitutional prohibition against establishment of
religion as set forth in the Establishment Clause. The Little Rock Ministerial Association supported
Epperson's challenge, declaring, "to use the Bible to support an irrational and an archaic concept of static
and undeveloping creation is not only to misunderstand the meaning of the Book of Genesis, but to do God
and religion a disservice by making both enemies of scientific advancement and academic freedom."[31] The
Court held that the United States Constitution prohibits a state from requiring, in the words of the majority
opinion, "that teaching and learning must be tailored to the principles or prohibitions of any religious sect or
dogma."[32] But the Supreme Court decision also suggested that creationism could be taught in addition to
evolution.[33]
 [edit] Daniel v. Waters
Main article: Daniel v. Waters
Daniel v. Waters was a 1975 legal case in which the United States Court of Appeals for the Sixth
Circuit struck down Tennessee's law regarding the teaching of "equal time" of evolution and creationism in
public school science classes because it violated the Establishment clause of the US Constitution. Following
this ruling, creationism was stripped of overt biblical references and renamed creation science, and several
states passed legislative acts requiring that this be given equal time with the teaching of evolution.

[edit] Creation Science

Main article: Creation Science
As biologists grew more and more confident in evolution as the central defining principle of biology,
[34] American membership in churches favoring increasingly literal interpretations of scripture rose, with the
Southern Baptist Convention and Lutheran Church - Missouri Synod outpacing all other denominations.[35]
With growth, these churches became better equipped to promulgate a creationist message, with their own
colleges, schools, publishing houses, and broadcast media.[36]
In 1961, the first major modern creationist book was published: Henry M. Morris and John C.
Whitcomb Jr.'s The Genesis Flood. Morris and Whitcomb argued that creation was literally 6 days long, that
humans lived concurrently with dinosaurs, and that God created each 'kind' of life individually.[37] On the
strength of this, Morris became a popular speaker, spreading anti-evolutionary ideas at fundamentalist
churches, colleges, and conferences.[38] Morris' Creation Science Research Center (CSRC) rushed
publication of biology text books that promoted creationism, and also published other books such as Kelly
Segrave's sensational Sons of God Return that dealt with UFOlogy, flood geology, and demonology against
Morris' objections.[39] Ultimately, the CSRC broke up over a divide between sensationalism and a more
intellectual approach, and Morris founded the Institute for Creation Research, which was promised to be
controlled and operated by scientists.[40] During this time, Morris and others who supported flood geology
adopted the terms scientific creationism and creation science.[41] The flood geologists effectively co-opted
"the generic creationist label for their hyperliteralist views".[42]

[edit] Court cases

[edit] McLean v. Arkansas

Main article: McLean v. Arkansas
In 1982 another case in Arkansas ruled that the Arkansas "Balanced Treatment for Creation-Science
and Evolution-Science Act" was unconstitutional because it violated the establishment clause of the U.S.
Constitution. Much of the transcript of the case was lost, including evidence from Francisco Ayala.

[edit] Edwards v. Aguillard

Main article: Edwards v. Aguillard
In the early 1980s, the Louisiana legislature passed a law titled the "Balanced Treatment for
Creation-Science and Evolution-Science in Public School Instruction Act". The act did not require teaching
either evolution or creationism as such, but did require that when evolutionary science was taught, creation
science had to be taught as well. Creationists had lobbied aggressively for the law, arguing that the act was
about academic freedom for teachers, an argument adopted by the state in support of the act. Lower courts
ruled that the State's actual purpose was to promote the religious doctrine of creation science, but the State
appealed to the Supreme Court.
In the similar case of McLean v. Arkansas (see above) the federal trial court had also decided against
creationism. Mclean v. Arkansas however was not appealed to the federal Circuit Court of Appeals,
creationists instead thinking that they had better chances with Edwards v. Aguillard. In 1987 the Supreme
Court of the United States ruled that the Louisiana act was unconstitutional, because the law was specifically
intended to advance a particular religion. At the same time, however, it stated in its opinion that "teaching a
variety of scientific theories about the origins of humankind to school children might be validly done with the
clear secular intent of enhancing the effectiveness of science instruction," leaving open the door for a handful
of proponents of creation science to evolve their arguments into the iteration of creationism that came to be
known as intelligent design.[43]

[edit] Intelligent Design

 The Discovery Institute's Center for the Renewal of Science and Culture used banners based on
"The Creation of Adam" from the Sistine Chapel. Later it used a less religious image, then was renamed the
Center for Science and Culture.[44]
Main article: Intelligent design
See also: Neo-creationism, Intelligent design movement, Teach the Controversy, and Critical
Analysis of Evolution
In response to Edwards v. Aguillard, the Neo-Creationist intelligent design movement was formed
around the Discovery Institute's Center for Science and Culture. Its goal is to restate creationism in terms
more likely to be well received by the public, policy makers, educators, and the scientific community, and
makes the claim that "certain features of the universe and of living things are best explained by an intelligent
cause, not an undirected process such as natural selection."[45] It has been viewed as a "scientific"
approach to creationism by creationists, but is widely rejected as unscientific by the science community —
primarily because intelligent design cannot be tested and rejected like scientific hypotheses (see for
example, list of scientific societies rejecting intelligent design).

[edit] Controversy in recent times

See also: Politics of creationism and Intelligent design in politics
The controversy continues to this day, with the mainstream scientific consensus on the origins and
evolution of life challenged by creationist organizations and religious groups who desire to uphold some form
of creationism (usually young earth creationism, creation science, old earth creationism or intelligent design)
as an alternative. Most of these groups are explicitly Christian, and more than one sees the debate as part of
the Christian mandate to evangelize.[46] Some see science and religion as being diametrically opposed
views which cannot be reconciled. More accommodating viewpoints, held by many mainstream churches and
many scientists, consider science and religion to be separate categories of thought, which ask fundamentally
different questions about reality and posit different avenues for investigating it.[47] Public opinion in regards
to the concepts of evolution, creationism, and intelligent design is fluctuating.
More recently, the Intelligent Design movement has taken an anti-evolution position which avoids
any direct appeal to religion. Scientists argue that Intelligent design does not represent any research program
within the mainstream scientific community, and is essentially creationism.[48] Its leading proponent, the
Discovery Institute, made widely publicised claims that it was a new science, though the only paper arguing
for it published in a scientific journal was accepted in questionable circumstances and quickly disavowed in
the Sternberg peer review controversy, with the Biological Society of Washington stating that it did not meet
the journal's scientific standards, was a "significant departure" from the journal's normal subject area and was
published at the former editor's sole discretion, "contrary to typical editorial practices".[49] President Bush
commented endorsing the teaching of Intelligent design alongside evolution "I felt like both sides ought to be
properly taught ... so people can understand what the debate is about."[50]

[edit] Kansas evolution hearings

Main article: Kansas evolution hearings
In the push by intelligent design advocates to introduce intelligent design in public school science
classrooms, the hub of the intelligent design movement, the Discovery Institute, arranged to conduct
hearings to review the evidence for evolution in the light of its Critical Analysis of Evolution lesson plans. The
Kansas Evolution Hearings were a series of hearings held in Topeka, Kansas 5 May to 12 May 2005. The
Kansas State Board of Education eventually adopted the institute's Critical Analysis of Evolution lesson plans
over objections of the State Board Science Hearing Committee, and electioneering on behalf of conservative
Republican candidates for the Board.[51] On 1 August 2006, 4 of the 6 conservative Republicans who
approved the Critical Analysis of Evolution classroom standards lost their seats in a primary election. The
moderate Republican and Democrats gaining seats vowed to overturn the 2005 school science standards
and adopt those recommended by a State Board Science Hearing Committee that were rejected by the
previous board,[52] and on 13 February 2007, the Board voted 6 to 4 to reject the amended science
standards enacted in 2005. The definition of science was once again limited to "the search for natural
explanations for what is observed in the universe."[53]

[edit] The Dover Trial

Main article: Kitzmiller v. Dover Area School District
Following the Edwards v. Aguillard decision by the Supreme Court of the United States, in which the
Court held that a Louisiana law requiring that creation science be taught in public schools whenever evolution
was taught was unconstitutional, because the law was specifically intended to advance a particular religion,
creationists renewed their efforts to introduce creationism into public school science classes. This effort
resulted in intelligent design, which sought to avoid legal prohibitions by leaving the source of creation to an
unnamed and undefined intelligent designer, as opposed to God.[54] This ultimately resulted in the "Dover
Trial," Kitzmiller v. Dover Area School District, which went to trial on 26 September 2005 and was decided on
20 December 2005 in favor of the plaintiffs, who charged that a mandate that intelligent design be taught in
public school science classrooms was an unconstitutional establishment of religion. The 139 page opinion of
Kitzmiller v. Dover held that intelligent design was not a subject of legitimate scientific research, and that it
"cannot uncouple itself from its creationist, and hence religious, antecedents".[55]
[edit] Viewpoints
[edit] Young Earth creationism
Main article: Young Earth creationism
See also: Creation science and Flood geology
Young Earth creationism is the belief that the Earth was created by God within the last 10,000 years,
literally as described in Genesis, within the approximate timeframe of biblical genealogies (detailed for
example in the Ussher chronology). Young Earth creationists often believe that the Universe has a similar
age to the Earth's. Creationist cosmologies are attempts by some creationist thinkers to give the universe an
age consistent with the Ussher chronology and other Young-Earth timeframes. This belief generally has a
basis in a literal and inerrant interpretation of the Bible.

[edit] Old Earth creationism

Main article: Old Earth creationism
See also: Gap creationism, Day-Age Creationism, and Progressive creationism
Old Earth creationism holds that the physical universe was created by God, but that the creation
event of Genesis is not to be taken strictly literally. This group generally believes that the age of the Universe
and the age of the Earth are as described by astronomers and geologists, but that details of the evolutionary
theory are questionable. Old Earth creationists interpret the Genesis creation myth in a number of ways, that
each differ from the six, consecutive, 24-hour day creation of the Young Earth Creationist view.
[edit] Neo-Creationism
Main article: Neo-Creationism
See also: Intelligent design
Neo-Creationists intentionally distance themselves from other forms of creationism, preferring to be
known as wholly separate from creationism as a philosophy. Their goal is to restate creationism in terms
more likely to be well received by the public, education policy makers and the scientific community. It aims to
re-frame the debate over the origins of life in non-religious terms and without appeals to scripture, and to
bring the debate before the public. Neo-creationists may be either Young Earth or Old Earth Creationists, and
hold a range of underlying theological viewpoints (e.g. on the interpretation of the Bible). Neo-Creationism
currently exists in the form of the intelligent design movement, which has a 'big tent' strategy making it
inclusive of many Young Earth Creationists (such as Paul Nelson and Percival Davis).

[edit] Theistic evolution

Main article: Theistic evolution
See also: Naturalism (philosophy), Evolution and the Roman Catholic Church, and Clergy Letter
Project
Theistic evolution is the general view that, instead of faith being in opposition to biological evolution,
some or all classical religious teachings about God and creation are compatible with some or all of modern
scientific theory, including, specifically, evolution. It generally views evolution as a tool used by a creator god,
who is both the first cause and immanent sustainer/upholder of the universe; it is therefore well accepted by
people of strong theistic (as opposed to deistic) convictions. Theistic evolution can synthesize with the day-
age interpretation of the Genesis creation myth; however most adherents consider that the first chapters of
Genesis should not be interpreted as a "literal" description, but rather as a literary framework or allegory.
This position does not generally exclude the viewpoint of methodological naturalism, a long standing
convention of the scientific method in science.
Theistic evolutionists have frequently been prominent in opposing creationism (including intelligent
design). Notable examples have been biologist Kenneth R. Miller and theologian John Haught, who testified
for the plaintiffs in Kitzmiller v. Dover Area School District. Another example is the Clergy Letter Project, an
organization that has created and maintains a statement signed by American Christian clergy of different
denominations rejecting creationism, with specific reference to points raised by intelligent design proponents.
Theistic evolutionists have also been active in Citizens Alliances for Science that oppose the introduction of
creationism into public school science classes (one example being evangelical Christian geologist Keith B.
Miller, who is a prominent board member of Kansas Citizens for Science).

[edit] Naturalistic evolution

Naturalistic evolution is the position of acceptance of biological evolution and of metaphysical
naturalism (and thus rejection of theism and theistic evolution). A prominent proponent of this viewpoint is
British evolutionary biologist Richard Dawkins.

This section requires expansion.

[edit] Arguments relating to the definition and limits of science
Critiques such as those based on the distinction between theory and fact are often leveled against
unifying concepts within scientific disciplines. Principles such as uniformitarianism, Occam's Razor or
parsimony, and the Copernican principle are claimed to be the result of a bias within science toward
philosophical naturalism, which is equated by many creationists with atheism.[56] In countering this claim,
philosophers of science use the term methodological naturalism to refer to the long standing convention in
science of the scientific method. The methodological assumption is that observable events in nature are
explained only by natural causes, without assuming the existence or non-existence of the supernatural, and
therefore supernatural explanations for such events are outside the realm of science.[57] Creationists claim
that supernatural explanations should not be excluded and that scientific work is paradigmatically close-
minded.[58]
Because modern science tries to rely on the minimization of a priori assumptions, error, and
subjectivity, as well as on avoidance of Baconian idols, it remains neutral on subjective subjects such as
religion or morality.[59] Mainstream proponents accuse the creationists of conflating the two in a form of
pseudoscience.[60]

[edit] Definitions
Fact: In science, an observation that has been repeatedly confirmed and for all practical
purposes is accepted as "true." Truth in science, however, is never final, and what is accepted
as a fact today may be modified or even discarded tomorrow.
 Hypothesis: A tentative statement about the natural world leading to deductions that can be
tested. If the deductions are verified, it becomes more probable that the hypothesis is correct. If
the deductions are incorrect, the original hypothesis can be abandoned or modified. Hypotheses
can be used to build more complex inferences and explanations.

Law: A descriptive generalization about how some aspect of the natural world behaves under
stated circumstances.

Theory: In science, a well-substantiated explanation of some aspect of the natural world that can
incorporate facts, laws, inferences, and tested hypotheses.

—National Academy of Sciences, Science and Creationism[61]

[edit] Limitations of the scientific endeavor

In science, explanations are limited to those based

on observations and experiments that can be substantiated by other scientists. Explanations

that cannot be based on empirical evidence are not a part of science.

—National Academy of Sciences, Science and Creationism[61]

 This section requires expansion.

[edit] Theory vs. fact

Main article: Evolution as theory and fact
The argument that evolution is a theory, not a fact, has often been made against the exclusive
teaching of evolution.[62] The argument is related to a common misconception about the technical meaning
of "theory" that is used by scientists. In common usage, "theory" often refers to conjectures, hypotheses, and
unproven assumptions. However, in science, "theory" usually means "a plausible or scientifically acceptable
general principle or body of principles offered to explain phenomena."[63]
Exploring this issue, paleontologist Stephen Jay Gould wrote:
Evolution is a theory. It is also a fact. And facts and theories are different things, not rungs in a
hierarchy of increasing certainty. Facts are the world's data. Theories are structures of ideas that
explain and interpret facts. Facts do not go away when scientists debate rival theories to explain
them. Einstein's theory of gravitation replaced Newton's, but apples did not suspend themselves
in mid-air, pending the outcome. And humans evolved from ape-like ancestors whether they did
so by Darwin's proposed mechanism or by some other yet to be discovered.
—Stephen Jay Gould, Evolution as Fact and Theory[64]
[edit] Falsifiability
Philosopher of science Karl R. Popper set out the concept of falsifiability as a way to distinguish
science and pseudoscience: Testable theories are scientific, but those that are untestable are not.[65]
However, in Unended Quest, Popper declared "I have come to the conclusion that Darwinism is not a
testable scientific theory but a metaphysical research programme, a possible framework for testable scientific
theories," while pointing out it had "scientific character".[66]
In what one sociologist derisively called "Popper-chopping,"[67] opponents of evolution seized upon
Popper's definition to claim evolution was not a science, and claimed creationism was an equally valid
metaphysical research program.[68] For example, Duane Gish, a leading Creationist proponent, wrote in a
letter to Discover magazine (July 1981): "Stephen Jay Gould states that creationists claim creation is a
scientific theory. While many Creationists claim creation is a scientific theory other Creationists have stated
that neither creation nor evolution is a scientific theory (and each is equally religious)."[69]
Popper responded to news that his conclusions were being used by anti-evolutionary forces by
affirming that evolutionary theories regarding the origins of life on earth were scientific because "their
hypotheses can in many cases be tested."[70] However, creationists claimed that a key evolutionary concept,
that all life on Earth is descended from a single common ancestor, was not mentioned as testable by Popper,
and claimed it never would be.[71]
In fact, Popper wrote admiringly of the value of Darwin's theory.[72] Only a few years later, Popper
wrote, "I have in the past described the theory as "almost tautological"...I still believe that natural selection
works in this way as a research programme. Nevertheless, I have changed my mind about the testability and
logical status of the theory of natural selection; and I am glad to have an opportunity to make a recantation".
His conclusion, later in the article is "The theory of natural selection may be so formulated that it is far from
tautological. In this case it is not only testable, but it turns out to be not strictly universally true."[73]
Debate among some scientists and philosophers of science on the applicability of falsifiability in
science continues.[74] However, simple falsifiability tests for common descent have been offered by some
scientists: For instance, biologist and prominent critic of creationism Richard Dawkins and J.B.S. Haldane
both pointed out that if fossil rabbits were found in the Precambrian era, a time before most similarly complex
lifeforms had evolved, "that would completely blow evolution out of the water."[75][76]
Falsifiability has also caused problems for creationists: In his 1982 decision McLean v. Arkansas
Board of Education, Judge William R. Overton used falsifiability as one basis for his ruling against the
teaching of creation science in the public schools, ultimately declaring it "simply not science."[77]

[edit] Conflation of science and religion

See also: Objection to evolution that it is a religion

[edit] Appeal to consequences

See also: Objection to evolution's moral implications
A number of creationists have blurred the boundaries between their disputes over the truth of the
underlying facts, and explanatory theories, of evolution, with their purported philosophical and moral
consequences. This type of argument is known as an appeal to consequences, and is a logical fallacy.
Examples of these arguments include those of prominent creationists such as Ken Ham[78] and Henry M.
Morris.[79]
[edit] Disputes relating to science
Many creationists strongly oppose certain scientific theories in a number of ways, including
opposition to specific applications of scientific processes, accusations of bias within the scientific community,
[80] and claims that discussions within the scientific community reveal or imply a crisis. In response to
perceived crises in modern science, creationists claim to have an alternative, typically based on faith,
creation science, and/or intelligent design. The scientific community has responded by pointing out that their
conversations are frequently misrepresented (e.g. by quote mining) in order to create the impression of a
deeper controversy or crisis, and that the creationists' alternatives are generally pseudoscientific.
[edit] Biology

A phylogenetic tree based on rRNA genes.

Disputes relating to evolutionary biology are central to the controversy between Creationists and the scientific
community. The aspects of evolutionary biology disputed include common descent (and particularly human
evolution from common ancestors with other members of the Great Apes), macroevolution, and the existence
of transitional fossils.

[edit] Common descent

Main article: Common descent
See also: Evidence of common descent and Tree of life (science)
[The] Discovery [Institute] presents common descent as controversial exclusively within the
animal kingdom, as it focuses on embryology, anatomy, and the fossil record to raise questions
about them. In the real world of science, common descent of animals is completely
noncontroversial; any controversy resides in the microbial world. There, researchers argued
over a variety of topics, starting with the very beginning, namely the relationship among the
three main branches of life.
—John Timmer, Evolution: what's the real controversy?[81]

A group of organisms is said to have common descent if they have a common ancestor. A theory of
universal common descent based on evolutionary principles was proposed by Charles Darwin and is now
generally accepted by biologists. The last universal common ancestor, that is, the most recent common
ancestor of all currently living organisms, is believed to have appeared about 3.9 billion years ago.
 With a few exceptions (e.g. Michael Behe), the vast majority of Creationists reject this theory.[82][83]
[84]
Evidence of common descent includes evidence from genetics, fossil records, comparative anatomy,
geographical distribution of species, comparative physiology and comparative biochemistry[citation needed].

[edit] Human evolution

Main article: Human evolution
See also: Paleoanthropology and Adam and Eve
Human evolution is the study of the biological evolution of humans as a distinct species from its
common ancestors with other animals. Analysis of fossil evidence and genetic distance are two of the means
by which scientists understand this evolutionary history.
Fossil evidence suggests that humans' earliest hominoid ancestors may have split from other
primates as early as the late Oligocene, circa 26-24 Ma, and that by the early Miocene, the adaptive radiation
of many different hominoid forms was well underway.[85] Evidence from the molecular dating of genetic
differences indicates that the gibbon lineage (family Hylobatidae) diverged between 18 and 12 Ma, and the
orangutan lineage (subfamily Ponginae) diverged about 12 Ma. While there is no fossil evidence thus far
clearly documenting the early ancestry of gibbons, fossil proto-orangutans may be represented by
Sivapithecus from India and Griphopithecus from Turkey, dated to around 10 Ma. Molecular evidence further
suggests that between 8 and 4 Ma, first the gorillas, and then the chimpanzee (genus Pan) split from the line
leading to the humans.[86] We have no fossil record of this divergence, but distinctively hominid fossils have
been found dating to 3.2 Ma (see Lucy) and possibly even earlier, at 6 or 7 Ma (see Toumaï).[87]
Comparisons of chimpanzee and human DNA show the two are approximately 98.4 percent identical, and
are taken as strong evidence of recent common ancestry.[88] Today, only one distinct human species
survives, but many earlier species have been found in the fossil record, including Homo erectus, Homo
habilis, and Homo neanderthalensis.
Creationists dispute there is evidence of shared ancestry in the fossil evidence, and argue either that
these are misassigned ape fossils (e.g. that Java man was a gibbon[89]) or too similar to modern humans to
designate them as distinct or transitional forms.[90] However Creationists frequently disagree where the
dividing lines would be.[91] Creation myths (such as the Book of Genesis) frequently posit a first man (Adam,
in the case of Genesis) as an alternative viewpoint to the scientific account.
Creationists also dispute science's interpretation of genetic evidence in the study of human evolution.
They argue that it is a "dubious assumption" that genetic similarities between various animals imply a
common ancestral relationship, and that scientists are coming to this interpretation only because they have
preconceived notions that such shared relationships exist. Creationists also argue that genetic mutations are
strong evidence against evolutionary theory because the mutations required for major changes to occur
would almost certainly be detrimental.[31]

[edit] Macroevolution
Main article: Macroevolution
See also: Speciation
Many creationists now accept the possibilities of microevolution within "kinds" but refuse to accept
and have long argued against the possibility of macroevolution. Macroevolution is defined by the scientific
community to be evolution that occurs at or above the level of species. Under this definition, macroevolution
can be considered to be a fact, as evidenced by observed instances of speciation. Creationists however tend
to apply a more restrictive, if vaguer, definition of macroevolution, often relating to the emergence of new
body forms or organs. The scientific community considers that there is strong evidence for even such more
restrictive definitions, but the evidence for this is more complex.
Recent arguments against (such restrictive definitions of) macroevolution include the intelligent
design arguments of irreducible complexity and specified complexity. However, neither argument has been
accepted for publication in a peer-reviewed scientific journal, and both arguments have been rejected by the
scientific community as pseudoscience.

This section requires expansion.

[edit] Transitional fossils

Main article: Transitional fossil
See also: List of transitional fossils, Bird evolution, and Evolution of the horse
It is commonly stated by critics of evolution that there are no known transitional fossils.[92][93] This
position is based on a misunderstanding of the nature of what represents a transitional feature. A common
creationist argument is that no fossils are found with partially functional features. It is plausible, however, that
a complex feature with one function can adapt a wholly different function through evolution. The precursor to,
for example, a wing, might originally have only been meant for gliding, trapping flying prey, and/or mating
display. Nowadays, wings can still have all of these functions, but they are also used in active flight.
 Reconstruction of Ambulocetus natans
As another example, Alan Haywood stated in Creation and Evolution that "Darwinists rarely mention
the whale because it presents them with one of their most insoluble problems. They believe that somehow a
whale must have evolved from an ordinary land-dwelling animal, which took to the sea and lost its legs ... A
land mammal that was in the process of becoming a whale would fall between two stools—it would not be
fitted for life on land or at sea, and would have no hope for survival."[94] The evolution of whales has
however been documented in considerable detail, with Ambulocetus, described as looking like a three-metre
long mammalian crocodile, as one of the transitional fossils.
Although transitional fossils elucidate the evolutionary transition of one life-form to another, they only
exemplify snapshots of this process. Due to the special circumstances required for preservation of living
beings, only a very small percentage of all life-forms that ever have existed can be expected to be
discovered. Thus, the transition itself can only be illustrated and corroborated by transitional fossils, but it will
never be known in detail. However, progressing research and discovery managed to fill in several gaps and
continues to do so. Critics of evolution often cite this argument as being a convenient way to explain off the
lack of 'snapshot' fossils that show crucial steps between species.
 The theory of punctuated equilibrium developed by Stephen Jay Gould and Niles Eldredge is often
mistakenly drawn into the discussion of transitional fossils. This theory, however, pertains only to well-
documented transitions within taxa or between closely related taxa over a geologically short period of time.
These transitions, usually traceable in the same geological outcrop, often show small jumps in morphology
between periods of morphological stability. To explain these jumps, Gould and Eldredge envisaged
comparatively long periods of genetic stability separated by periods of rapid evolution. For example the
change from a creature the size of a mouse, to one the size of an elephant, could be accomplished over
60,000 years, with a rate of change too small to be noticed over any human lifetime. 60,000 years is too
small a gap to be identified or identifiable in the fossil record.[95]
Furthermore, promoters of evolution (such as Richard Dawkins) have pointed out that, even if it were
possible for enough fossils to survive to show a close transitional change, evolution's critics will never be
satisfied, as the discovery of one "missing link" itself creates two more so-called "missing links" on either side
of the discovery. Dawkins goes on to say that the reason for this "losing battle" is that many of these critics
are theists who "simply don't want to see the truth".

[edit] Geology
Main article: Flood Geology
See also: Creation geophysics, Geochronology, and Age of the Earth
Many believers in Young Earth Creationism – a position held by the majority of proponents of Flood
Geology – accept biblical chronogenealogies (such as the Ussher chronology which in turn is based on the
Masoretic version of the Genealogies of Genesis).[96][97] They believe that God created the universe
approximately 6000 years ago, in the space of six days. Much of creation geology is devoted to debunking
the dating methods used in anthropology, geology, and planetary science that give ages in conflict with the
young Earth idea. In particular, creationists dispute the reliability of radiometric dating and isochron analysis,
both of which are central to mainstream geological theories of the age of the Earth. They usually dispute
these methods based on uncertainties concerning initial concentrations of individually considered species
and the associated measurement uncertainties caused by diffusion of the parent and daughter isotopes.
However, a full critique of the entire parameter-fitting analysis, which relies on dozens of radionuclei parent
and daughter pairs, has not been done by creationists hoping to cast doubt on the technique.
The consensus of professional scientific organisations worldwide is that no scientific evidence
contradicts the age of approximately 4.5 billion years.[98] Young Earth creationists reject these ages on the
grounds of what they regard as being tenuous and untestable assumptions in the methodology. Apparently
inconsistent radiometric dates are often quoted to cast doubt on the utility and accuracy of the method.
Mainstream proponents who get involved in this debate point out that dating methods only rely on the
assumptions that the physical laws governing radioactive decay have not been violated since the sample was
formed (harking back to Lyell's doctrine of uniformitarianism). They also point out that the "problems" that
creationists publicly mentioned can be shown to either not be problems at all, are issues with known
contamination, or simply the result of incorrectly evaluating legitimate data.

[edit] Other sciences

[edit] Cosmology
See also: Age of the universe
Whilst Young Earth Creationists believe that the Universe was created by the Judeo-Chrisitian God
approximately 6000 years ago, the current scientific consensus is that the Universe as we know it emerged
from the Big Bang 13.7 billion years ago. The recent science of nucleocosmochronology is extending the
approaches used for Carbon-14 dating to the dating of astronomical features. For example based upon this
emerging science, the Galactic thin disk of the Milky Way galaxy is estimated to have been formed 8.3 ± 1.8
billion years ago.[99]
Old Earth Creationists and some others do not necessarily dispute these figures.

[edit] Nuclear physics

See also: radiometric dating
Creationists point to experiments they have performed, which they claim demonstrate that 1.5 billion
years of nuclear decay took place over a short period of time, from which they infer that "billion-fold speed-
ups of nuclear decay" have occurred, a massive violation of the principle that radioisotope decay rates are
constant, a core principle underlying nuclear physics generally, and radiometric dating in particular.[100]
The scientific community points to numerous flaws in these experiments, to the fact that their results
have not been accepted for publication by any peer-reviewed scientific journal, and to the fact that the
creationist scientists conducting them were untrained in experimental geochronology.[101][102]
In refutation of young-Earth claims of inconstant decay rates affecting the reliability of radiometric
dating, Roger C. Wiens, a physicist specialising in isotope dating states:
There are only three quite technical instances where a half-life changes, and these do not affect
the dating methods [under discussion]":[103]
 1. Only one technical exception occurs under terrestrial conditions, and this is not for an
isotope used for dating. ... The artificially-produced isotope, beryllium-7 has been shown
to change by up to 1.5%, depending on its chemical environment. ... [H]eavier atoms are
even less subject to these minute changes, so the dates of rocks made by electron-
capture decays would only be off by at most a few hundredths of a percent.
2. ... Another case is material inside of stars, which is in a plasma state where electrons
are not bound to atoms. In the extremely hot stellar environment, a completely different
kind of decay can occur. 'Bound-state beta decay' occurs when the nucleus emits an
electron into a bound electronic state close to the nucleus. ... All normal matter, such as
everything on Earth, the Moon, meteorites, etc. has electrons in normal positions, so
these instances never apply to rocks, or anything colder than several hundred thousand
degrees. ...
3. The last case also involves very fast-moving matter. It has been demonstrated by
atomic clocks in very fast spacecraft. These atomic clocks slow down very slightly (only
a second or so per year) as predicted by Einstein's theory of relativity. No rocks in our
solar system are going fast enough to make a noticeable change in their dates. ...

—Roger C. Wiens, Radiometric Dating, A Christian Perspective[104]

[edit] Misrepresentations of science

[edit] Quote mining

Main article: Quote mining
 As a means to criticise mainstream science, creationists have been known to quote, at length,
scientists who ostensibly support the mainstream theories, but appear to acknowledge criticisms similar to
those of creationists.[105] However, almost universally these have been shown to be quote mines that do not
accurately reflect the evidence for evolution or the mainstream scientific community's opinion of it, or highly
out-of-date.[106][107] Many of the same quotes used by creationists have appeared so frequently in Internet
discussions due to the availability of cut and paste functions, that the TalkOrigins Archive has created "The
Quote Mine Project" for quick reference to the original context of these quotations.[106]

[edit] Public policy issues

[edit] Science education
Main article: Creation and evolution in public education
See also: Teach the Controversy
Creationists promote that evolution is a theory in crisis[108][109] with scientists criticizing
evolution[110] and claim that fairness and equal time requires educating students about the alleged scientific
controversy.
Opponents, being the overwhelming majority of the scientific community and science education
organizations,[111] reply that there is no scientific controversy and that the controversy exists solely in terms
of religion and politics.[108][110] The American Association for the Advancement of Science and other
science and education professional organizations say that Teach the Controversy proponents seek to
undermine the teaching of evolution[108][112] while promoting intelligent design,[113][114][115] and to
advance an education policy for US public schools that introduces creationist explanations for the origin of
life to public-school science curricula.[116][117] This viewpoint was supported by the December 2005 ruling
in the Kitzmiller v. Dover Area School District trial.[117]
George Mason University Biology Department introduced a course on the creation/evolution
controversy, and apparently as students learn more about biology, they find objections to evolution less
convincing, suggesting that “teaching the controversy” rightly as a separate elective course on philosophy or
history of science, or "politics of science and religion," would undermine creationists’ criticisms, and that the
scientific community’s resistance to this approach was bad public relations.[118]
On March 27, 2009, the Texas Board of Education, by a vote of 13 to 2, voted that at least in Texas,
textbooks must teach intelligent design alongside evolution, and question the validity of the fossil record. Don
McLeroy, a dentist and chair of the board, said, "I think the new standards are wonderful ... dogmatism about
evolution" has sapped "America's scientific soul." According to Science magazine, "Because Texas is the
second-largest textbook market in the United States, publishers have a strong incentive to be certified by the
board as 'conforming 100% to the state's standards'."[119]

[edit] Freedom of speech

Creationists have claimed that preventing them from teaching Creationism violates their right of
Freedom of speech. However court cases (such as Webster v. New Lenox School District and Bishop v.
Aronov) have upheld school districts' and universities' right to restrict teaching to a specified curriculum.

[edit] Issues relating to religion

See also: Relationship between religion and science and Evolution and the Roman Catholic Church
[edit] Theological arguments
See also: Allegorical interpretations of Genesis and Evolutionary argument against naturalism
This section requires expansion.

[edit] Religion and historical scientists

Creationists often argue that Christianity and literal belief in the Bible are either foundationally
significant or directly responsible for scientific progress.[120] To that end, Institute for Creation Research
founder Henry M. Morris has enumerated scientists such as astronomer and philosopher Galileo,
mathematician and theoretical physicist James Clerk Maxwell, mathematician and philosopher Blaise Pascal,
geneticist monk Gregor Mendel, and Isaac Newton as believers in a biblical creation narrative.[121]
This argument usually involves scientists either who were no longer alive when evolution was
proposed or whose field of study didn't include evolution. The argument is generally rejected as specious by
those who oppose creationism.[122]
Many of the scientists in question did some early work on the mechanisms of evolution, e.g., the
Modern evolutionary synthesis combines Darwin's Evolution with Mendel's theories of inheritance and
genetics. Though biological evolution of some sort had become the primary mode of discussing speciation
within science by the late-19th century, it was not until the mid-20th century that evolutionary theories
stabilized into the modern synthesis. Some of the historical scientists marshalled by creationists were dealing
with quite different issues than any are engaged with today: Louis Pasteur, for example, opposed the theory
of spontaneous generation with biogenesis, an advocacy some creationists describe as a critique on
chemical evolution and abiogenesis. Pasteur accepted that some form of evolution had occurred and that the
Earth was millions of years old.[123]
The relationship between science and religion was not portrayed in antagonistic terms until the late-
19th century, and even then there have been many examples of the two being reconcilable for evolutionary
scientists.[124] Many historical scientists wrote books explaining how pursuit of science was seen by them as
fulfillment of spiritual duty in line with their religious beliefs. Even so, such professions of faith were not
insurance against dogmatic opposition by certain religious people.
Some extensions to this creationist argument have included the incorrect suggestions that Einstein's
deism was a tacit endorsement of creationism or that Charles Darwin converted on his deathbed and
recanted evolutionary theory.

[edit] Forums for the controversy

[edit] Debates
Many creationists and scientists engage in frequent public debates regarding the origin of human life,
hosted by a variety of institutions. However, some scientists disagree with this tactic, arguing that by openly
debating supporters of supernatural origin explanations (creationism and intelligent design), scientists are
lending credibility and unwarranted publicity to creationists, which could foster an inaccurate public
perception and obscure the factual merits of the debate.[125] For example, in May 2004 Dr. Michael Shermer
debated creationist Kent Hovind in front of a predominantly creationist audience. In Shermer's online
reflection while he was explaining that he won the debate with intellectual and scientific evidence he felt it
was "not an intellectual exercise," but rather it was "an emotional drama", with scientists arguing from "an
impregnable fortress of evidence that converges to an unmistakable conclusion", whilst for creationists it is "a
spiritual war".[126] While receiving positive responses from creationist observers, Shermer concluded
"Unless there is a subject that is truly debatable (evolution v. creation is not), with a format that is fair, in a
forum that is balanced, it only serves to belittle both the magisterium of science and the magisterium of
religion."[126] (see: scientific method). Others, like evolutionary biologist Massimo Pigliucci, have debated
Hovind, and have expressed surprise to hear Hovind try "to convince the audience that evolutionists believe
humans came from rocks" and at Hovind's assertion that biologists believe humans "evolved from
bananas."[127]
Eugenie Scott of the National Center for Science Education, a non-profit organization dedicated to
defending the teaching of evolution in the public schools, claimed debates are not the sort of arena to
promote science to creationists.[126] Scott says that "Evolution is not on trial in the world of science," and
"the topic of the discussion should not be the scientific legitimacy of evolution" but rather should be on the
lack of evidence in creationism. Stephen Jay Gould adopted a similar position, explaining:
Debate is an art form. It is about the winning of arguments. It is not about the discovery of truth.
There are certain rules and procedures to debate that really have nothing to do with establishing
fact — which [creationists] are very good at. Some of those rules are: never say anything positive
about your own position because it can be attacked, but chip away at what appear to be the
weaknesses in your opponent's position. They are good at that. I don't think I could beat the
creationists at debate. I can tie them. But in courtrooms they are terrible, because in courtrooms
you cannot give speeches. In a courtroom you have to answer direct questions about the
positive status of your belief.
—Stephen Jay Gould, lecture 1985[128]
[edit] Political lobbying
See also: Politics of creationism, Kansas evolution hearings, Santorum Amendment, and List of
scientific societies rejecting intelligent design
A wide range of organisations, on both sides of the controversy, are involved in lobbying in an
attempt to influence political decisions relating to the teaching of evolution, at a number of levels. These
include the Discovery Institute, the National Center for Science Education, the National Science Teachers
Association, state Citizens Alliances for Science, and numerous national science associations and state
Academies of Science.[129]

This section requires expansion.

[edit] In the media

The controversy has been discussed in numerous newspaper articles, reports, op-eds and letters to
the editor, as well as a number of radio and television programmes (including the PBS series, Evolution and
Coral Ridge Ministries' Darwin's Deadly Legacy). This has led some commentators to express a concern at
what they see as a highly inaccurate and biased understanding of evolution among the general public.
Edward Humes states:
There are really two theories of evolution. There is the genuine scientific theory and there is the
talk-radio pretend version, designed not to enlighten but to deceive and enrage.
The talk-radio version had a packed town hall up in arms at the Why Evolution Is Stupid lecture.
In this version of the theory, scientists supposedly believe that all life is accidental, a random
crash of molecules that magically produced flowers, horses and humans – a scenario as unlikely
as a tornado in a junkyard assembling a 747. Humans come from monkeys in this theory, just
popping into existence one day. The evidence against Darwin is overwhelming, the purveyors of
talk-radio evolution rail, yet scientists embrace his ideas because they want to promote atheism.

—Edward Humes, Unintelligent Designs on Darwin[130]

[edit] Outside the United States
 Views on human evolution in various countries.[131][132]
While the controversy has been prominent in the United States, it has flared up in other countries as
well.[133][134][135]

[edit] Europe
Europeans have often regarded the creation-evolution controversy as an American matter.[134]
However, in recent years the conflict has become an issue in a variety of countries including Germany, the
United Kingdom, Italy, the Netherlands, Poland, and Serbia.[134][135][136][137]
On 17 September 2007 the Committee on Culture, Science and Education of the Parliamentary
Assembly of the Council of Europe issued a report on the attempt by American-inspired creationists to
promote creationism in European schools. It concludes "If we are not careful, creationism could become a
threat to human rights which are a key concern of the Council of Europe.... The war on the theory of evolution
and on its proponents most often originates in forms of religious extremism which are closely allied to
extreme right-wing political movements... some advocates of creationism are out to replace democracy by
theocracy."[138]

[edit] Australia
With declining church attendance, there has been some growth in fundamentalist and Pentecostal
Christian denominations.[139] Under the former Queensland state government of Joh Bjelke-Petersen, in
1980 lobbying was so successful that Queensland allowed the teaching of creationism as science to school
children[citation needed]. In 2010 the Queensland state government introduced the topic of creationism into
school classes within the "ancient history" subject where its origins and nature are discussed as a significant
controversy.[140] Public lectures have been given in rented rooms at Universities, by visiting American
speakers, and speakers with doctorates purchased by mail from Florida sites.[141] One of the most
acrimonious aspects of the Australian debate was featured on the science television program Quantum,
about a long-running and ultimately unsuccessful court case by Ian Plimer, Professor of Geology at
Melbourne University, against an ordained minister, Dr. Allen Roberts, who had claimed that there were
remnants of Noah's Ark in eastern Turkey. Although the court found that Dr Roberts had made false and
misleading claims, they were not made in the course of trade or commerce, so the case failed.[142]

[edit] Islamic countries

See also: Islamic creationism
In recent times, the controversy has become more prominent in Islamic countries.[143] Currently, in
Egypt evolution is taught in schools but Saudi Arabia and Sudan have both banned the teaching of evolution
in schools.[133] Creation science has also been heavily promoted in Turkey and in immigrant communities in
Western Europe, primarily by Harun Yahya.[135]

[edit] See also

Science portal

Main article: Outline of the creation-evolution controversy

Book:Evolution
Books are collections of articles that can be downloaded or ordered in print.
• Abiogenesis
• Allegorical interpretations of Genesis
• Anti-intellectualism
• Biogenesis
• Clergy Letter Project
• Creation science
• Creationism
• Evidence of common descent
• Evolution and the Roman Catholic Church
• Evolution Sunday
• Evolutionary origin of religions
• Hindu views on evolution
• History of the creation-evolution controversy
• Intelligent design
• Jainism and non-creationism
• Jewish views on evolution
• Level of support for evolution
• List of participants in the creation-evolution controversy
• Mormonism and evolution
• Natural theology
• Objections to evolution
 • Politics of creationism
• Project Steve
• Relationship between religion and science
• Teach the Controversy

[edit] Notes
[note 1]
1. ^ While the term myth is often used colloquially to refer to "a false story", this article uses the
term "creation myth" in the formal sense, common in academic literature, meaning "a religious or
supernatural story or explanation that describes the beginnings of humanity, earth, life, and the
universe."

[edit] Footnotes
1. ^ See Hovind 2006, for example.
2. ^ An Index to Creationist Claims , Mark Isaak, Talkorigins Archive,Copyright © 2006.
3. ^ Curry, Andrew (27 February 2009). "Creationist Beliefs Persist in Europe". Science 323
(5918): 1159. doi:10.1126/science.323.5918.1159. PMID 19251601. "News coverage of the
creationism-versus-evolution debate tends to focus on the United States ... But in the past 5 years,
political clashes over the issue have also occurred in countries all across Europe. ... "This isn't just
an American problem," says Dittmar Graf of the Technical University of Dortmund, who organized the
meeting".
 4. ^ Larson 2004, pp. 247-263 Chapter titled Modern Culture Wars. See also Ruse 1999, p. 26,
who writes "One thing that historians delighted in showing is that, contrary to the usually held tale of
science and religion being always opposed...religion and theologically inclined philosophy have
frequently been very significant factors in the forward movement of science."
5. ^ Numbers 1992, p. 3-240
6. ^ See:
• Peters & Hewlett 2005, p. 1;
• Kitzmiller v. Dover Area School District, page 20
7. ^ See:
• Battle on Teaching Evolution Sharpens, Peter Slevin, Washington Post, Monday, 14
March 2005, Page A01;
• The Political Design of Intelligent Design , Russell D. Renka, 16 November 2005;
• Politicized Scholars Put Evolution on the Defensive , Jody Wilgoren, The New York
Times, 21 April 2005
• The Newest Evolution of Creationism, Barbara Forrest, Natural History, April, 2002,
page 80;
• Kitzmiller vs. Dover Area School District, pages 7-9, also pages 64-90
8. ^ Myers 2006; NSTA 2007; IAP 2006; AAAS 2006; and Pinholster 2006; Ruling, Kitzmiller v.
Dover page 83
9. ^ Larson 2004, p. 258 "Virtually no secular scientists accepted the doctrines of creation
science; but that did not deter creation scientists from advancing scientific arguments for their
position." See also Martz & McDaniel 1987, p. 23, a Newsweek article which states "By one count
there are some 700 scientists (out of a total of 480,000 U.S. earth and life scientists) who give
credence to creation-science, the general theory that complex life forms did not evolve but appeared
'abruptly'."
10.^ Committee on Revising Science and Creationism, A View from the National Academy of
Sciences, National Academy of Sciences and Institute of Medicine of the National Academies (2008).
Science, Evolution, and Creationism. National Academy of Sciences. p. 12. ISBN 0-309-10586-2.
http://www.nap.edu/catalog.php?record_id=11876.
11.^ Desmond & Moore 1991, p. 34-35
12.^ See":
• van Wyhe 2006;
• Desmond & Moore 1991, p. 321-323, 503-505.
13.^ van Wyhe 2006
14.^ a b c d AAAS Evolution Dialogues: Science, Ethics and Religion study guide (pdf)
15.^ See:
• Hodge 1874, p. 177;
• Numbers 1992, p. 14;
• Burns, Ralph, Lerner, & Standish 1982, p. 965;
• Huxley 1902
16.^ a b Witham 2002
17.^ Barbour 1997
18.^ Numbers 1992, p. 14
19.^ Numbers 1992, p. 14-15
20.^ a b Numbers 1992, p. 17
 21.^ Numbers 1992, p. 18, noting that this applies to published or public skeptics. Many or most
Christians may have held on to a literal six days of creation, but these views were rarely expressed in
books and journals. Exceptions are also noted, such as literal interpretations published by Eleazar
Lord (1788-1871) and David Nevins Lord (1792-1880). However, the observation that evolutionary
critics had a relaxed interpretation of Genesis is supported by specifically enumerating: Louis
Agassiz (1807-1873); Arnold Henry Guyot (1807-1884); John William Dawson (1820-1899); Enoch
Fitch Burr (1818-1907); George D. Armstrong (1813-1899); Charles Hodge, theologian (1797-1878);
James Dwight Dana (1813-1895); Edward Hitchcock, clergyman and respected Amherst College
geologist, (1793-1864); Reverend Herbert W. Morris (1818-1897); H. L. Hastings (1833?-1899);
Luther T. Townsend (1838-1922; Alexander Patterson, Presbyterian evangelist who published The
Other Side of Evolution Its Effects and Fallacy
22.^ Numbers(2006) p161
23.^ Similar legislation was passed in two other states prior to the Scopes trial, in Oklahoma and
Florida. The efforts to enact "Butler Acts" in other jurisdictions were abandoned after the Scopes trial.
See HistoryNet (describing the Florida and Oklahoma acts and Scientific American Article.
24.^ The statute required a minimum fine of $100, and the state Constitution required all fines
over $50 to be assessed by a jury. Court's opinion.
25.^ In his closing argument, Darrow told the jury that they essentially had no choice but to
convict Scopes on the evidence, and he did not hold that against them. The jury was out for a total of
nine minutes (including time for egress and ingress). The jury informed the Court that they "passed"
the question of the sentence to the Court, a move which the judge accepted despite the
Constitutional provision establishing a $50 limit on judge-imposed fines.
26.^ The Establishment Clause of the First Amendment of the United States Constitution was
not, at the time of the Scopes decision in the 1920s, deemed applicable to the states. Thus, Scopes'
constitutional defense on establishment grounds rested solely on the state constitution. See Court's
opinion. See generally Incorporation doctrine and Everson v. Board of Education (seminal U.S.
Supreme Court opinion finally applying the Establishment Clause against states in 1947). See also
Book Review of Professor Larson's Pulitzer Prize winning Summer for the Gods at point 4 ("The
constitutional case was largely based on state constitutional law; this was before most of the Bill of
Rights had been incorporated and applied to the states."); Cantwell v. Connecticut (1940 Supreme
Court case stating that the Free Exercise Clause of the First Amendment is incorporated into the
Fourteenth Amendment and is therefore applicable against the states); Cookson, Catharine (2003).
Encyclopedia of religious freedom. Taylor & Francis. pp. 132 et seq.. ISBN 0415941814. (explaining
incorporation doctrine relative to First Amendment); Incorporation Doctrine Explained by
Constitutional Rights Foundation.
27.^ The Court accordingly did not address the question of whether the teaching of Creationism
in the public schools was unconstitutional.
28.^ The Court stated in its opinion that "England and Scotland maintained State churches as
did some of the Colonies, and it was intended by this clause of the Constitution [the Religious
Preference Clause] to prevent any such undertaking in Tennessee."
29.^ See:
• s:Kitzmiller v. Dover Area School District/2:Context#Page 19 of 139 ;
• Understanding the Intelligent Design Creationist Movement: Its True Nature and
Goals. (pdf) A Position Paper from the Center for Inquiry, Office of Public Policy Barbara
Forrest. May, 2007;
• TalkOrigins Archive: Post of the Month: March 2006 , The History of Creationism by
Lenny Flank.
30.^ Timeline and Myths of Scopes Trial
 31.^ a b Nelkin, Dorothy (2000). The Creation Controversy: Science or Scripture in Schools. New
York: iUniverse. pp. 242. ISBN 0-595-00194-7.
32.^ Epperson et al. v. Arkansas, 393 U.S. 97 (U.S. Supreme Court 1968-11-12).
33.^ Larson, Edward J. (2003). Trial and Error: The American Controversy Over Creation and
Evolution. Oxford University Press. pp. 276. ISBN 0195154703.
34.^ Larson 2004, p. 248,250, see also Dobzhansky 1973
35.^ Larson 2004, p. 251
36.^ Larson 2004, p. 252
37.^ Larson 2004, p. 255,Numbers 1992, p. xi,200-208
38.^ Larson 2004, p. 255
39.^ Numbers 1992, p. 284-285
40.^ Numbers 1992, p. 284-6
41.^ Quoting Larson 2004, p. 255-256: "Fundamentalists no longer merely denounced
Darwinism as false; they offered a scientific-sounding alternative of their own, which they called
either 'scientific creationism (as distinct from religious creationism) or 'creation science' (as opposed
to evolution science."
42.^ Larson 2004, p. 254-255, Numbers 1998, p. 5-6
43.^ Ruling, Kitzmiller v. Dover Area School District pp 7-9.
44.^ "NCSE Resource". Evolving Banners at the Discovery Institute. 2002-08-29.
http://ncse.com/creationism/general/evolving-banners-at-discovery-institute. Retrieved 2009-04-07.
45.^ "Top Questions-1.What is the theory of intelligent design?". Discovery Institute.
http://www.discovery.org/csc/topQuestions.php#questionsAboutIntelligentDesign. Retrieved 2007-
05-13. .
46.^ Verderame 2007,Simon 2006
 47.^ Dewey 1994, p. 31, and Wiker 2003, summarizing Gould.
48.^ Larson 2004, p. 258 "Virtually no secular scientists accepted the doctrines of creation
science; but that did not deter creation scientists from advancing scientific arguments for their
position." See also Martz & McDaniel 1987, p. 23, a Newsweek article which states "By one count
there are some 700 scientists (out of a total of 480,000 U.S. earth and life scientists) who give
credence to creation-science, the general theory that complex life forms did not evolve but appeared
'abruptly'."
49.^ Statement from the Council of the Biological Society of Washington at the Wayback
Machine (archived September 26, 2007).
50.^ Bumiller 2005, Peters & Hewlett 2005, p. 3
51.^ Some question group's move with elections nearing 6News Lawrence, Lawrence Journal-
World. 7 July 2006.
52.^ Evolution’s foes lose ground in Kansas MSNBC, 2 August 2006.
53.^ Evolution of Kansas science standards continues as Darwin's theories regain prominence
The Associated Press, via the International Herald Tribune, 13 February 2007.
54.^ The "Evolution" of Creationism Timeline: how creationism has "evolved". People for the
American Way.
55.^ Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., Conclusion of
Ruling.
56.^ Johnson 1998, Hodge 1874, p. 177, Wiker 2003, Peters & Hewlett 2005, p. 5--Peters and
Hewlett argue that the atheism of many evolutionary supporters must be removed from the debate
57.^ Lenski 2000, p. Conclusions
58.^ Johnson 1998
59.^ Einstein 1930, p. 1-4
 60.^ Dawkins 1997
61.^ a b Free Executive Summary, Science and Creationism: A View from the National Academy
of Sciences, Second Edition, Steering Committee on Science and Creationism, National Academy of
Sciences, 1999, ISBN 978-0-309-06406-4.
62.^ Johnson 1993, p. 63, Tolson 2005, Moran 1993 ; Selman v. Cobb County School District.
US District Court for the Northern District of Georgia (2005); Talk. Origins; Bill Moyers et al., 2004.
"Now with Bill Moyers." PBS. Accessed 2006-01-29. Interview with Richard Dawkins
63.^ Merriam-Webster online dictionary. www.m-w.com
64.^ Gould 1981
65.^ See:
• Number 1992, p. 247;
• Wilkins, John S, Evolution and Philosophy: Is Evolution Science, and What Does
'Science' Mean?, TalkOrigins Archive
66.^ Popper 1976, p. 168 and 172 quoted in Kofahl 1981
67.^ Unknown sociologist quoted in Numbers 1992, p. 247
68.^ Kofahl 1989 as quoted by Numbers 1992, p. 247
69.^ Lewin 1982
70.^ Numbers(2006) p274
71.^ Kofahl 1981, p. 873
72.^ Talkorigins summary of Karl Popper attitudes towards evolution
73.^ Karl Popper (1978). "Natural selection and the emergence of mind". Dialectica 32(3/4): 339
—355.
http://www.informationphilosopher.com/solutions/philosophers/popper/natural_selection_and_the_e
mergence_of_mind.html. , Massimo Pigliucci (Sept-October 2004). "Did Popper refute evolution?".
Skeptical Inquirer.
http://www.lehman.edu/deanhum/philosophy/platofootnote/PlatoFootnote.org/Outreach_files/2004-
Did%20Popper%20refute%20evolution%3F-Skeptical%20Inquirer.pdf.
74.^ Ruse 1999, p. 13-37, which discusses conflicting ideas about science among Karl Popper,
Thomas Samuel Kuhn, and their disciples.
75.^ As quoted by Wallis 2005, p. 32. Also see Dawkins 1986 and Dawkins 1995
76.^ Wallis 2005, p. 6 Dawkins quoting Haldane
77.^ Dorman 1996
78.^ "One thing we have come to realise in Creation Science is that the Lord has not just called
us to knock down evolution, but to help in restoring the foundation of the gospel in our society. We
believe that if the churches took up the tool of Creation Evangelism in society, not only would we see
a stemming of the tide of humanistic philosophy, but we would also see the seeds of revival sown in
a culture which is becoming increasingly more pagan each day.
...
It is also worth noting the comment in the book, ‘By Their Blood-Christian Martyrs of the 20th
Century’ (Most Media) by James and Marti Helfi, on page 49 and 50: ‘New philosophies and
theologies from the West also helped to erode Chinese confidence in Christianity. A new wave of so-
called missionaries from mainline Protestant denominations came teaching evolution and a non-
supernatural view of the Bible. Methodist, Presbyterian, Congregationalist, and Northern Baptist
schools were especially hard hit. Bertrand Russell came from England preaching atheism and
socialism. Destructive books brought by such teachers further undermined orthodox Christianity. The
Chinese Intelligentsia who had been schooled by Orthodox Evangelical Missionaries were thus
softened for the advent of Marxism.’ Evolution is destroying the Church and society, and Christians
need to be awakened to that fact!"Ham, Ken. Creation Evangelism (Part II of Relevance of Creation).
Creation Magazine '6'(2):17, November 1983.
79.^ "...I want to list seventeen summary statements which, if true, provide abundant reason why
the reader should reject evolution and accept special creation as his basic world-view. ...
13. Belief in special creation has a salutary influence on mankind, since it encourages responsible
obedience to the Creator and considerate recognition of those who were created by Him. ...
16. Belief in evolution and animal kinship leads normally to selfishness, aggressiveness, and fighting
between groups, as well as animalistic attitudes and behaviour by individuals." Henry M. Morris, The
Remarkable Birth of Planet Earth (Creation-Life Publishers, 1972), pp. vi-viii. Cited in Appeal to
Consequences, Fallacy Files
80.^ Johnson 1993, p. 69 where Johnson cites three pages spent in Issac Asimov's New Guide
to Science that take creationists to task, while only spending one half page on evidence of evolution.
81.^ Evolution: what's the real controversy?, John Timmer, Nobel Intent, 7 May 2008
82.^ The Discontinuity of Life, Kurt Wise, Answers in Genesis
83.^ A Primer on the Tree of Life, Casey Luskin, Center for Science and Culture, Discovery
Institute
84.^ Evolution versus the People, Henry M. Morris, Institute for Creation Research
85.^ Stringer, Chris; Andrews, Peter (2005). The complete world of human evolution. London:
Thames & Hudson. pp. 240. ISBN 0-500-05132-1.
86.^ Relethford, John (2003). Reflections of our past: how human history is revealed in our
genes. Boulder, Colo: Westview Press. pp. 257. ISBN 0-8133-3958-8.
 87.^ "Toumaï the Human Ancestor: Skull of Oldest Known Hominid Unearthed in Chad". NPR:
All Things Considered. http://www.npr.org/programs/atc/features/2002/july/toumai/index.html.
Retrieved 2009-02-21.
88.^ Chimps are human, gene study implies New Scientist, website, 19 May 2003
89.^ Was Java Man a gibbon?, Jim Foley, TalkOrigins website, 30 April 2003.
90.^ See disputes over the classification of Neanderthals in The Counter-Creationism
Handbook, Mark Isaak, University of California Press (2007), ISBN 978-0520249264
91.^ Comparison of all skulls, Jim Foley, TalkOrigins website, 8 August 2005.
92.^ Scientific Creationism, Henry M. Morris, 1985, pp. 78-90
93.^ Life—How Did It Get Here?, Watchtower Bible and Tract Society, 1985, pp. 57-59
94.^ Haywood, Alan (1985) Creation and Evolution.Triangle Books, London. Quoted in Hooking
Leviathan by Its Past, Stephen Jay Gould
95.^ Exploring the Way Life Works, Mahlon B. Hoagland, Bert Dodson, Judith Hauck, p298
96.^ Biblical chronogenealogies
97.^ The Meaning of the Chronogenealogies of Genesis 5 and 11
98.^ IAP Statement on the Teaching of Evolution, Interacademy Panel on Global Issues, 21
June 2006.
99.^ Del Peloso, E.F.; Da Silva, L.; De Mello, G.F.P.; Arany-prado, L.I. (2005). "The age of the
Galactic thin disk from Th/Eu nucleocosmochronology". A&A 434: 301–308. doi:10.1051/0004-
6361:20047060.
100.^ Nuclear Decay: Evidence For A Young World, D. Russell Humphreys, Impact, Number
352, October 2002.
 101.^ Young-Earth Creationist Helium Diffusion "Dates" Fallacies Based on Bad Assumptions
and Questionable Data, Kevin R. Henke, TalkOrigins website, Original version: 17 March 2005,
Revision: 24 November 2005.
102.^ R.A.T.E: More Faulty Creation Science from The Institute for Creation Research , J. G.
Meert, Gondwana Research, The Official Journal of the International Association for Gondwana, 13
November 2000 (updated 6 February 2003).
103.^ Dating methods discussed were potassium-argon dating, argon-argon dating, rubidium-
strontium dating, samarium-neodymium dating, lutetium-hafnium, rhenium-osmium dating, and
uranium-lead dating.
104.^ Radiometric Dating, A Christian Perspective, Roger C. Wiens, American Scientific
Affiliation, p20-21
105.^ Dobzhansky 1973
106.^ a b Pieret 2006
107.^
• Isaak, Mark (2004). "Index to Creationist Claims: Claim CA113". Talk.origins.
http://www.talkorigins.org/indexcc/CA/CA113.html. Retrieved 2007-12-27.
• Dunford, Mike (2007-07-02). "A new (mis)take on an old paper (and other posts)".
[[The Panda's Thumb (blog)|]]. http://www.pandasthumb.org/archives/quote_mines/.
Retrieved 2007-12-26.
• Myers, PZ (2004-09-11). "I'm shocked, shocked to find that quote mining is going on
in there!". Pharyngula.
http://pharyngula.org/index/weblog/comments/im_shocked_shocked_to_find_that_quote_min
ing_is_going_on_in_there. Retrieved 2007-12-27.
 108.^ a b c "Some bills seek to discredit evolution by emphasizing so-called "flaws" in the theory
of evolution or "disagreements" within the scientific community. Others insist that teachers have
absolute freedom within their classrooms and cannot be disciplined for teaching non-scientific
"alternatives" to evolution. A number of bills require that students be taught to "critically analyze"
evolution or to understand "the controversy." But there is no significant controversy within the
scientific community about the validity of the theory of evolution. The current controversy surrounding
the teaching of evolution is not a scientific one." AAAS Statement on the Teaching of Evolution
American Association for the Advancement of Science. 16 February 2006 (PDF file)
109.^ Ruling, Kitzmiller v. Dover Area School District, page 89
110.^ a b "That this controversy is one largely manufactured by the proponents of creationism
and intelligent design may not matter, and as long as the controversy is taught in classes on current
affairs, politics, or religion, and not in science classes, neither scientists nor citizens should be
concerned." Intelligent Judging — Evolution in the Classroom and the Courtroom George J. Annas,
New England Journal of Medicine, Volume 354:2277-25 May 2281 2006
111.^ See: 1) List of scientific societies rejecting intelligent design 2) Kitzmiller v. Dover page 83.
The Discovery Institute's Dissent From Darwin Petition has been signed by about 500 scientists. The
AAAS, the largest association of scientists in the U.S., has 120,000 members, and firmly rejects
intelligent design and denies that there is a legitimate scientific controversy. More than 70,000
Australian scientists and educators condemn teaching of intelligent design in school science classes.
Statements from Scientific Organizations on the status intelligent design and other forms of
creationism.
112.^ "In summary, the disclaimer singles out the theory of evolution for special treatment,
misrepresents its status in the scientific community, causes students to doubt its validity without
scientific justification, presents students with a religious alternative masquerading as a scientific
theory, directs them to consult a creationist text as though it were a science resource, and instructs
students to forgo scientific inquiry in the public school classroom and instead to seek out religious
instruction elsewhere." Ruling - disclaimer, pg. 49 Kitzmiller v. Dover Area School District.
113.^ "ID's home base is the Center for Science and Culture at Seattle's conservative Discovery
Institute. Meyer directs the center; former Reagan adviser Bruce Chapman heads the larger institute,
with input from the Christian supply-sider and former American Spectator owner George Gilder (also
a Discovery senior fellow). From this perch, the ID crowd has pushed a "teach the controversy"
approach to evolution that closely influenced the Ohio State Board of Education's recently proposed
science standards, which would require students to learn how scientists "continue to investigate and
critically analyze" aspects of Darwin's theory." Chris Mooney. The American Prospect. 2 December
2002 Survival of the Slickest: How anti-evolutionists are mutating their message
114.^ Teaching Intelligent Design: What Happened When? by William A. Dembski"The clarion
call of the intelligent design movement is to "teach the controversy." There is a very real controversy
centering on how properly to account for biological complexity (cf. the ongoing events in Kansas),
and it is a scientific controversy."
115.^ Nick Matzke's analysis shows how teaching the controversy using the Critical Analysis of
Evolution model lesson plan is a means of teaching all the intelligent design arguments without using
the intelligent design label.No one here but us Critical Analysis-ists... Nick Matzke. The Panda's
Thumb, 11 July 2006
116.^ "has the effect of implicitly bolstering alternative religious theories of origin by suggesting
that evolution is a problematic theory even in the field of science." ... The effect of Defendants’
actions in adopting the curriculum change was to impose a religious view of biological origins into the
biology course, in violation of the Establishment Clause. Kitzmiller v. Dover Area School District,
Conclusion, page 134
 117.^ a b "ID's backers have sought to avoid the scientific scrutiny which we have now
determined that it cannot withstand by advocating that the controversy, but not ID itself, should be
taught in science class. This tactic is at best disingenuous, and at worst a canard. The goal of the
IDM is not to encourage critical thought, but to foment a revolution which would supplant evolutionary
theory with ID."Kitzmiller v. Dover Area School District, whether ID is science, page 89
118.^ AAAS Dialogue on Science, Ethics, and Religion, 20 April 2006, Emmett Holman,
Associate Professor of Philosophy from George Mason University, retrieved 2007-04-29
119.^ Science, 3 APRIL 2009
120.^ Woods 2005, p. 67-114, Chapter Five: The Church and Science
121.^ Morris 1982
122.^ Index to Creationist Claims - Claim CA114 edited by Mark Isaak. 2005
123.^ Index to Creationist Claims - Claim CA114.22 edited by Mark Isaak. 2005
124.^ [1] Science and religion: Conflicts & occasional agreements
125.^ Why I Won't Debate Creationists, Richard Dawkins, Reason: In the News,
richarddawkins.net, the official Richard Dawkins website, Monday, 15 May 2006.
126.^ a b c Shermer, Michael (2004-05-10). "Then a Miracle Occurs: An Obstreperous Evening
with the Insouciant Kent Hovind, Young Earth Creationist and Defender of the Faith". eSkeptic
Online. http://www.skeptic.com/eskeptic/04-05-10.html#miracle. Retrieved 2007-02-11.
127.^ Massimo Pigliucci. Denying Evolution: Creationism, Scientism, and the Nature of Science.
(Sinauer, 2002): ISBN 0878936599 page 102.
128.^ Shermer, Michael. 'Why People Believe Weird Things', Owl Books, 2002. Paperback ed,
p. 153.
129.^ Statements from Scientific Organizations, NCSE
130.^ Unintelligent designs on Darwin, Edward Humes, Pittsburgh Tribune-Review
 131.^ Michael Le Page (19 April 2008). "Evolution myths: It doesn't matter if people don't grasp
evolution". New Scientist 198 (2652): 31. http://www.newscientist.com/article/mg19826523.000-
evolution-myths-it-doesnt-matter-if-people-dont-grasp-evolution.html.
132.^ Jeff Hecht (19 August 2006). "Why doesn't America believe in evolution?". New Scientist
191 (2565): 11. http://www.newscientist.com/article/dn9786-why-doesnt-america-believe-in-
evolution.html.
133.^ a b Pitock, Todd (2007). "Science and Islam". Discover: 36–45.
134.^ a b c Gregory Katz (2008-02-16). "Clash Over Creationism Is Evolving In Europe's
Schools". Associated Press. http://www2.tbo.com/content/2008/feb/16/na-clash-over-creationism-is-
evolving-in-europes-s. Retrieved 2008-02-17. /
135.^ a b c Taner Edis. "Cloning Creationism in Turkey". RNCSE 19 (6): 30-35. National Center
for Science Education. http://ncse.com/rncse/19/6/cloning-creationism-turkey. Retrieved 2008-02-
17.
136.^ "Serbia reverses Darwin suspension". BBC. 2004-09-09.
http://news.bbc.co.uk/1/hi/world/europe/3642460.stm. Retrieved 2008-02-17.
137.^ Roger Highfield (2007-02-10). "Creationists rewrite natural history". London: The
Telegraph. http://www.telegraph.co.uk/earth/main.jhtml?
view=DETAILS&grid=&xml=/earth/2007/10/02/scihist102.xml. Retrieved 2008-02-17.
138.^ New Scientist 10 November 2007, p. 72
139.^ Christianity – Pentecostalism Australian Broadcasting Corporation
140.^ Carly Hennessy (May 30, 2010). "Creationism to be taught in Queensland classrooms".
Herald Sun. http://www.heraldsun.com.au/news/national/creationism-to-be-taught-in-queensland-
classrooms/story-e6frf7l6-1225873019548. Retrieved July 22, 2010.
141.^ Plimer, Ian "Telling lies for God- Reason versus Creationism", (Random House)
 142.^ "Telling Lies for God"? - One Man's Crusade, accessed 2008-02-05, Quantum. See
transcript link for detail.
143.^ "Evolution and religion: In the beginning". The Economist. 2007-04-19.
http://www.economist.com/world/displaystory.cfm?story_id=9036706. Retrieved 2007-04-25. This
article gives a worldwide overview of recent developments on the subject of the controversy.

[edit] References
• AAAS, American Association for the Advancement of Science (2006-02-16) (pdf), Statement
on the Teaching of Evolution, aaas.org,
http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf, retrieved 2007-01-14
• Bumiller, Elisabeth (2005-08-03), "Bush Remarks Roil Debate on Teaching of Evolution",
The New York Times (2005-08-03), http://www.nytimes.com/2005/08/03/politics/03bush.html?
ex=1280721600&en=8bbf73d2f5204260&ei=5088&partner=r, retrieved 2007-02-03
• Barbour, Ian G. (1997), Religion and Science: Historical and Contemporary Issues (first
revised ed.), HarperSanFrancisco, pp. 58,65, ISBN 0060609389
• Burns, Edward M.; Ralph, Philip Lee; Lerner, Robert E.; Standish, Meacham (1982), World
Civilizations Their History and Their Culture (Sixth ed.), W.W. Norton & Company, ISBN 0-393-
95077-8
• Dawkins, Richard (1986), The Blind Watchmaker, W. W. Norton & Company, Inc., ISBN 0-
393-31570-3
• Dawkins, Richard (1995), River Out of Eden, Basic Books, ISBN 0-465-06990-8
• Dawkins, Richard (January 1997), "Is Science a Religion?", Humanist,
http://www.thehumanist.org/humanist/articles/dawkins.html, retrieved 2007-01-30
 • Desmond, Adrian; Moore, James (1991), Darwin, London: Michael Joseph, Penguin Group,
ISBN 0-7181-3430-3
• Dewey, John (1994), "The Influence of Darwinism on Philosophy", in Martin Gardner, Great
Essays in Science, Prometheus Books, ISBN 0-87975-853-8
• Dobzhansky, Theodosius (1973), "Nothing in Biology Makes Sense Except in the Light of
Evolution", American Biology Teacher (National Association of Biology Teachers) 35: 125–129,
http://www.2think.org/dobzhansky.shtml, retrieved 2007-01-14
• Transcription of McLean v. Arkansas Board of Education Decision by U.S. District Court
Judge William R. Overton, TalkOrigins Archive Foundation, 1996-01-30,
http://www.talkorigins.org/faqs/mclean-v-arkansas.html, retrieved 2007-01-31
• Einstein, Albert (1930-11-09), "Religion and Science", New York Times Magazine: 1–4,
http://www.sacred-texts.com/aor/einstein/einsci.htm, retrieved 2007-01-30
• Gould, Stephen Jay (1981), Evolution as Fact and Theory, The Unofficial Stephen Jay Gould
Archive, http://www.stephenjaygould.org/library/gould_fact-and-theory.html, retrieved 2007-01-17
• Hodge, Charles (1874), What is Darwinism?, Scribner, Armstrong, and Company, ASIN
B0006AEEMO, http://www.gutenberg.org/files/19192/19192-8.txt, retrieved 2007-01-14
• Hovind, Kent (2006), Dr. Hovind's $250,000 Offer, Creation Science Evangelism,
http://www.drdino.com/articles.php?spec=67, retrieved 2007-01-17
• Huxley, Thomas H. (1902), "An Episcopal Trilogy 1887", Collected Essays Science and
Christian Tradition (Kessinger Publishing) V: 126–159, ISBN 978-1417973729,
http://www.gutenberg.org/files/15905/15905-8.txt, retrieved 2007-01-14
 • IAP, Interacademy Panel (2006-06-21), IAP Statement on the Teaching of Evolution,
interacademies.net, http://www.interacademies.net/Object.File/Master/6/150/Evolution
%20statement.pdf, retrieved 2007-01-14
• Johnson, Phillip E. (1998), Reason in the Balance: The Case Against Naturalism in Science,
Law & Education, IntervaVsity Press, ISBN 0-8308-1929-0
• Johnson, Phillip E. (1993), Darwin on Trial (2nd ed.), InterVarsity Press, ISBN 0-8308-1324-
1
• Kofahl, Robert E. (22 May 1981), "Popper on Darwinism", Science (New Series) 212 (4497)
• Kofahl, Robert E. (June, 1989), "The Hierarchy of Conceptual Levels For Scientific Thought
And Research", CRS Quarterly (The Creation Research Society) 26 (1),
http://creationresearch.org/crsq/abstracts/sum26_1.html, retrieved 2007-01-29
• Larson, Edward J. (2004). Evolution. Modern Library. ISBN 0-679-64288-9.
• Lenski, Richard E. (2000), Evolution: Fact and Theory, AcitionBioscience.org,
http://www.actionbioscience.org/evolution/lenski.html, retrieved 2007-01-23
• Martz, Larry; McDaniel, Ann (1987-06-29), "Keeping God out of the Classroom (Washington
and bureau reports)", Newsweek (Newsweek Inc.) CIX (26): 23–24, ISBN 0028-9604
• Moore, James (2006), "Evolution and Wonder — Understanding Charles Darwin", Speaking
of Faith (Radio Program), American Public Media,
http://speakingoffaith.publicradio.org/programs/darwin/transcript.shtml, retrieved 2007-01-24
• Moran, Laurence (1993), Evolution is a Fact and a Theory, TalkOrigins Archive Foundation,
http://www.talkorigins.org/faqs/cre-error.html, retrieved 2007-01-23
 • Morris, Henry M. (1982-01-01), "Bible-Believing Scientists of the Past", Impact (Institute for
Creation Research) 103, http://www.icr.org/index.php?module=articles&action=view&ID=185,
retrieved 2007-01-20
• Myers, P.Z. (2006-06-18), "Ann Coulter: No Evidence for Evolution?", Pharyngula
(ScienceBlogs), http://scienceblogs.com/pharyngula/2006/06/ann_coulter_no_evidence_for_ev.php,
retrieved 2007-09-12
• NSTA, National Science Teachers Association (2007). "An NSTA Evolution Q&A".
http://www.nsta.org/publications/evolution.aspx. Retrieved 2008-02-01.
• Numbers, Ronald L. (1992). Creationists: The Evolution of Scientific Creationism. Alfred A.
Knopf, Inc.. pp. 224. ISBN 0-679-40104-0.
• Numbers, Ronald (2006-11-30). The Creationists: From Scientific Creationism to Intelligent
Design, Expanded Edition. Harvard University Press. p. 624 pages. ISBN 0674023390.
• Numbers, Ronald L. (1998-11-15), Darwinism Comes to America, Harvard University Press,
pp. 224, ISBN 0674193121
• Peters, Ted; Hewlett, Martinez (2005-12-22) (pdf), The Evolution Controversy: Who's
Fighting with Whom about What?, Pacific Lutheran Theological Seminary,
http://www.plts.edu/docs/EvolutionBrief2.pdf, retrieved 2007-01-28
• Pieret, John (2006), The Quote Mine Project Or, Lies, Damned Lies and Quote Mines ,
TalkOrigins Archive Foundation, http://www.talkorigins.org/faqs/quotes/mine/project.html, retrieved
2007-01-23
• Pinholster, Ginger (2006-02-19), AAAS Denounces Anti-Evolution Laws as Hundreds of K-
12 Teachers Convene for 'Front Line' Event, aaas.org,
http://www.aaas.org/news/releases/2006/0219boardstatement.shtml, retrieved 2007-01-14
 • Popper, Karl (1976), Unended Quest: An Intellectual Autobiography, Open Court Publishing
Co., ISBN 0-87548-366-6
• Popper, Karl (1980), "Letter to the Editor", New Scientist 87
• Ruse, Michael (1999-04-30), Mystery of Mysteries: Is Evolution a Social Construction ,
Harvard University Press, pp. 320, ISBN 0-674-46706-X
• Simon, Stephanie (2006), "Their Own Version of a Big Bang: Those who believe in
creationism – children and adults – are being taught to challenge evolution's tenets in an in-your-face
way.", Los Angeles Times (2006-02-11), http://philosophy.tamucc.edu/article.pl?
sid=06/02/12/1727208&mode=thread, retrieved 2007-02-03
• Tolson, Jay (2005-09-05), Religion in America, "Religion in America: Intelligent Design on
Trial", U.S. News & World Report,
http://www.usnews.com/usnews/culture/articles/050928/28religion.htm, retrieved 2007-01-06
• van Wyhe, John (2006), Charles Darwin: gentleman naturalist: A biographical sketch ,
http://darwin-online.org.uk/darwin.html, retrieved 2007-01-24
• Verderame, John (2007), Creation evangelism: cutting through the excess,
answersingenesis.org, http://www.answersingenesis.org/docs2001/0510news.asp, retrieved 2007-
02-07
• Wallis, Claudia (2005-08-07), "The Evolution Wars", Time Magazine,
http://www.time.com/time/magazine/article/0,9171,1090909-1,00.html Retrieved on 2007-01-31
• Wiker, Benjamin D. (July/August 2003), "Does Science Point to God? Part II: The Christian
Critics", Crisis Politics, Culture, and the Church (CRISIS Magazine),
http://www.crisismagazine.com/julaug2003/feature1.htm, retrieved 2007-01-21
 • Witham, Larry A. (2002). "From Broadway to Biophilia". Where Darwin Meets the Bible:
Creationists and Evolutionists in America (kindle ed.). Oxford University Press. Chapter 2, sect.
"From Broadway to Biophilia", para. 9. ISBN 978-0195150452.
• Woods, Thomas E., Jr. (2005), How the Catholic Church Built Western Civilization , Regnery
Publishing, Inc., ISBN 0-89526-038-7

[edit] Published books and other resources

• Burian, RM: 1994. Dobzhansky on Evolutionary Dynamics: Some Questions about His
Russian Background; in Adams, Mark A. (1994). The Evolution of Theodosius Dobzhansky: essays
on his life and thought in Russia and America. Princeton, N.J.: Princeton University Press. ISBN 0-
691-03479-6.
• Butler, Samuel (2007). Evolution, Old And New: Or The Theories Of Buffon, Dr. Erasmus
Darwin And Lamarck, As Compared With That Of Charles Darwin (1911) . Kessinger Publishing, LLC.
ISBN 0548641323.
• Beer, Gillian; Darwin, Charles (1996). The origin of species. Oxford [Oxfordshire]: Oxford
University Press. ISBN 019283438X.
• Gould, Stephen Jay; Dobzhansky, Theodosius Grigorievich (1982). Genetics and the origin
of species. New York: Columbia University Press. ISBN 0231054750.
• Haught, John F. (2010). Making Sense of Evolution: Darwin, God, and the Drama of Life .
Louisville: Westminster John Knox Press. pp. 144. ISBN 9780664232856.
• Henig, Robin Marantz (2000). The monk in the garden: the lost and found genius of Gregor
Mendel, the father of genetics. Boston: Houghton Mifflin. ISBN 0395977657.
 • Kutschera U, Niklas KJ (2004). "The modern theory of biological evolution: an expanded
synthesis". Naturwissenschaften 91 (6): 255–76. doi:10.1007/s00114-004-0515-y. PMID 15241603.
• Mayr, Ernst (1985). The Growth of Biological Thought: Diversity, Evolution, and Inheritance .
Cambridge, Mass: Belknap Press. ISBN 0674364465.
• Miller, James H. (2001). An evolving dialogue: theological and scientific perspectives on
evolution. Valley Forge, Pa: Trinity Press International. ISBN 1-56338-349-7.
• Morris, HR (1963). The Twilight of Evolution. Grand Rapids, MI: Baker Pub Group.
ISBN 0801058627.
• Numbers, Ronald L. (1993). The creationists. Berkeley: University of California Press.
ISBN 0520083938.
• Pennock RT (2003). "Creationism and intelligent design". Annu Rev Genomics Hum Genet 4:
143–63. doi:10.1146/annurev.genom.4.070802.110400. PMID 14527300.
• Sagan, Carl (1997). The demon-haunted world: science as a candle in the dark . New York:
Ballantine Books. ISBN 0345409469.
• Scott, E.C. (1997). "Antievolution And Creationism In The United States". Annual Review of
Anthropology 26 (1): 263–289. doi:10.1146/annurev.anthro.26.1.263.
• Maynard Smith, "The status of neo-Darwinism," in "Towards a Theoretical Biology" (C.H.
Waddington, ed., University Press, Edinburgh, 1969.
• Hull, D.L. (1999). "The Use and Abuse of Sir Karl Popper". Biology and Philosophy 14 (4):
481–504. doi:10.1023/A:1006554919188. http://www.springerlink.com/index/T4220X33T0284444.pdf.
Retrieved 2008-04-11.
• Strobel, Lee (2004). The case for a Creator: a journalist investigates scientific evidence that
points toward God. Grand Rapids, Mich: Zondervan/Willow Creek Resources. ISBN 0310240506.
[edit] External links
Wikibooks has a book on the topic of
Introduction to Paleoanthropology
• Gallup public opinion poll in regards to the concepts of Evolution, Creationism, and Intelligent
Design as of May 2007
• Data by country regarding the percentage of the population that believes in evolution
• Opinion of Tennessee Supreme Court in Scopes v. State
Creationism as social policy
• Isaac Asimov, The "Threat" of Creationism.
Creationist beliefs
• Answers in Genesis, Young Earth Creationism
• Reasons to Believe, Old Earth Creationism
Scientific rebuttals
• Talk.origins Index to Creationist Claims
• Creation/Evolution journal online 1980-1996 (full run)
Evolution versus creationism debates
Location and Year Pro-evolution Pro-creationism
 link

Stanford
1994 Will Provine Phillip E. Johnson
University

Nova on-line 1996 Kenneth R. Miller Phillip E. Johnson

Kenneth R. Miller, Phillip E. Johnson, Michael

Firing Line 1997 Michael Ruse, Eugenie Scott & Behe, David Berlinski & William F.
Barry Lynn Buckley

University of
2007 Lewis Wolpert Steve Fuller
London (transcript)
Retrieved from "http://en.wikipedia.org/wiki/Creation%E2%80%93evolution_controversy"

Categories: Creationism | Evolution and religion | Intelligent design controversies

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 Intelligent design

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 movement

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 Intelligent design is the proposition that "certain features of the universe and of living things are best
explained by an intelligent cause, not an undirected process such as natural selection."[1][2] It is a modern
form of the traditional teleological argument for the existence of God, but one which purposefully avoids
specifying the nature or identity of the designer.[3]The idea was developed by a group of American
creationists who reformulated their argument in the creation–evolution controversy to circumvent court rulings
that prohibit the teaching of creationism as science.[4][5][6] Intelligent design's leading proponents – all of
whom are associated with the Discovery Institute, a politically conservative think tank[7][8] – believe the
designer to be the God of Christianity.[9][10]
Advocates of intelligent design seek to fundamentally redefine science to accept supernatural
explanations,[11] arguing that intelligent design is a scientific theory under this new definition of science.[1]
The unequivocal consensus in the scientific community is that intelligent design is not science.[12][13][14]
[15] The U.S. National Academy of Sciences has stated that "creationism, intelligent design, and other claims
of supernatural intervention in the origin of life or of species are not science because they are not testable by
the methods of science."[16] The U.S. National Science Teachers Association and the American Association
for the Advancement of Science have termed it pseudoscience.[17][18][19] Others in the scientific community
have concurred,[20] and some have called it junk science.[21][22]
Intelligent design originated in response to the 1987 United States Supreme Court Edwards v.
Aguillard ruling involving separation of church and state.[4] Its first significant published use was in Of
Pandas and People, a 1989 textbook intended for high-school biology classes.[23] Several additional books
on the subject were published in the 1990s. By the mid-1990s, intelligent design proponents had begun
clustering around the Discovery Institute and more publicly advocating the inclusion of intelligent design in
public school curricula.[24] With the Discovery Institute and its Center for Science and Culture serving a
central role in planning and funding, the "intelligent design movement" grew increasingly visible in the late
1990s and early 2000s, culminating in the 2005 Dover trial which challenged the intended use of intelligent
design in public school science classes.[7]
In Kitzmiller v. Dover Area School District, a group of parents of high-school students challenged a
public school district requirement for teachers to present intelligent design in biology classes as an
alternative "explanation of the origin of life". U.S. District Judge John E. Jones III ruled that intelligent design
is not science, that it "cannot uncouple itself from its creationist, and thus religious, antecedents", and that
the school district's promotion of it therefore violated the Establishment Clause of the First Amendment to the
U.S. Constitution.[25]
Contents
[hide]
• 1 History
• 1.1 Origin of the concept
• 1.2 Development of its modern form
• 1.3 Origin of the term
• 2 Integral concepts
• 2.1 Irreducible complexity
• 2.2 Specified complexity
• 2.3 Fine-tuned Universe
• 2.4 Intelligent designer
• 3 Movement
• 3.1 Religion and leading proponents
• 3.2 Reaction from other creationist groups
• 3.3 Polls
• 3.4 Film
• 4 Creating and teaching the controversy
• 4.1 Defining science
• 4.2 Peer review
• 4.3 Intelligence as an observable quality
• 4.4 Arguments from ignorance
[edit] History
[edit] Origin of the concept
Further information: Argument from poor design, Teleological argument, and Watchmaker analogy

A first-century bust of Socrates. The idea of intelligent design, or the argument from design, is an
ancient one, held in some form by Plato, and Aristotle.
Whether the complexity of nature indicates purposeful design has been the subject of debate since
the Greeks. In the 4th century BCE, Plato posited a good and wise "demiurge" as the creator and first cause
of the cosmos in his Timaeus.[26][27] In his Metaphysics, Aristotle developed the idea of an "Unmoved
Mover".[28] In De Natura Deorum (On the Nature of the Gods, 45 BCE) Cicero wrote that "the divine power is
to be found in a principle of reason which pervades the whole of nature."[29] This line of reasoning has come
to be known as the teleological argument for the existence of God. Some well-known forms of it were
expressed in the 13th century by Thomas Aquinas and in the 19th century by William Paley. Aquinas, in his
Summa Theologiae, used the concept of design in his "fifth proof" for God's existence.[30]
In the early 19th century, Paley's argument from design in Natural Theology (1802), used the
watchmaker analogy,[31] and such arguments led to the development of what was called natural theology,
the study of nature as way of understanding "the mind of God". This movement fueled the passion for
collecting fossils and other biological specimens, which ultimately led to Charles Darwin's On the Origin of
Species (1859). Similar reasoning postulating a divine designer is embraced today by many believers in
theistic evolution, who consider modern science and the theory of evolution to be compatible with the
concept of a supernatural designer. In correspondence about the question with Asa Gray, Darwin wrote that
"I cannot honestly go as far as you do about Design. I am conscious that I am in an utterly hopeless muddle. I
cannot think that the world, as we see it, is the result of chance; & yet I cannot look at each separate thing as
the result of Design."[32] Though he had studied Paley's work while at university, by the end of his life he
came to regard it as useless for scientific development.[33]

[edit] Development of its modern form

Intelligent design in the late 20th and early 21st century is a development of natural theology that
seeks to change the basis of science and undermine evolutionary theory.[34] As evolutionary theory
expanded to explain more phenomena, the examples held up as evidence of design changed, though the
essential argument remains the same: complex systems imply a designer. Past examples have included the
eye and the feathered wing; current examples are typically biochemical: protein functions, blood clotting, and
bacterial flagella; see irreducible complexity.
Philosopher Barbara Forrest writes that the intelligent design movement began in 1984 with the
publication by Jon A. Buell's the Foundation for Thought and Ethics of The Mystery of Life's Origin by Charles
B. Thaxton, a chemist and creationist. Thaxton held a conference in 1988, "Sources of Information Content in
DNA," which attracted creationists such as Stephen C. Meyer. Forrest writes that, in December 1988,
Thaxton decided to use the term "intelligent design," instead of creationism, for the movement.[35]
In March 1986 a review by Meyer used information theory to suggest that messages transmitted by
DNA in the cell show "specified complexity" specified by intelligence, and must have originated with an
intelligent agent.[36] In November of that year Thaxton described his reasoning as a more sophisticated form
of Paley's argument from design.[37] At the Sources of Information Content in DNA conference in 1988 he
said that his intelligent cause view was compatible with both metaphysical naturalism and supernaturalism,
[38]
Intelligent design avoids identifying or naming the agent of creation—it merely states that one (or
more) must exist—but leaders of the movement have said the designer is the Christian God.[9][10][39][40][41]
Whether this lack of specificity about the designer's identity in public discussions is a genuine feature of the
concept, or just a posture taken to avoid alienating those who would separate religion from the teaching of
science, has been a matter of great debate between supporters and critics of intelligent design. The
Kitzmiller v. Dover Area School District court ruling held the latter to be the case.

[edit] Origin of the term

See also: Timeline of intelligent design
 Of Pandas and People was the first modern intelligent design book. Rethinking Schools magazine
characterizes it as a "creationist treatise dressed up to look like a legitimate discussion of science".[42]
 Use of the terms "creationism" versus "intelligent design" in sequential drafts of the book Of Pandas
and People[43]
The phrase "intelligent design" can be found in an 1847 issue of Scientific American,[44] in an 1850
book by Patrick Edward Dove,[45] and in an 1861 letter from Charles Darwin.[46] The phrase was used in an
address to the 1873 annual meeting of the British Association for the Advancement of Science by Paleyite
botanist George James Allman:
 No physical hypothesis founded on any indisputable fact has yet explained the origin of the
primordial protoplasm, and, above all, of its marvellous properties, which render evolution
possible—in heredity and in adaptability, for these properties are the cause and not the effect of
evolution. For the cause of this cause we have sought in vain among the physical forces which
surround us, until we are at last compelled to rest upon an independent volition, a far-seeing
intelligent design.[47]

The phrase can be found again in Humanism, a 1903 book by one of the founders of classical
pragmatism, F.C.S. Schiller: "It will not be possible to rule out the supposition that the process of evolution
may be guided by an intelligent design". A derivative of the phrase appears in the Macmillan Encyclopedia of
Philosophy (1967) in the article titled, "Teleological argument for the existence of God": "Stated most
succinctly, the argument runs: The world exhibits teleological order (design, adaptation). Therefore, it was
produced by an intelligent designer".[48] Robert Nozick (1974) wrote: "Consider now complicated patterns
which one would have thought would arise only through intelligent design".[49] The phrases "intelligent
design" and "intelligently designed" were used in a 1979 philosophy book Chance or Design? by James
Horigan[50] and the phrase "intelligent design" was used in a 1982 speech by Sir Fred Hoyle in his promotion
of panspermia.[51]
The modern use of the words "intelligent design", as a term intended to describe a field of inquiry,
began after the Supreme Court of the United States, in the case of Edwards v. Aguillard (1987), ruled that
creationism is unconstitutional in public school science curricula. A Discovery Institute report says that
Charles Thaxton, editor of Of Pandas and People, had picked the phrase up from a NASA scientist, and
thought "That's just what I need, it's a good engineering term".[52] In drafts of the book over one hundred
uses of the root word "creation", such as "creationism" and "creation science", were changed, almost without
exception, to "intelligent design",[23] while "creationists" was changed to "design proponents" or, in one
instance, "cdesign proponentsists". [sic][43] In June 1988 Thaxton held a conference titled "Sources of
Information Content in DNA" in Tacoma, Washington,[38] and in December decided to use the label
"intelligent design" for his new creationist movement.[53] Stephen C. Meyer was at the conference, and later
recalled that "the term came up".[54]
Of Pandas and People was published in 1989, and was the first book to make frequent use of the
phrases "intelligent design," "design proponents," and "design theory", thus representing the beginning of the
modern "intelligent design" movement.[55] "Intelligent design” was the most prominent of around fifteen new
terms it introduced as a new lexicon of creationist terminology to oppose evolution without using religious
language.[56] It was the first place where the phrase "intelligent design" appeared in its present use, as
stated both by its publisher Jon Buell,[57][58] and by William A. Dembski in his expert witness report.[59] The
book presented all of the basic arguments of intelligent design proponents before any research had been
done to support these arguments, and was actively promoted by creationists for public school use.[55]
Rethinking Schools magazine has criticized the book, saying it was a "creationist treatise" packaged to look
like a high quality science textbook, with a "glossy cover, full-color illustrations, and chapter titles such as
'Homology' and 'Genetics and Macroevolution'", with numerous "professionally prepared charts and
illustrations appear to show how concrete scientific evidence supports the existence of the unnamed
'designer'". Philosopher of science Michael Ruse believes the contents were "worthless and dishonest", and
it was described by an ACLU lawyer as a political tool aimed at students who did not "know science or
understand the controversy over evolution and creationism".[42]
[edit] Integral concepts
[edit] Irreducible complexity
Further information: Irreducible complexity

The concept of irreducible complexity was popularised by Michael Behe, in his 1996 book, Darwin's
Black Box.
The term "irreducible complexity" was introduced by biochemist Michael Behe in his 1996 book
Darwin's Black Box, though he had already described the concept in his contributions to the 1993 revised
edition of Of Pandas and People.[55] Behe defines it as "a single system which is composed of several well-
matched interacting parts that contribute to the basic function, wherein the removal of any one of the parts
causes the system to effectively cease functioning".[60]
 Behe uses the analogy of a mousetrap to illustrate this concept. A mousetrap consists of several
interacting pieces—the base, the catch, the spring and the hammer—all of which must be in place for the
mousetrap to work. Removal of any one piece destroys the function of the mousetrap. Intelligent design
advocates assert that natural selection could not create irreducibly complex systems, because the selectable
function is present only when all parts are assembled. Behe argued that irreducibly complex biological
mechanisms include the bacterial flagellum of E. coli, the blood clotting cascade, cilia, and the adaptive
immune system.[61][62]
Critics point out that the irreducible complexity argument assumes that the necessary parts of a
system have always been necessary and therefore could not have been added sequentially.[63][64] They
argue that something which is at first merely advantageous can later become necessary as other
components change. Furthermore, they argue, evolution often proceeds by altering preexisting parts or by
removing them from a system, rather than by adding them. This is sometimes called the "scaffolding
objection" by an analogy with scaffolding, which can support an "irreducibly complex" building until it is
complete and able to stand on its own.[65] Behe has acknowledged using "sloppy prose", and that his
"argument against Darwinism does not add up to a logical proof".[66] Irreducible complexity has remained a
popular argument among advocates of intelligent design; in the Dover trial, the court held that "Professor
Behe's claim for irreducible complexity has been refuted in peer-reviewed research papers and has been
rejected by the scientific community at large".[67]

[edit] Specified complexity

Further information: Specified complexity
In 1986 the creationist chemist Charles Thaxton used the term "specified complexity" from
information theory when claiming that messages transmitted by DNA in the cell were specified by
intelligence, and must have originated with an intelligent agent.[36] The intelligent design concept of
"specified complexity" was developed in the 1990s by mathematician, philosopher, and theologian William
Dembski. Dembski states that when something exhibits specified complexity (i.e., is both complex and
"specified", simultaneously), one can infer that it was produced by an intelligent cause (i.e., that it was
designed) rather than being the result of natural processes. He provides the following examples: "A single
letter of the alphabet is specified without being complex. A long sentence of random letters is complex
without being specified. A Shakespearean sonnet is both complex and specified".[68] He states that details
of living things can be similarly characterized, especially the "patterns" of molecular sequences in functional
biological molecules such as DNA.

William Dembski proposed the concept of specified complexity.[69]

 Dembski defines complex specified information (CSI) as anything with a less than 1 in 10150 chance
of occurring by (natural) chance. Critics say that this renders the argument a tautology: complex specified
information cannot occur naturally because Dembski has defined it thus, so the real question becomes
whether or not CSI actually exists in nature.[70][71][72]
The conceptual soundness of Dembski's specified complexity/CSI argument has been widely
discredited by the scientific and mathematical communities.[73][74][75] Specified complexity has yet to be
shown to have wide applications in other fields as Dembski asserts. John Wilkins and Wesley Elsberry
characterize Dembski's "explanatory filter" as eliminative, because it eliminates explanations sequentially:
first regularity, then chance, finally defaulting to design. They argue that this procedure is flawed as a model
for scientific inference because the asymmetric way it treats the different possible explanations renders it
prone to making false conclusions.[76]
Richard Dawkins, another critic of intelligent design, argues in The God Delusion that allowing for an
intelligent designer to account for unlikely complexity only postpones the problem, as such a designer would
need to be at least as complex.[77] Other scientists have argued that evolution through selection is better
able to explain the observed complexity, as is evident from the use of selective evolution to design certain
electronic, aeronautic and automotive systems which are considered problems too complex for human
"intelligent designers".[78]

[edit] Fine-tuned Universe

Further information: Fine-tuned Universe
Intelligent design proponents have also occasionally appealed to broader teleological arguments
outside of biology, most notably an argument based on the fine-tuning of universal constants that make
matter and life possible and which are argued not to be solely attributable to chance. These include the
values of fundamental physical constants, the relative strength of nuclear forces, electromagnetism, and
gravity between fundamental particles, as well as the ratios of masses of such particles. Intelligent design
proponent and Center for Science and Culture fellow Guillermo Gonzalez argues that if any of these values
were even slightly different, the universe would be dramatically different, making it impossible for many
chemical elements and features of the Universe, such as galaxies, to form.[79] Thus, proponents argue, an
intelligent designer of life was needed to ensure that the requisite features were present to achieve that
particular outcome.
Scientists have generally responded that this argument cannot be tested and is therefore not science
but metaphysics. Some scientists argue that even when taken as mere speculation, these arguments are
poorly supported by existing evidence.[80] Victor J. Stenger and other critics say both intelligent design and
the weak form of the anthropic principle are essentially a tautology; in his view, these arguments amount to
the claim that life is able to exist because the Universe is able to support life.[81][82][83] The claim of the
improbability of a life-supporting universe has also been criticized as an argument by lack of imagination for
assuming no other forms of life are possible. Life as we know it might not exist if things were different, but a
different sort of life might exist in its place. A number of critics also suggest that many of the stated variables
appear to be interconnected and that calculations made by mathematicians and physicists suggest that the
emergence of a universe similar to ours is quite probable.[84]
Proponent Granville Sewell argues that the evolution of complex forms of life represents a decrease
of entropy, and that it thus violates the second law of thermodynamics and so supports intelligent design.[85]
This, however, is a misapplication of thermodynamic principles.[86] The second law applies to closed
systems only. If Granville's argument were valid, living things could not be born and grow, as this also would
be a decrease in entropy. Neither evolution nor the growth of living things violates the second law of
thermodynamics because living things are not closed systems—they have external energy sources (e.g. food,
oxygen, sunlight) whose production requires an offsetting net increase in entropy.

[edit] Intelligent designer

Further information: Intelligent designer
Intelligent design arguments are formulated in secular terms and intentionally avoid identifying the
intelligent agent (or agents) they posit. Although they do not state that God is the designer, the designer is
often implicitly hypothesized to have intervened in a way that only a god could intervene. Dembski, in The
Design Inference, speculates that an alien culture could fulfill these requirements. The authoritative
description of intelligent design,[87] however, explicitly states that the Universe displays features of having
been designed. Acknowledging the paradox, Dembski concludes that "no intelligent agent who is strictly
physical could have presided over the origin of the universe or the origin of life".[88] The leading proponents
have made statements to their supporters that they believe the designer to be the Christian God, to the
exclusion of all other religions.[9][10][39]
Beyond the debate over whether intelligent design is scientific, a number of critics argue that existing
evidence makes the design hypothesis appear unlikely, irrespective of its status in the world of science. For
example, Jerry Coyne asks why a designer would "give us a pathway for making vitamin C, but then destroy
it by disabling one of its enzymes" (see pseudogene) and why he or she would not "stock oceanic islands
with reptiles, mammals, amphibians, and freshwater fish, despite the suitability of such islands for these
species". Coyne also points to the fact that "the flora and fauna on those islands resemble that of the nearest
mainland, even when the environments are very different" as evidence that species were not placed there by
a designer.[89] Previously, in Darwin's Black Box, Behe had argued that we are simply incapable of
understanding the designer's motives, so such questions cannot be answered definitively. Odd designs
could, for example, "have been placed there by the designer ... for artistic reasons, to show off, for some as-
yet undetectable practical purpose, or for some unguessable reason". Coyne responds that in light of the
evidence, "either life resulted not from intelligent design, but from evolution; or the intelligent designer is a
cosmic prankster who designed everything to make it look as though it had evolved".[89]
Asserting the need for a designer of complexity also raises the question "What designed the
designer?"[90] Intelligent design proponents say that the question is irrelevant to or outside the scope of
intelligent design.[91] Richard Wein counters that the unanswered questions an explanation creates "must be
balanced against the improvements in our understanding which the explanation provides. Invoking an
unexplained being to explain the origin of other beings (ourselves) is little more than question-begging. The
new question raised by the explanation is as problematic as the question which the explanation purports to
answer".[72] Richard Dawkins sees the assertion that the designer does not need to be explained, not as a
contribution to knowledge, but as a thought-terminating cliché.[92][93] In the absence of observable,
measurable evidence, the very question "What designed the designer?" leads to an infinite regression from
which intelligent design proponents can only escape by resorting to religious creationism or logical
contradiction.[94]

[edit] Movement
Further information: Intelligent design movement
 The Discovery Institute's Center for the Renewal of Science and Culture used banners based on
"The Creation of Adam" from the Sistine Chapel. Later it used a less religious image, then was renamed the
Center for Science and Culture.[95]
The intelligent design movement is a direct outgrowth of the creationism of the 1980s.[6] The
scientific and academic communities, along with a U.S. federal court, view intelligent design as either a form
of creationism or as a direct descendant that is closely intertwined with traditional creationism;[18] [19][96]
[97][98][99] and several authors explicitly refer to it as "intelligent design creationism".[6][100][101][102]
The movement is headquartered in the Center for Science and Culture (CSC), established in 1996 as
the creationist wing of the Discovery Institute to promote a religious agenda[103] calling for broad social,
academic and political changes. The Discovery Institute's intelligent design campaigns have been staged
primarily in the United States, although efforts have been made in other countries to promote intelligent
design. Leaders of the movement say intelligent design exposes the limitations of scientific orthodoxy and of
the secular philosophy of naturalism. Intelligent design proponents allege that science should not be limited
to naturalism and should not demand the adoption of a naturalistic philosophy that dismisses out-of-hand any
explanation which contains a supernatural cause. The overall goal of the movement is to "defeat [the]
materialist world view" represented by the theory of evolution in favor of "a science consonant with Christian
and theistic convictions".[103]
Phillip E. Johnson stated that the goal of intelligent design is to cast creationism as a scientific
concept.[40][104] All leading intelligent design proponents are fellows or staff of the Discovery Institute and
its Center for Science and Culture.[105] Nearly all intelligent design concepts and the associated movement
are the products of the Discovery Institute, which guides the movement and follows its wedge strategy while
conducting its Teach the Controversy campaign and their other related programs.
Leading intelligent design proponents have made conflicting statements regarding intelligent design.
In statements directed at the general public, they say intelligent design is not religious; when addressing
conservative Christian supporters, they state that intelligent design has its foundation in the Bible.[104]
Recognizing the need for support, the institute affirms its Christian, evangelistic orientation: "Alongside a
focus on influential opinion-makers, we also seek to build up a popular base of support among our natural
constituency, namely, Christians. We will do this primarily through apologetics seminars. We intend these to
encourage and equip believers with new scientific evidences that support the faith, as well as to 'popularize'
our ideas in the broader culture."[103]
Barbara Forrest, an expert who has written extensively on the movement, describes this as being
due to the Discovery Institute's obfuscating its agenda as a matter of policy. She has written that the
movement's "activities betray an aggressive, systematic agenda for promoting not only intelligent design
creationism, but the religious world-view that undergirds it".[106]
[edit] Religion and leading proponents
Although arguments for intelligent design are formulated in secular terms and intentionally avoid
positing the identity of the designer,[107] the majority of principal intelligent design advocates are publicly
religious Christians who have stated that in their view the designer proposed in intelligent design is the
Christian conception of God. Stuart Burgess, Phillip E. Johnson, William Dembski, and Stephen C. Meyer are
evangelical Protestants, and Michael Behe is a Roman Catholic, while Jonathan Wells is a member of the
Unification Church. Phillip E. Johnson has stated that cultivating ambiguity by employing secular language in
arguments that are carefully crafted to avoid overtones of theistic creationism is a necessary first step for
ultimately reintroducing the Christian concept of God as the designer. Johnson explicitly calls for intelligent
design proponents to obfuscate their religious motivations so as to avoid having intelligent design identified
"as just another way of packaging the Christian evangelical message".[108] Johnson emphasizes that "the
first thing that has to be done is to get the Bible out of the discussion"; "after we have separated materialist
prejudice from scientific fact [...] only then can 'biblical issues' be discussed".[109]
The strategy of deliberately disguising the religious intent of intelligent design has been described by
William Dembski in The Design Inference.[110] In this work Dembski lists a god or an "alien life force" as two
possible options for the identity of the designer; however, in his book Intelligent Design: The Bridge Between
Science and Theology, Dembski states that "Christ is indispensable to any scientific theory, even if its
practitioners don't have a clue about him. The pragmatics of a scientific theory can, to be sure, be pursued
without recourse to Christ. But the conceptual soundness of the theory can in the end only be located in
Christ."[111] Dembski also stated, "ID is part of God's general revelation [...] Not only does intelligent design
rid us of this ideology (materialism), which suffocates the human spirit, but, in my personal experience, I've
found that it opens the path for people to come to Christ".[112] Both Johnson and Dembski cite the Bible's
Gospel of John as the foundation of intelligent design.[39][104]
 Barbara Forrest contends such statements reveal that leading proponents see intelligent design as
essentially religious in nature, not merely a scientific concept that has implications with which their personal
religious beliefs happen to coincide.[113] She writes that the leading proponents of intelligent design are
closely allied with the ultra-conservative Christian Reconstructionism movement. She lists connections of
(current and former) Discovery Institute Fellows Phillip Johnson, Charles Thaxton, Michael Behe, Richard
Weikart, Jonathan Wells and Francis Beckwith to leading Christian Reconstructionist organizations, and the
extent of the funding provided the Institute by Howard Ahmanson Jr., a leading figure in the Reconstructionist
movement.[114]

[edit] Reaction from other creationist groups

Not all creationist organizations have embraced the intelligent design movement. Hugh Ross of
Reasons to Believe, a proponent of Old Earth creationism, believes that the efforts of intelligent design
proponents to divorce the concept from Biblical Christianity make its hypothesis too vague. In 2002 he wrote:
"Winning the argument for design without identifying the designer yields, at best, a sketchy origins model.
Such a model makes little if any positive impact on the community of scientists and other scholars… The time
is right for a direct approach, a single leap into the origins fray. Introducing a biblically based, scientifically
verifiable creation model represents such a leap."[115]
Likewise, two of the most prominent Young Earth creationism organizations in the world have
attempted to distinguish their views from intelligent design. Henry M. Morris of the Institute for Creation
Research (ICR) wrote, in 1999, that ID, "even if well-meaning and effectively articulated, will not work! It has
often been tried in the past and has failed, and it will fail today. The reason it won't work is because it is not
the Biblical method.” According to Morris: “The evidence of intelligent design… must be either followed by or
accompanied by a sound presentation of true Biblical creationism if it is to be meaningful and lasting."[116] In
2002, Carl Wieland of Answers in Genesis (AiG) criticized design advocates who, though well-intentioned,
"left the Bible out of it" and thereby unwittingly aided and abetted the modern rejection of the Bible. Wieland
explained that "AiG's major 'strategy' is to boldly, but humbly, call the church back to its Biblical foundations…
[so] we neither count ourselves a part of this movement nor campaign against it."[117]

[edit] Polls
Several surveys were conducted prior to the December 2005 decision in Kitzmiller v. Dover, which
sought to determine the level of support for intelligent design among certain groups. According to a 2005
Harris poll, 10% of adults in the United States viewed human beings as "so complex that they required a
powerful force or intelligent being to help create them".[118] Although Zogby polls commissioned by the
Discovery Institute show more support, these polls suffer from considerable flaws, such as having a very low
response rate (248 out of 16,000), being conducted on behalf of an organization with an expressed interest in
the outcome of the poll, and containing leading questions.[119][120][121]
A May 2005 survey of nearly 1500 physicians in the United States conducted by the Louis Finkelstein
Institute and HCD Research showed that 63% of the physicians agreed more with evolution than with
intelligent design.[122]
A series of Gallup polls in the United States from 1982 through 2008 on "Evolution, Creationism,
Intelligent Design" found support for "human beings have developed over millions of years from less
advanced formed of life, but God guided the process" of between 35% and 40%, support for "God created
human beings in pretty much their present form at one time within the last 10,000 years or so" varied from
43% to 47%, and support for "human beings have developed over millions of years from less advanced
formed of life, but God had no part in the process" varied from 9% to 14%. The polls also noted answers to a
series of more detailed questions.[123]
[edit] Film
The film Expelled: No Intelligence Allowed sparked further controversy in 2008. This documentary,
hosted by Ben Stein, spends much time focusing on professors who have been asked to leave or have left
numerous institutions because, the film insinuates, of their beliefs in Intelligent Design. One of the film's first
screenings resulted in Paul "PZ" Myers, an interviewee in the film, being asked to leave the theater. There
have also been allegations from some interviewees that interviews were recorded many times in order to get
the exact phrasing required by the producer. The production company, Premise Media, also has helped
finance some religious films such as The Passion of the Christ. [124] [125]

[edit] Creating and teaching the controversy

Main article: Teach the Controversy
A key strategy of the intelligent design movement is convincing the general public that there is a
debate among scientists about whether life evolved. The intelligent design movement creates this
controversy in order to convince the public, politicians and cultural leaders that schools should "Teach the
Controversy".[126] But in fact, there is no such controversy in the scientific community; the scientific
consensus is that life evolved.[127][128][129] Intelligent design is widely viewed as a stalking horse for its
proponents' campaign against what they say is the materialist foundation of science, which they argue leaves
no room for the possibility of God.[130][131]
Advocates of intelligent design seek to keep God and the Bible out of the discussion, and present
intelligent design in the language of science as a scientific hypothesis.[107][109] However, among a very
significant proportion of the general public in the United States the major concern is whether conventional
evolutionary biology is compatible with belief in God and in the Bible, and how this issue is taught in schools.
[132] The public controversy was given widespread media coverage in the United States, particularly during
the Kitzmiller v. Dover trial in late 2005 and after President George W. Bush expressed support for the idea of
teaching intelligent design alongside evolution in August 2005. In response to Bush’s statement and the
pending federal trial, Time magazine ran an eight-page cover story on the Evolution Wars in which they
examined the issue of teaching intelligent design in the classroom.[133][134] The cover of the magazine
featured a parody of The Creation of Adam from the Sistine Chapel. Rather than pointing at Adam,
Michelangelo’s God points at the image of a chimpanzee contemplating the caption which read " The push to
teach "intelligent design" raises a question: Does God have a place in science class?" .[135] In the Kitzmiller
v. Dover case, the court ruled that intelligent design was a religious and creationist position, finding that God
and intelligent design were both distinct from the material that should be covered in a science class.[5]
Empirical science uses the scientific method to create a posteriori knowledge based on observation
and repeated testing of hypotheses and theories. Intelligent design proponents seek to change this
fundamental basis of science[136] by eliminating "methodological naturalism" from science[137] and
replacing it with what the leader of the intelligent design movement, Phillip E. Johnson, calls "theistic
realism".[138] Some have called this approach "methodological supernaturalism", which means belief in a
transcendent, nonnatural dimension of reality inhabited by a transcendent, nonnatural deity.[139] Intelligent
design proponents argue that naturalistic explanations fail to explain certain phenomena and that
supernatural explanations provide a very simple and intuitive explanation for the origins of life and the
universe.[140] Proponents say evidence exists in the forms of irreducible complexity and specified
complexity that cannot be explained by natural processes.[1] They also hold that religious neutrality requires
the teaching of both evolution and intelligent design in schools, saying that teaching only evolution unfairly
discriminates against those holding creationist beliefs. Teaching both, they argue, allows for the possibility of
religious belief, without causing the state to actually promote such beliefs. Many intelligent design followers
believe that "Scientism" is itself a religion that promotes secularism and materialism in an attempt to erase
theism from public life, and they view their work in the promotion of intelligent design as a way to return
religion to a central role in education and other public spheres. Some allege that this larger debate is often
the subtext for arguments made over intelligent design, though others note that intelligent design serves as
an effective proxy for the religious beliefs of prominent intelligent design proponents in their efforts to
advance their religious point of view within society.[141][142][143]
Intelligent design has not presented a credible scientific case and is an attempt to teach religion in
public schools, substituting public support for scientific research.[144] If the argument to give "equal time for
all theories" were actually practiced, there would be no logical limit to the number of mutually incompatible
supernatural "theories" regarding the origins and diversity of life to be taught in the public school system,
including intelligent design parodies such as the Flying Spaghetti Monster "theory"; intelligent design does
not provide a mechanism for discriminating among them. Philosopher of biology Elliott Sober, for example,
states that intelligent design is not falsifiable because "[d]efenders of ID always have a way out".[145][146]
Intelligent design proponent Michael Behe concedes "You can't prove intelligent design by experiment".[132]
Intelligent design proponents cannot legitimately infer that an intelligent designer is behind the part of
the process that is not understood scientifically, since they have not shown that anything supernatural has
occurred. The inference that an intelligent designer created life on Earth, which advocate William Dembski
has said could alternately be an "alien" life force,[110] has been compared to the a priori claim that aliens
helped the ancient Egyptians build the pyramids.[147][148] In both cases, the effect of this outside
intelligence is not repeatable, observable or falsifiable, and it violates the principle of parsimony. From a
strictly empirical standpoint, one may list what is known about Egyptian construction techniques, but one
must admit ignorance about exactly how the Egyptians built the pyramids.
Supporters of intelligent design have also reached out to other faith groups with similar accounts of
creation with the hope that the broader coalition will have greater influence in supporting science education
that does not contradict their religious views.[140] Many religious bodies have responded by expressing
support for evolution. The Roman Catholic church has stated that religious faith is fully compatible with
science, which is limited to dealing only with the natural world[149] — a position described by the term theistic
evolution.[150] As well as pointing out that intelligent design is not science, they also reject it for various
philosophical and theological reasons.[151][152] The arguments of intelligent design have been directly
challenged by the over 10,000 clergy who signed the Clergy Letter Project. Prominent scientists who strongly
express religious faith, such as the astronomer George Coyne and the biologist Ken Miller, have been at the
forefront of opposition to intelligent design. While creationist organizations have welcomed intelligent
design's support against naturalism, they have also been critical of its refusal to identify the designer,[153]
[154][155] and have pointed to previous failures of the same argument.[156]
Rabbi Natan Slifkin directly criticized the advocates of intelligent design as presenting a perspective
of God that is dangerous to religion.[157] Those who promote it as parallel to religion, he asserts, do not truly
understand it. Slifkin criticizes intelligent design's advocacy of teaching their perspective in biology classes,
wondering why no one claims that God's hand should be taught in other secular classes, such as history,
physics or geology. Slifkin also asserts that the intelligent design movement is inordinately concerned with
portraying God as "in control" when it comes to things that cannot be easily explained by science, but not in
control in respect to things which can be explained by scientific theory.[157] Kenneth Miller expressed a view
similar to Slifkin's: "[T]he struggles of the Intelligent Design movement are best understood as clamorous and
disappointing double failures - rejected by science because they do not fit the facts, and having failed religion
because they think too little of God.[158]
[edit] Defining science
The scientific method is a body of techniques for investigating phenomena and acquiring new
knowledge of the natural world without assuming the existence or nonexistence of the supernatural, an
approach sometimes called methodological naturalism. Intelligent design proponents believe that this can be
equated to materialist metaphysical naturalism, and have often said that not only is their own position
scientific, but it is even more scientific than evolution, and that they want a redefinition of science as a
revived natural theology or natural philosophy to allow "non-naturalistic theories such as intelligent design".
[159] This presents a demarcation problem, which in the philosophy of science is about how and where to
draw the lines around science.[160] For a theory to qualify as scientific,[161][162][163] it is expected to be:
• Consistent
• Parsimonious (sparing in its proposed entities or explanations, see Occam's Razor)
• Useful (describes and explains observed phenomena, and can be used predictively)
• Empirically testable and falsifiable (see Falsifiability)
• Based on multiple observations, often in the form of controlled, repeated experiments
• Correctable and dynamic (modified in the light of observations that do not support it)
• Progressive (refines previous theories)
• Provisional or tentative (is open to experimental checking, and does not assert certainty)
For any theory, hypothesis or conjecture to be considered scientific, it must meet most, and ideally
all, of these criteria. The fewer criteria are met, the less scientific it is; and if it meets only a few or none at all,
then it cannot be treated as scientific in any meaningful sense of the word. Typical objections to defining
intelligent design as science are that it lacks consistency,[164] violates the principle of parsimony,[165] is not
scientifically useful,[166] is not falsifiable,[167] is not empirically testable,[168] and is not correctable,
dynamic, provisional or progressive.[169][170][171]
Critics also say that the intelligent design doctrine does not meet the Daubert Standard,[172] the
criteria for scientific evidence mandated by the US Supreme Court. The Daubert Standard governs which
evidence can be considered scientific in United States federal courts and most state courts. Its four criteria
are:
• The theoretical underpinnings of the methods must yield testable predictions by means of
which the theory could be falsified.
• The methods should preferably be published in a peer-reviewed journal.
• There should be a known rate of error that can be used in evaluating the results.
• The methods should be generally accepted within the relevant scientific community.
In Kitzmiller v. Dover Area School District, using these criteria and others mentioned above, Judge
Jones ruled that "... we have addressed the seminal question of whether ID is science. We have concluded
that it is not, and moreover that ID cannot uncouple itself from its creationist, and thus religious,
antecedents".
Against this, the philosopher Thomas Nagel argues that intelligent design is very different from
creation science, in that it does not depend on distortion of evidence, or on the assumption that it is immune
to empirical evidence. It depends only on the idea that the hypothesis of a designer makes sense. Whatever
the merits of the positions, he argues that it is a scientific disagreement, not a disagreement between science
and something else.[173]
[edit] Peer review
The failure to follow the procedures of scientific discourse and the failure to submit work to the
scientific community that withstands scrutiny have weighed against intelligent design's being considered as
valid science.[174] The intelligent design movement has not published a properly peer-reviewed article in a
scientific journal.[174]
Intelligent design, by appealing to a supernatural agent, directly conflicts with the principles of
science, which limit its inquiries to empirical, observable and ultimately testable data and which require
explanations to be based on empirical evidence. Dembski, Behe and other intelligent design proponents say
bias by the scientific community is to blame for the failure of their research to be published.[175] Intelligent
design proponents believe that their writings are rejected for not conforming to purely naturalistic, non-
supernatural mechanisms rather than because their research is not up to "journal standards", and that the
merit of their articles is overlooked. Some scientists describe this claim as a conspiracy theory.[176] Michael
Shermer has rebutted the claim, noting "Anyone who thinks that scientists do not question Darwinism has
never been to an evolutionary conference." He noted that scientists such as Joan Roughgarden and Lynn
Margulis have challenged certain Darwinist theories and offered explanations of their own and despite this
they "have not been persecuted, shunned, fired or even expelled. Why? Because they are doing science, not
religion."[177] The issue that supernatural explanations do not conform to the scientific method became a
sticking point for intelligent design proponents in the 1990s, and is addressed in the wedge strategy as an
aspect of science that must be challenged before intelligent design can be accepted by the broader scientific
community.
The debate over whether intelligent design produces new research, as any scientific field must, and
has legitimately attempted to publish this research, is extremely heated. Both critics and advocates point to
numerous examples to make their case. For instance, the Templeton Foundation, a former funder of the
Discovery Institute and a major supporter of projects seeking to reconcile science and religion, says that it
asked intelligent design proponents to submit proposals for actual research, but none were ever submitted.
Charles L. Harper Jr., foundation vice-president, said: "From the point of view of rigor and intellectual
seriousness, the intelligent design people don't come out very well in our world of scientific review".[178]
The only article published in a peer-reviewed scientific journal that made a case for intelligent design
was quickly withdrawn by the publisher for having circumvented the journal's peer-review standards.[179]
Written by the Discovery Institute's Center for Science & Culture Director Stephen C. Meyer, it appeared in
the peer-reviewed journal Proceedings of the Biological Society of Washington in August 2004.[180] The
article was a literature review, which means that it did not present any new research, but rather culled
quotations and claims from other papers to argue that the Cambrian explosion could not have happened by
natural processes. The choice of venue for this article was also considered problematic, because it was so
outside the normal subject matter (see Sternberg peer review controversy[181]). Dembski has written that
"perhaps the best reason [to be skeptical of his ideas] is that intelligent design has yet to establish itself as a
thriving scientific research program."[182] In a 2001 interview, Dembski said that he stopped submitting to
peer-reviewed journals because of their slow time-to-print and that he makes more money from publishing
books.[183]
In the Dover trial, the judge found that intelligent design features no scientific research or testing.
[184] There, intelligent design proponents cited just one paper, on simulation modeling of evolution by Behe
and Snoke,[185] which mentioned neither irreducible complexity nor intelligent design and which Behe
admitted did not rule out known evolutionary mechanisms.[186] Michael Lynch called the conclusions of the
article "an artifact of unwarranted biological assumptions, inappropriate mathematical modeling, and faulty
logic".[187] In sworn testimony, however, Behe said: "There are no peer reviewed articles by anyone
advocating for intelligent design supported by pertinent experiments or calculations which provide detailed
rigorous accounts of how intelligent design of any biological system occurred".[188] As summarized by the
judge, Behe conceded that there are no peer-reviewed articles supporting his claims of intelligent design or
irreducible complexity. In his ruling, the judge wrote: "A final indicator of how ID has failed to demonstrate
scientific warrant is the complete absence of peer-reviewed publications supporting the theory".[174]
The Discovery Institute insists that a number of intelligent design articles have been published in
peer-reviewed journals,[189] including in its list the two articles mentioned above. Critics, largely members of
the scientific community, reject this claim, stating that no established scientific journal has yet published an
intelligent design article. Instead, intelligent design proponents have set up their own journals with "peer
review" which lack impartiality and rigor,[190] consisting entirely of intelligent design supporters.[191]

[edit] Intelligence as an observable quality

The phrase intelligent design makes use of an assumption of the quality of an observable
intelligence, a concept that has no scientific consensus definition. William Dembski, for example, has written
that "Intelligence leaves behind a characteristic signature". The characteristics of intelligence are assumed
by intelligent design proponents to be observable without specifying what the criteria for the measurement of
intelligence should be. Dembski, instead, asserts that "in special sciences ranging from forensics to
archaeology to SETI (the Search for Extraterrestrial Intelligence), appeal to a designing intelligence is
indispensable".[192] How this appeal is made and what this implies as to the definition of intelligence are
topics left largely unaddressed. Seth Shostak, a researcher with the SETI Institute, disputed Dembski's
comparison of SETI and intelligent design, saying that intelligent design advocates base their inference of
design on complexity—the argument being that some biological systems are too complex to have been made
by natural processes—while SETI researchers are looking primarily for artificiality.[193]
 Critics say that the design detection methods proposed by intelligent design proponents are radically
different from conventional design detection, undermining the key elements that make it possible as
legitimate science. Intelligent design proponents, they say, are proposing both searching for a designer
without knowing anything about that designer's abilities, parameters, or intentions (which scientists do know
when searching for the results of human intelligence), as well as denying the very distinction between
natural/artificial design that allows scientists to compare complex designed artifacts against the background
of the sorts of complexity found in nature.[194]
As a means of criticism, certain skeptics have pointed to a challenge of intelligent design derived
from the study of artificial intelligence. The criticism is a counter to intelligent design claims about what
makes a design intelligent, specifically that "no preprogrammed device can be truly intelligent, that
intelligence is irreducible to natural processes".[195] This claim is similar in type to an assumption of
Cartesian dualism that posits a strict separation between "mind" and the material Universe. However, in
studies of artificial intelligence, while there is an implicit assumption that supposed "intelligence" or creativity
of a computer program is determined by the capabilities given to it by the computer programmer, artificial
intelligence need not be bound to an inflexible system of rules. Rather, if a computer program can access
randomness as a function, this effectively allows for a flexible, creative, and adaptive intelligence.
Evolutionary algorithms, a subfield of machine learning (itself a subfield of artificial intelligence), have been
used to mathematically demonstrate that randomness and selection can be used to "evolve" complex, highly
adapted structures that are not explicitly designed by a programmer. Evolutionary algorithms use the
Darwinian metaphor of random mutation, selection and the survival of the fittest to solve diverse
mathematical and scientific problems that are usually not solvable using conventional methods. Intelligence
derived from randomness is essentially indistinguishable from the "innate" intelligence associated with
biological organisms, and poses a challenge to the intelligent design conception that intelligence itself
necessarily requires a designer. Cognitive science continues to investigate the nature of intelligence along
these lines of inquiry. The intelligent design community, for the most part, relies on the assumption that
intelligence is readily apparent as a fundamental and basic property of complex systems.[196]

[edit] Arguments from ignorance

Eugenie Scott, along with Glenn Branch and other critics, has argued that many points raised by
intelligent design proponents are arguments from ignorance.[197] In the argument from ignorance, a lack of
evidence for one view is erroneously argued to constitute proof of the correctness of another view. Scott and
Branch say that intelligent design is an argument from ignorance because it relies on a lack of knowledge for
its conclusion: lacking a natural explanation for certain specific aspects of evolution, we assume intelligent
cause. They contend most scientists would reply that the unexplained is not unexplainable, and that "we
don't know yet" is a more appropriate response than invoking a cause outside science.[197] Particularly,
Michael Behe's demands for ever more detailed explanations of the historical evolution of molecular systems
seem to assume a false dichotomy, where either evolution or design is the proper explanation, and any
perceived failure of evolution becomes a victory for design. Scott and Branch also contend that the
supposedly novel contributions proposed by intelligent design proponents have not served as the basis for
any productive scientific research.
Intelligent design has also been characterized as a "god of the gaps" argument, which has the
following form:
• There is a gap in scientific knowledge.
• The gap is filled with acts of God (or intelligent designer) and therefore proves the
existence of God (or intelligent designer).
 A god-of-the-gaps argument is the theological version of an argument from ignorance. A key feature
of this type of argument is that it merely answers outstanding questions with explanations (often
supernatural) that are unverifiable and ultimately themselves subject to unanswerable questions.[198]

[edit] Kitzmiller trial

Main article: Kitzmiller v. Dover Area School District
Kitzmiller v. Dover Area School District was the first direct challenge brought in the United States
federal courts against a public school district that required the presentation of intelligent design as an
alternative to evolution. The plaintiffs successfully argued that intelligent design is a form of creationism, and
that the school board policy thus violated the Establishment Clause of the First Amendment to the United
States Constitution.[199]
Eleven parents of students in Dover, Pennsylvania, sued the Dover Area School District over a
statement that the school board required be read aloud in ninth-grade science classes when evolution was
taught. The plaintiffs were represented by the American Civil Liberties Union (ACLU), Americans United for
Separation of Church and State (AU) and Pepper Hamilton LLP. The National Center for Science Education
(NCSE) acted as consultants for the plaintiffs. The defendants were represented by the Thomas More Law
Center.[200] The suit was tried in a bench trial from September 26, 2005 to November 4, 2005 before Judge
John E. Jones III. Ken Miller, Kevin Padian, Brian Alters, Robert Pennock, Barbara Forrest and John Haught
served as expert witnesses for the prosecution. Michael Behe, Steve Fuller and Scott Minnich served as
expert witnesses for the defense.
On December 20, 2005 Judge Jones issued his 139-page findings of fact and decision, ruling that
the Dover mandate was unconstitutional, and barring intelligent design from being taught in Pennsylvania's
Middle District public school science classrooms. The eight Dover school board members who voted for the
intelligent design requirement were all defeated in a November 8, 2005 election by challengers who opposed
the teaching of intelligent design in a science class, and the current school board president stated that the
board does not intend to appeal the ruling.[201]
In his finding of facts, Judge Jones made the following condemnation of the Teach the Controversy
strategy:
Moreover, ID’s backers have sought to avoid the scientific scrutiny which we have now
determined that it cannot withstand by advocating that the controversy, but not ID itself, should
be taught in science class. This tactic is at best disingenuous, and at worst a canard.

[edit] Reaction
Judge Jones himself anticipated that his ruling would be criticized, saying in his decision that:
Those who disagree with our holding will likely mark it as the product of an activist judge. If so,
they will have erred as this is manifestly not an activist Court. Rather, this case came to us as
the result of the activism of an ill-informed faction on a school board, aided by a national public
interest law firm eager to find a constitutional test case on ID, who in combination drove the
Board to adopt an imprudent and ultimately unconstitutional policy. The breathtaking inanity of
the Board's decision is evident when considered against the factual backdrop which has now
been fully revealed through this trial. The students, parents, and teachers of the Dover Area
School District deserved better than to be dragged into this legal maelstrom, with its resulting
utter waste of monetary and personal resources.
 As Jones had predicted, John G. West, Associate Director of the Center for Science and Culture at
Discovery Institute, said:
The Dover decision is an attempt by an activist federal judge to stop the spread of a scientific
idea and even to prevent criticism of Darwinian evolution through government-imposed
censorship rather than open debate, and it won't work. He has conflated Discovery Institute's
position with that of the Dover school board, and he totally misrepresents intelligent design and
the motivations of the scientists who research it.[202]

Newspapers have noted with interest that the judge is "a Republican and a churchgoer".[203][204]
[205][206]
Subsequently, the decision has been examined in a search for flaws and conclusions, partly by
intelligent design supporters aiming to avoid future defeats in court. In the Spring of 2007 the University of
Montana Law review published three articles.[207] In the first, David K. DeWolf, John G. West and Casey
Luskin, all of the Discovery Institute, argued that intelligent design is a valid scientific theory, the Jones court
should not have addressed the question of whether it was a scientific theory, and that the Kitzmiller decision
will have no effect at all on the development and adoption of intelligent design as an alternative to standard
evolutionary theory.[134] In the second Peter Irons responded, arguing that the decision was extremely well
reasoned and spells the death knell for the intelligent design efforts to introduce creationism in public
schools,[208] while in the third, DeWolf et al. answer the points made by Irons.[209] However, fear of a
similar lawsuit has resulted in other school boards abandoning intelligent design "teach the controversy"
proposals.[6]
 In April 2010, the American Academy of Religion issued Guidelines for Teaching About Religion in K‐
12 Public Schools in the United States which included guidance that creation science or intelligent design
should not be taught in science classes, as "Creation science and intelligent design represent worldviews
that fall outside of the realm of science that is defined as (and limited to) a method of inquiry based on
gathering observable and measurable evidence subject to specific principles of reasoning." However, they,
as well as other "worldviews that focus on speculation regarding the origins of life represent another
important and relevant form of human inquiry that is appropriately studied in literature or social sciences
courses. Such study, however, must include a diversity of worldviews representing a variety of religious and
philosophical perspectives and must avoid privileging one view as more legitimate than others."[210]

[edit] Status outside the United States

[edit] Europe
In June 2007 the Council of Europe's "Committee on Culture, Science and Education" issued a
report, The dangers of creationism in education, which states "Creationism in any of its forms, such as
'intelligent design', is not based on facts, does not use any scientific reasoning and its contents are
pathetically inadequate for science classes."[211] In describing the dangers posed to education by teaching
creationism, it described intelligent design as "anti-science" and involving "blatant scientific fraud" and
"intellectual deception" that "blurs the nature, objectives and limits of science" and links it and other forms of
creationism to denialism. On October 4, 2007, the Council of Europe's Parliamentary Assembly approved a
resolution stating that schools should "resist presentation of creationist ideas in any discipline other than
religion", including "intelligent design" which it described as "the latest, more refined version of creationism",
"presented in a more subtle way". The resolution emphasises that the aim of the report is not to question or
to fight a belief, but to "warn against certain tendencies to pass off a belief as science".[212]
In the United Kingdom, public education includes Religious Education as a compulsory subject, and
many "faith schools" that teach the ethos of particular denominations. When it was revealed that a group
called Truth in Science had distributed DVDs produced by the Discovery Institute affiliate Illustra Media[213]
featuring Discovery Institute fellows making the case for design in nature,[214] and claimed they were being
used by 59 schools,[215] the Department for Education and Skills (DfES) stated that "Neither creationism nor
intelligent design are taught as a subject in schools, and are not specified in the science curriculum" (part of
the National Curriculum which does not apply to independent schools or to Education in Scotland).[216][217]
The DfES subsequently stated that "Intelligent design is not a recognised scientific theory; therefore, it is not
included in the science curriculum", but left the way open for it to be explored in religious education in relation
to different beliefs, as part of a syllabus set by a local Standing Advisory Council on Religious Education.
[218] In 2006 the Qualifications and Curriculum Authority produced a Religious Education model unit in which
pupils can learn about religious and nonreligious views about creationism, intelligent design and evolution by
natural selection.[219][220]
On June 25, 2007, the UK Government responded to an e-Petition by saying that creationism and
intelligent design should not be taught as science, though teachers would be expected to answer pupils'
questions within the standard framework of established scientific theories.[221] Detailed government
"Creationism teaching guidance" for schools in England was published on September 18, 2007. It states that
"Intelligent design lies wholly outside of science", has no underpinning scientific principles, or explanations,
and is not accepted by the science community as a whole. Though it should not be taught as science,
"questions about creationism and intelligent design which arise in science lessons, for example as a result of
media coverage, could provide the opportunity to explain or explore why they are not considered to be
scientific theories and, in the right context, why evolution is considered to be a scientific theory". However,
"Teachers of subjects such as RE, history or citizenship may deal with creationism and intelligent design in
their lessons".[13]
The British Centre for Science Education lobbying group has the goal of "countering creationism
within the UK" and has been involved in government lobbying in the UK in this regard.[222] However, in
Northern Ireland the Democratic Unionist Party claims that the revised curriculum provides an opportunity for
alternative theories to be taught, and has sought assurances that pupils will not lose marks if they give
creationist or intelligent design answers to science questions.[223] In Lisburn the DUP has arranged that the
City Council will write to post primary schools asking what their plans are to develop teaching material in
relation to "creation, intelligent design and other theories of origin".[224]
Plans by Dutch Education Minister Maria van der Hoeven to "stimulate an academic debate" on the
subject in 2005 caused a severe public backlash.[225] After the 2007 elections she was succeeded by
Ronald Plasterk, described as a "molecular geneticist, staunch atheist and opponent of intelligent design".
[226] As a reaction on this situation in the Netherlands, in Belgium the President of the Flemish Catholic
Educational Board (VSKO) Mieke Van Hecke declared that: "Catholic scientists already accepted the theory
of evolution for a long time and that intelligent design and creationism doesn't belong in Flemish Catholic
schools. It's not the tasks of the politics to introduce new ideas, that's task and goal of science."[227]

[edit] Relation to Islam

Creationism has strong political support in many Islamic countries, and antievolutionary views are
mainstream among academic theologians and scientists.[228] In general, Muslim creationists have partnered
with the Institute for Creation Research for ideas and materials which they adapted to their own theological
positions. Similarly, some use was made of intelligent design antievolution resources. Muzaffar Iqbal, a
notable Muslim in Canada, signed the Scientific Dissent list of the Discovery Institute.[229] Ideas similar to
intelligent design have been considered respected intellectual options among Muslims, and in Turkey many
intelligent design books have been translated. In Istanbul in 2007, public meetings promoting intelligent
design were sponsored by the local government,[228] and David Berlinski of the Discovery Institute was the
keynote speaker at a meeting in May 2007.[230]

[edit] Australia
The status of intelligent design in Australia is somewhat similar to that in the UK (see: Education in
Australia). When the former Australian Federal Education Minister, Brendan Nelson, raised the notion of
intelligent design being taught in science classes, the public outcry caused the minister to quickly concede
that the correct forum for intelligent design, if it were to be taught, is in religious or philosophy classes.[231]

[edit] See also

• Abiogenesis • List of works on Creationism
• Argument from intelligent design portal
poor design • Neo-Creationism
• Cosmological • Old Earth
argument creationism
• Creation • Santorum
science Amendment
• Flock of Dodos • Watchmaker
• Intelligent analogy
 falling

[edit] Notes
1. ^ a b c "Top Questions-1.What is the theory of intelligent design?". Discovery Institute.
http://www.discovery.org/csc/topQuestions.php#questionsAboutIntelligentDesign. Retrieved 2007-
05-13.
2. ^ "Primer: Intelligent Design Theory in a Nutshell" (PDF). Intelligent Design and Evolution
Awareness Center. 2004.
http://www.ideacenter.org/stuff/contentmgr/files/393410a2d36e9b96329c2faff7e2a4df/miscdocs/intell
igentdesigntheoryinanutshell.pdf. Retrieved 2007-05-13.
• "Intelligent Design". Intelligent Design network. 2007.
http://www.intelligentdesignnetwork.org/. Retrieved 2007-05-13.
3. ^ The Creationists, Expanded Edition. Harvard University Press; 2006. ISBN 0674023390. p.
373, 379–380.
4. ^ a b Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., Context pg.
32 ff, citing Edwards v. Aguillard, 482 U.S. 578 .
5. ^ a b "ID is not a new scientific argument, but is rather an old religious argument for the
existence of God. He traced this argument back to at least Thomas Aquinas in the 13th century, who
framed the argument as a syllogism: Wherever complex design exists, there must have been a
designer; nature is complex; therefore nature must have had an intelligent designer." "This argument
for the existence of God was advanced early in the 19th century by Reverend Paley" (the teleological
argument) "The only apparent difference between the argument made by Paley and the argument for
ID, as expressed by defense expert witnesses Behe and Minnich, is that ID's 'official position' does
not acknowledge that the designer is God." Kitzmiller v. Dover Area School District, 04 cv 2688
(December 20, 2005)., Ruling, p. 24.
6. ^ a b c d Forrest, Barbara (May 2007). "Understanding the Intelligent Design Creationist
Movement: Its True Nature and Goals." (PDF). Washington, D.C.: Center for Inquiry, Office of Public
Policy. http://www.centerforinquiry.net/uploads/attachments/intelligent-design.pdf. Retrieved 2007-
08-06.
7. ^ a b "Q. Has the Discovery Institute been a leader in the intelligent design movement? A.
Yes, the Discovery Institute's Center for Science and Culture. Q. And are almost all of the individuals
who are involved with the intelligent design movement associated with the Discovery Institute? A. All
of the leaders are, yes." Barbara Forrest, 2005, testifying in the Kitzmiller v. Dover Area School
District trial. "Kitzmiller v. Dover Area School District Trial transcript: Day 6 (October 5), PM Session,
Part 1.". TalkOrigins Archive. 2005. http://www.talkorigins.org/faqs/dover/day6pm.html. Retrieved
2007-07-19.
• "The Discovery Institute is the ideological and strategic backbone behind the eruption
of skirmishes over science in school districts and state capitals across the country". In:
Wilgoren, J (August 21, 2005). "Politicized Scholars Put Evolution on the Defensive" (PDF).
New York Times. http://www.msu.edu/course/te/407/FS05Sec3/te408/files/Politicized
%20Scholars%20Put%20Evolution%20on%20the%20Defensive%20-%20New%20York
%20Times.pdf. Retrieved 2007-07-19.
• "Who is behind the ID movement?". Frequently Asked Questions About "Intelligent
Design". American Civil Liberties Union. September 16, 2005.
http://www.aclu.org/religion/schools/16371res20050916.html. Retrieved 2007-07-20.
 • Kahn, JP (July 27, 2005). "The Evolution of George Gilder. The Author And Tech-
Sector Guru Has A New Cause To Create Controversy With: Intelligent Design". The Boston
Globe. http://www.discovery.org/scripts/viewDB/index.php?command=view&program=DI
%20Main%20Page%20-%20News&id=2745. Retrieved 2007-07-19.
• "Who's Who of Intelligent Design Proponents" (PDF). Science & Religion Guide.
Science & Theology News. November 2005.
http://www.discovery.org/scripts/viewDB/filesDB-download.php?
command=download&id=602. Retrieved 2007-07-20.
• "The engine behind the ID movement is the Discovery Institute". Attie, Alan D.; Elliot
Sober, Ronald L. Numbers, Richard M. Amasino, Beth Cox4, Terese Berceau, Thomas
Powell and Michael M. Cox (2006). "Defending science education against intelligent design:
a call to action". Journal of Clinical Investigation 116:1134–1138 . A publication of the
American Society for Clinical Investigation.. doi:10.1172/JCI28449.
http://www.jci.org/articles/view/28449. Retrieved 2007-07-20.
8. ^ "Science and Policy: Intelligent Design and Peer Review". American Association for the
Advancement of Science. 2007.
http://www.aaas.org/spp/dser/03_Areas/evolution/issues/peerreview.shtml. Retrieved 2007-07-19.
9. ^ a b c "the writings of leading ID proponents reveal that the designer postulated by their
argument is the God of Christianity". Kitzmiller v. Dover Area School District, 04 cv 2688 (December
20, 2005)., Ruling p. 26. A selection of writings and quotes of intelligent design supporters
demonstrating this identification of the Christian God with the intelligent designer are found in the pdf
Horse's
Mouthhttp://web.archive.org/web/20080627021627/http://home.kc.rr.com/bnpndxtr/download/Horses
Mouth-BP007.pdf[dead link] (PDF) by Brian Poindexter, dated 2003.
 10.^ a b c William A. Dembski, when asked in an interview whether his research concluded that
God is the Intelligent Designer, stated "I believe God created the world for a purpose. The Designer
of intelligent design is, ultimately, the Christian God". Devon Williams (December 14, 2007).
"CitizenLink: Friday Five: William A. Dembski". Focus on the Family.
http://www.citizenlink.org/content/A000006139.cfm. Retrieved 2007-12-15.
11.^ Stephen C. Meyer and Paul A. Nelson (May 1, 1996). "CSC – Getting Rid of the Unfair
Rules, A book review, Origins & Design"]. http://www.discovery.org/scripts/viewDB/index.php?
command=view&id=1685. Retrieved 2007-05-20.
• Phillip E. Johnson (August 31, 1996). "Starting a Conversation about Evolution".
Phillip Johnson Files. Access Research Network.
http://www.arn.org/docs/johnson/ratzsch.htm. Retrieved 2007-05-20.
• Stephen C. Meyer (December 1, 2002). "The Scientific Status of Intelligent Design:
The Methodological Equivalence of Naturalistic and Non-Naturalistic Origins Theories".
Ignatius Press. http://www.discovery.org/scripts/viewDB/index.php?
command=view&id=1780.
• Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., Whether
ID Is Science
• Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., Lead
defense expert Professor Behe admitted that his broadened definition of science, which
encompasses ID, would also include astrology.
• See also "Evolution of Kansas science standards continues as Darwin's theories
regain prominence". International Herald Tribune. February 13, 2007.
http://www.iht.com/articles/ap/2007/02/13/america/NA-GEN-US-Kansas-Evolution-
History.php. Retrieved 2007-05-20.
 12.^ See: 1) List of scientific societies explicitly rejecting intelligent design 2) Kitzmiller v. Dover
page 83. 3) The Discovery Institute's A Scientific Dissent From Darwinism petition begun in 2001 has
been signed by "over 700 scientists" as of August 20, 2006. A four day A Scientific Support for
Darwinism petition gained 7733 signatories from scientists opposing ID. The AAAS, the largest
association of scientists in the U.S., has 120,000 members, and firmly rejects ID. More than 70,000
Australian scientists and educators condemn teaching of intelligent design in school science classes
List of statements from scientific professional organizations on the status intelligent design and other
forms of creationism. According to The New York Times "There is no credible scientific challenge to
the theory of evolution as an explanation for the complexity and diversity of life on earth". Dean,
Cordelia (September 27, 2007). "Scientists Feel Miscast in Film on Life's Origin". New York Times.
http://www.nytimes.com/2007/09/27/science/27expelled.html?_r=2&oref=slogin&oref=slogin.
Retrieved 2007-09-28.
13.^ a b "Teachernet, Document bank". Creationism teaching guidance. UK Department for
Children, Schools and Families. September 18, 2007.
http://www.teachernet.gov.uk/docbank/index.cfm?id=11890. Retrieved 2007-10-01. "The intelligent
design movement claims there are aspects of the natural world that are so intricate and fit for
purpose that they cannot have evolved but must have been created by an 'intelligent designer'.
Furthermore they assert that this claim is scientifically testable and should therefore be taught in
science lessons. Intelligent design lies wholly outside of science. Sometimes examples are quoted
that are said to require an 'intelligent designer'. However, many of these have subsequently been
shown to have a scientific explanation, for example, the immune system and blood clotting
mechanisms.
Attempts to establish an idea of the 'specified complexity' needed for intelligent design are
surrounded by complex mathematics. Despite this, the idea seems to be essentially a modern
version of the old idea of the "God-of-the-gaps". Lack of a satisfactory scientific explanation of some
phenomena (a 'gap' in scientific knowledge) is claimed to be evidence of an intelligent designer."
14.^ Nature Methods Editorial. An intelligently designed response. Nat. Methods.
2007;4(12):983. doi:10.1038/nmeth1207-983.
15.^ Mark Greener. Taking on creationism. Which arguments and evidence counter
pseudoscience?. EMBO Reports. 2007;8(12):1107–1109. doi:10.1038/sj.embor.7401131. PMID
18059309.
16.^ "Science and Creationism: A View from the National Academy of Sciences". National
Academy of Sciences. 1999. http://www.nap.edu/openbook.php?isbn=0309064066&page=25.
17.^ National Science Teachers Association, a professional association of 55,000 science
teachers and administrators National Science Teachers Association (August 3, 2005). "National
Science Teachers Association Disappointed About Intelligent Design Comments Made by President
Bush". Press release. http://www.nsta.org/about/pressroom.aspx?id=50794. "We stand with the
nation's leading scientific organizations and scientists, including Dr. John Marburger, the president's
top science advisor, in stating that intelligent design is not science. ...It is simply not fair to present
pseudoscience to students in the science classroom."
18.^ a b David Mu. Trojan Horse or Legitimate Science: Deconstructing the Debate over
Intelligent Design [PDF]. Harvard Science Review. Fall 2005;19(1). "For most members of the
mainstream scientific community, ID is not a scientific theory, but a creationist pseudoscience"."
19.^ a b "Professional Ethics Report" (PDF). American Association for the Advancement of
Science. 2001. http://www.aaas.org/spp/sfrl/per/per26.pdf. "Creationists are repackaging their
message as the pseudoscience of intelligent design theory."
20.^ Evolution critics seek role for unseen hand in education. Nature. 2002;416(6878):250.
doi:10.1038/416250a. PMID 11907537. "But many scientists regard ‘intelligent design’ as
pseudoscience, and say that it is being used as a Trojan Horse to introduce the teaching of
creationism into schools"
21.^ Attie, A. D.. Defending science education against intelligent design: a call to action. Journal
of Clinical Investigation. 2006;116(5):1134–1138. doi:10.1172/JCI28449. PMID 16670753. PMC
1451210.
• H. Allen Orr (May 2005). "Devolution—Why intelligent design isn't". Annals of
Science. New Yorker. http://www.newyorker.com/archive/2005/05/30/050530fa_fact.
"Biologists aren't alarmed by intelligent design's arrival in Dover and elsewhere because they
have all sworn allegiance to atheistic materialism; they're alarmed because intelligent design
is junk science."
• Robert T. Pennock Tower of Babel: The Evidence Against the New Creationism.
• Mark Bergin (February 25, 2006). "Junk science". World Magazine.
http://www.worldmag.com/articles/11553.
22.^ Junk Science. Macmillan; 2006. ISBN 9780312352417. p. 210 ff.
23.^ a b Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., pp. 31 – 33.
24.^ "Media Backgrounder: Intelligent Design Article Sparks Controversy". Discovery Institute.
September 7, 2004. http://www.discovery.org/scripts/viewDB/index.php?command=view&id=2190.
• James M. Kushiner (June 2002). "Berkeley's Radical". Touchstone Magazine.
http://touchstonemag.com/archives/article.php?id=15-05-037-i.
• Jodi Wilgoren (August 21, 2005). "Politicized Scholars Put Evolution on the
Defensive" (PDF). The New York Times.
http://www.msu.edu/course/te/407/FS05Sec3/te408/files/Politicized%20Scholars%20Put
%20Evolution%20on%20the%20Defensive%20-%20New%20York%20Times.pdf.
 • Downey, Roger (February 1, 2006). "Discovery's Creation". Seattle Weekly.
http://seattleweekly.com/2006-02-01/news/discovery-s-creation.php. Retrieved 2007-07-27.
25.^ Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., Conclusion of
Ruling.
26.^ "Plato's Timaeus". The Stanford Encyclopedia of Philosophy. The Metaphysics Research
Lab, Stanford University. October 25, 2005. http://plato.stanford.edu/entries/plato-timaeus/. Retrieved
2007-07-22.
27.^ Plato. "Timaeus". Internet Classics Archive. classics.mit.edu.
http://classics.mit.edu/Plato/timaeus.html. Retrieved 2007-07-22.
28.^ Aristotle, Metaphysics Bk. 12
29.^ Linda Trinkaus Zagzebski (2006). The Philosophy of Religion: An Historical Introduction ,
31; Cicero, De Natura Deorum, Book I, 36–37, Latin Library.
30.^ Thomas Aquinas, Summa Theologiae "Thomas Aquinas' 'Five Ways' (archive link)" in
faithnet.org.uk.
31.^ William Paley, Natural Theology: or, Evidences of the Existence and Attributes of the Deity ,
1809, London, Twelfth Edition.
32.^ "Letter 2998 — Darwin, C. R. to Gray, Asa, 26 Nov (1860)". Darwin Correspondence
Project. http://www.darwinproject.ac.uk/entry-2998. Retrieved 2010-08-11.
33.^ Gerard Radnitzky. Evolutionary epistemology, rationality, and the sociology of knowledge .
Open Court Publishing; 1993. ISBN 0812690397. p. 140.
34.^ See, e.g., the publisher's editorial description of the 2006 paperback printing of William
Paley (1803) Natural Theology" : "William Paley's classic brings depth to the history of intelligent
design arguments. The contrivance of the eye, the ear, and numerous other anatomical features
throughout the natural world are presented as arguments for God's presence and concern. While
there are distinctive differences between Paley's argument and those used today by intelligent
design theorists and creationists, it remains a fascinating glimpse of the nineteenth-century's debate
over the roles of religion and science".
• David C. Steinmetz (2005) "The Debate on Intelligent Design" in The Christian
Century, (December 27, 2005, pp. 27–31.)
• Leading intelligent design proponent William Dembski (2001) argues the opposing
view in Is Intelligent Design a form of natural theology?
35.^ Forrest, Barbara. Know Your Creationists: Know Your Allies
36.^ a b Stephen C. Meyer (March 1986). "We Are Not Alone". Eternity. Access Research
Network. http://www.arn.org/docs/meyer/sm_notalone.htm. Retrieved 2007-10-10.
37.^ Charles B. Thaxton, Ph.D. (November 13–16, 1986). "DNA, Design and the Origin of Life".
Christian Leadership Ministries. http://www.origins.org/articles/thaxton_dnadesign.html. Retrieved
2007-10-10.
38.^ a b Charles B. Thaxton (June 23–26, 1988, revised July 1988 and May 1991). "In Pursuit of
Intelligent Causes: Some Historical Background".
http://www.leaderu.com/offices/thaxton/docs/inpursuit.html. Retrieved 2007-10-06.
39.^ a b c Dembski: "Intelligent design is just the Logos theology of John's Gospel restated in the
idiom of information theory," Touchstone Magazine. Volume 12, Issue4: July/August, 1999
40.^ a b Phillip Johnson: "Our strategy has been to change the subject a bit so that we can get
the issue of Intelligent Design, which really means the reality of God, before the academic world and
into the schools." Johnson 2004. Christianity.ca. Let's Be Intelligent About Darwin[dead
link]http://web.archive.org/web/20070608233455/http://www.christianity.ca/news/social-
issues/2004/03.001.html. "This isn't really, and never has been a debate about science. It's about
religion and philosophy." Johnson 1996. World Magazine. Witnesses For The Prosecution. "So the
question is: "How to win?" That's when I began to develop what you now see full-fledged in the
"wedge" strategy: "Stick with the most important thing"—the mechanism and the building up of
information. Get the Bible and the Book of Genesis out of the debate because you do not want to
raise the so-called Bible-science dichotomy. Phrase the argument in such a way that you can get it
heard in the secular academy and in a way that tends to unify the religious dissenters. That means
concentrating on, "Do you need a Creator to do the creating, or can nature do it on its own?" and
refusing to get sidetracked onto other issues, which people are always trying to do." Johnson 2000.
Touchstone magazine. Berkeley's Radical An Interview with Phillip E. Johnson at the Wayback
Machine (archived June 9, 2007).
41.^ Stephen C. Meyer: "I think the designer is God ..." (Darwin, the marketing of Intelligent
Design . Nightline ABC News, with Ted Koppel, August 10, 2005); Nancy Pearcey: "By contrast,
design theory demonstrates that Christians can sit in the supernaturalist’s “chair” even in their
professional lives, seeing the cosmos through the lens of a comprehensive biblical worldview.
Intelligent Design steps boldly into the scientific arena to build a case based on empirical data. It
takes Christianity out of the ineffectual realm of value and stakes out a cognitive claim in the realm of
objective truth. It restores Christianity to its status as genuine knowledge, equipping us to defend it in
the public arena". (Total Truth, Crossway Books, June 29, 2004, ISBN 1581344589, pp. 204-205)
42.^ a b Leon Lynn (Winter 1997/98). "Rethinking Schools Online". Creationists Push Pseudo-
Science Text. http://www.rethinkingschools.org/archive/12_02/panda.shtml. Retrieved 2009-02-08.
43.^ a b Nick Matzke (2006). "NCSE Resource -- 9.0. Matzke (2006): The Story of the Pandas
Drafts". National Center for Science Education. http://ncse.com/rncse/26/1-2/design-trial. Retrieved
2009-11-18. *Nick Matzke (2006). "Missing Link discovered!". National Center for Science
Education. Archived from the original on 2007-01-14.
http://web.archive.org/web/20070114121029/http://www2.ncseweb.org/wp/?p=80. Retrieved 2009-
11-18.
44.^ Nick Matzke (August 14, 2007). "The true origin of "intelligent design"". The Panda's
Thumb. http://pandasthumb.org/archives/2007/08/the-true-origin.html. Retrieved 2010-01-21.
"Journals: Scientific American (1846 - 1869)". http://digital.library.cornell.edu/cgi/t/text/pageviewer-
idx?c=scia;cc=scia;rgn=full%20text;idno=scia0002-48;didno=scia0002-
48;view=image;seq=00383;node=scia0002-48%3A1. Retrieved 2010-01-21.
45.^ Dove, Patrick Edward, The theory of human progression, and natural probability of a reign
of justice. London, Johnstone & Hunter, 1850. LC 08031381 "Intelligence-Intelligent Design".
46.^ Charles Darwin (May 23, 1861). "Letter 3154—Darwin, C. R. to Herschel, J. F. W., 23 May
1861". Darwin Correspondence Project. http://www.darwinproject.ac.uk/darwinletters/calendar/entry-
3154.html.
47.^ "The British Association". The Times. September 20, 1873. pp. 10; col A..
48.^ William P. Alston. In: Paul Edwards. Encyclopedia of Philosophy. New York City, London:
Macmillan Publishing Company, The Free Press, Collier Macmillan Publishers; 1967. ISBN
0028949900.
49.^ Robert Nozick. Anarchy, State, and Utopia. USA: Basic Books; 1974. ISBN 0465097200. p.
19.
50.^ James E. Horigan. Chance or Design?. Philosophical Library; 1979.
51.^ Nicholas Timmins (January 13, 1982). "Evolution according to Hoyle: Survivors of disaster
in an earlier world". The Times (61130): p. 22. http://telicthoughts.com/sir-fred-hoyle-and-the-origins-
of-id/. "F. Hoyle stated in a 1982 speech: '...one arrives at the conclusion that biomaterials with their
amazing measure or order must be the outcome of intelligent design.'"
 52.^ Jonathan Witt (December 20, 2005). "Evolution News & Views: Dover Judge Regurgitates
Mythological History of Intelligent Design". Discovery Institute.
http://www.evolutionnews.org/2005/12/post_6.html. Retrieved 2007-10-05.
53.^ DarkSyde (March 11, 2006). "Daily Kos: Know Your Creationists: Know Your Allies".
interview with Barbara Forrest. http://www.dailykos.com/story/2006/3/11/8448/52824. Retrieved
2007-10-05.
54.^ William Safire (August 21, 2005). "On Language: Neo-Creo". New York Times.
http://www.nytimes.com/2005/08/21/magazine/21ONLANGUAGE.html?
position=&ei=5090&en=f2de0d764cc7e0e8&ex=1282276800&adxnnl=1&partner=rssuserland&emc=
rss&pagewanted=print&adxnnlx=1132902202-gyP0H4EZfG7IeNHPMWlcBw.
55.^ a b c Nick Matzke (2004). "NCSE Resource". Introduction: Of Pandas and People, the
foundational work of the 'Intelligent Design' movement . National Center for Science Education.
http://ncse.com/creationism/analysis/critique-pandas-people. Retrieved 2007-09-24.
56.^ Richard P. Aulie (1998). "A Reader's Guide to Of Pandas and People". National
Association of Biology Teachers. http://www.stephenjaygould.org/ctrl/archive/design/aulie_of-
pandas.html. Retrieved 2007-10-05.
57.^ Eugenie C. Scott. Biological design in science classrooms. Proceedings of the National
Academy of Sciences. May 15, 2007 [cited 2009-06-02];104(Suppl 1):8669–8676.
doi:10.1073/pnas.0701505104. PMID 17494747. PMC 1876445.
58.^ Nick Matzke; Jon Buell (October 13, 2005). "I guess ID really was "Creationism's Trojan
Horse" after all". The Panda's Thumb.
http://www.pandasthumb.org/archives/2005/10/i_guess_id_real.html. Retrieved 2009-06-02.
59.^ William A. Dembski (March 29, 2005) (PDF). Expert Witness Report: The Scientific Status
of Intelligent Design. Kitzmiller v. Dover Area School District trial document.
http://www.designinference.com/documents/2005.09.Expert_Report_Dembski.pdf. Retrieved 2009-
06-02.
60.^ Behe, Michael (1997): Molecular Machines: Experimental Support for the Design Inference
[1]
61.^ Irreducible complexity of these examples is disputed; see Kitzmiller, pp. 76–78, and Ken
Miller Webcast
62.^ The Collapse of "Irreducible Complexity" Kenneth R. Miller Brown University [2]
63.^ John H. McDonald's "reducibly complex mousetrap"
64.^ David Ussery, "A Biochemist's Response to 'The Biochemical Challenge to Evolution'"
65.^ For example, Bridgham et al. showed that gradual evolutionary mechanisms can produce
complex protein-protein interaction systems from simpler precursors. Bridgham et al.. Evolution of
Hormone-Receptor Complexity by Molecular Exploitation. Science. 2006;312(5770):97–101.
doi:10.1126/science.1123348. PMID 16601189.
66.^ Orr, H. Allen (May 30, 2005). "Devolution". The New Yorker.
http://www.newyorker.com/archive/2005/05/30/050530fa_fact. This article draws from the following
exchange of letters in which Behe admits to sloppy prose and non-logical proof: Behe, M.; Dembski,
Wells, Nelson, Berlinski (March 26, 2003). "Has Darwin met his match? Letters—An exchange over
ID". Discovery Institute. http://www.discovery.org/scripts/viewDB/index.php?program=CRSC
%20Responses&command=view&id=1406. Retrieved 2006-11-30.
67.^ Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005)., p. 64.
68.^ Dembski. Intelligent Design, p. 47
69.^ William Dembski, Photo by Wesley R. Elsberry, taken at lecture given at University of
California at Berkeley, 2006/03/17.
 70.^ Branden Fitelson, Christopher Stephens, Elliott Sober. How Not to Detect Design: A review
of William A. Dembski's The Design Inference—Eliminating Chance Through Small Probabilities
[PDF]. Cambridge University Press; 1998.
71.^ Some of Dembski's responses to assertions of specified complexity being a tautology can
be found at William A. Dembski. "Another way to detect design". ARN.
http://www.arn.org/docs/dembski/wd_anotherwaytodetectdesign.htm.
72.^ a b Richard Wein (2002). "Not a Free Lunch But a Box of Chocolates: A critique of William
Dembski's book No Free Lunch". http://www.talkorigins.org/design/faqs/nfl/.
73.^ Rich Baldwin (2005). "Information Theory and Creationism".
http://www.talkorigins.org/faqs/information/dembski.html.
74.^ Mark Perakh (2005). "Dembski 'displaces Darwinism' mathematically -- or does he?".
http://www.talkreason.org/articles/newmath.cfm.
75.^ Jason Rosenhouse. How Anti-Evolutionists Abuse Mathematics [PDF]. The Mathematical
Intelligencer. Fall 2001;23(4):3–8.
76.^ John S. Wilkins, Wesley R. Elsberry. The Advantages of Theft over Toil: The Design
Inference and Arguing from Ignorance. Biology and Philosophy. 2001;16:711–724.
77.^ Richard Dawkins. The God Delusion. Boston: Houghton Mifflin; 2006. ISBN 0618680004.
78.^ Evolutionary algorithms now surpass human designers. New Scientist. July 28, 2007.
79.^ Guillermo Gonzalez. The Privileged Planet: How Our Place in the Cosmos is Designed for
Discovery. Washington, DC: Regnery Publ.; 2004. ISBN 0-89526-065-4.
80.^ The Panda's Thumb. review of The Privileged Planet
81.^ Is The Universe Fine-Tuned For Us? Victor J. Stenger. University of Colorado. (PDF file)
82.^ Victor J. Stenger. "The Anthropic Principle" (PDF). University of Colorado.
http://www.colorado.edu/philosophy/vstenger/Cosmo/ant_encyc.pdf.
 83.^ Joseph Silk. Our place in the Multiverse. Nature. September 14, 2006;443(7108).
84.^ See, e.g., Gerald Feinberg and Robert Shapiro. A Puddlian Fable. In: Huchingson. Religion
and the Natural Sciences. 1993. p. 220–221.
85.^ Sewell, Granville (December 28, 2005). "Evolution's Thermodynamic Failure". The
American Spectator. http://spectator.org/archives/2005/12/28/evolutions-thermodynamic-failu.
Retrieved 2007-02-16. (Also available from the Discovery Institute)
86.^ "Entropy, Disorder and Life". TalkOrigins Archive.
http://www.talkorigins.org/faqs/thermo/entropy.html. Retrieved 2007-07-17.
87.^ Dembski. "Questions About Intelligent Design". What is Intelligent Design?. Discovery
Institute. http://www.discovery.org/csc/topQuestions.php#questionsAboutIntelligentDesign. "The
theory of Intelligent Design holds that certain features of the universe and of living things are best
explained by an intelligent cause, not an undirected process such as natural selection. "
88.^ "The Act of Creation: Bridging Transcendence and Immanence". LeaderU.
http://www.leaderu.com/offices/dembski/docs/bd-the_ac.html.
89.^ a b The Case Against Intelligent Design [PDF]. The New Republic. August 22–29,
2005;233(8/9):21–
33.http://web.archive.org/web/20060618235930/http://pondside.uchicago.edu/cluster/pdf/coyne/New
_Republic_ID.pdf
90.^ Donald E. Simanek. "Intelligent Design: The Glass is Empty".
http://www.lhup.edu/~dsimanek/philosop/empty.htm.
91.^ IDEA "One need not fully understand the origin or identity of the designer to determine that
an object was designed. Thus, this question is essentially irrelevant to intelligent design theory,
which merely seeks to detect if an object was designed ... Intelligent design theory cannot address
the identity or origin of the designer—it is a philosophical / religious question that lies outside the
domain of scientific inquiry. Christianity postulates the religious answer to this question that the
designer is God who by definition is eternally existent and has no origin. There is no logical
philosophical impossibility with this being the case (akin to Aristotle's 'unmoved mover') as a religious
answer to the origin of the designer..." FAQ: Who designed the designer? FAQ: Who designed the
designer?
92.^ Jason Rosenhouse. "Who Designed the Designer?". Creation & Intelligent Design Watch.
Committee for the Scientific Investigation of Claims of the Paranormal.
http://www.csicop.org/intelligentdesignwatch/designer.html.
93.^ Richard Dawkins. The Blind Watchmaker: Why the Evidence of Evolution Reveals a
Universe without Design. p. 141.
94.^ See, e.g., Joseph Manson (September 27, 2005). "Intelligent design is pseudoscience".
UCLA Today. http://www.today.ucla.edu/portal/ut/050927voices_pseudoscience.aspx. ; Rev Max
(July–August 2006). "The Incredibly Strange Story of Intelligent Design". New Dawn Magazine (97).
95.^ "NCSE Resource". Evolving Banners at the Discovery Institute. August 29, 2002.
http://ncse.com/creationism/general/evolving-banners-at-discovery-institute. Retrieved 2007-10-07.
96.^ Conclusion of Kitzmiller v. Dover Area School District Ruling
97.^ Wise, D.U., 2001, Creationism's Propaganda Assault on Deep Time and Evolution, Journal
of Geoscience Education, v. 49, n. 1, p. 30–35.
98.^ Marcus R. Ross. Who Believes What? Clearing up Confusion over Intelligent Design and
Young-Earth Creationism [PDF]. Journal of Geoscience Education. May, 2005;53(3):319–323.
99.^ Ronald L. Numbers. The Creationists: From Scientific Creationism to Intelligent Design,
Expanded Edition. Cambridge, Massachusetts: Harvard University Press; November 30, 2006. ISBN
0674023390.
 100.^ Forrest, B.C. and Gross, P.R.. Evolution and the Wedge of Intelligent Design: The Trojan
Horse Strategy. Oxford: Oxford University Press; 2003. ISBN 0195157427.
101.^ Robert T. Pennock. Wizards of ID: Reply to Dembski. In: Robert T. Pennock. Intelligent
Design Creationism and Its Critics: Philosophical, Theological, and Scientific Perspectives .
Cambridge, Massachusetts: MIT Press; 2001. ISBN 0262661241. "Dembski chides me for never
using the term "intelligent design" without conjoining it to "creationism". He implies (though never
explicitly asserts) that he and others in his movement are not creationists and that it is incorrect to
discuss them in such terms, suggesting that doing so is merely a rhetorical ploy to "rally the troops".
(2) Am I (and the many others who see Dembski's movement in the same way) misrepresenting their
position? The basic notion of creationism is the rejection of biological evolution in favor of special
creation, where the latter is understood to be supernatural. Beyond this there is considerable
variability..." p. 645–667.; Pennock, Robert T.. Tower of Babel: Evidence Against the New
Creationism. Cambridge, Massachusetts: MIT Press; 1999.
102.^ Eugenie Scott. The Creation/Evolution Continuum. NCSE Reports. 1999;19(4):16–17, 23–
25.; Scott, Eugenie C.. Evolution vs. Creationism: An Introduction. Westport, Connecticut:
Greenwood Press; 2004. ISBN 0520246500.
103.^ a b c "The social consequences of materialism have been devastating. As symptoms,
those consequences are certainly worth treating. However, we are convinced that in order to defeat
materialism, we must cut it off at its source. That source is scientific materialism. This is precisely our
strategy. If we view the predominant materialistic science as a giant tree, our strategy is intended to
function as a 'wedge' that, while relatively small, can split the trunk when applied at its weakest
points. The very beginning of this strategy, the 'thin edge of the wedge,' was Phillip Johnson's
critique of Darwinism begun in 1991 in Darwinism on Trial, and continued in Reason in the Balance
and Defeating Darwinism by Opening Minds. Michael Behe's highly successful Darwin's Black Box
followed Johnson's work. We are building on this momentum, broadening the wedge with a positive
scientific alternative to materialistic scientific theories, which has come to be called the theory of
intelligent design (ID). Design theory promises to reverse the stifling dominance of the materialist
worldview, and to replace it with a science consonant with Christian and theistic convictions". Wedge
Document Discovery Institute, 1999. (PDF file)
104.^ a b c "I have built an intellectual movement in the universities and churches that we call
The Wedge, which is devoted to scholarship and writing that furthers this program of questioning the
materialistic basis of science. [...] Now the way that I see the logic of our movement going is like this.
The first thing you understand is that the Darwinian theory isn't true. It's falsified by all of the
evidence and the logic is terrible. When you realize that, the next question that occurs to you is, well,
where might you get the truth? [...] I start with John 1:1. In the beginning was the word. In the
beginning was intelligence, purpose, and wisdom. The Bible had that right. And the materialist
scientists are deluding themselves." Johnson 1999. Reclaiming America for Christ Conference. How
the Evolution Debate Can Be Won
105.^ "Discovery Institute fellows and staff". http://www.discovery.org/fellows/. "Center for
Science and Culture fellows and staff". http://www.discovery.org/csc/fellows.php.
106.^ Barbara Forrest (2001). "The Wedge at Work: Intelligent Design Creationism and Its
Critics". http://www.infidels.org/library/modern/barbara_forrest/wedge.html.
107.^ a b "...intelligent design does not address metaphysical and religious questions such as
the nature or identity of the designer," and "...the nature, moral character and purposes of this
intelligence lie beyond the competence of science and must be left to religion and philosophy". In:
"Truth Sheet # 09-05 Does intelligent design postulate a "supernatural creator? ". Discovery Institute.
http://www.discovery.org/scripts/viewDB/filesDB-download.php?command=download&id=565.
Retrieved 2007-07-19.
 108.^ Phillip Johnson (April 1999). "'Keeping the Darwinists Honest' an interview with Phillip
Johnson". Citizen Magazine. "Intelligent Design is an intellectual movement, and the Wedge
strategy stops working when we are seen as just another way of packaging the Christian evangelical
message. [...] The evangelists do what they do very well, and I hope our work opens up for them
some doors that have been closed".
109.^ a b Phillip Johnson. Touchstone: A Journal of Mere Christianity. 1999. "...the first thing that
has to be done is to get the Bible out of the discussion. ...This is not to say that the biblical issues are
unimportant; the point is rather that the time to address them will be after we have separated
materialist prejudice from scientific fact". The Wedge
110.^ a b William Dembski, 1998. The Design Inference.
111.^ Dembski, 1999. Intelligent Design: The Bridge Between Science and Theology, p. 210.
112.^ William Dembski (2005). "Intelligent Design's Contribution to the Debate Over Evolution: A
Reply to Henry Morris".
http://www.designinference.com/documents/2005.02.Reply_to_Henry_Morris.htm.
113.^ Barbara Forrest. Expert Testimony. Kitzmiller v. Dover Area School District trial transcript,
Day 6 (October 5) "What I am talking about is the essence of intelligent design, and the essence of it
is theistic realism as defined by Professor Johnson. Now that stands on its own quite apart from what
their motives are. I'm also talking about the definition of intelligent design by Dr. Dembski as the
Logos theology of John's Gospel. That stands on its own. [...] Intelligent design, as it is understood
by the proponents that we are discussing today, does involve a supernatural creator, and that is my
objection. And I am objecting to it as they have defined it, as Professor Johnson has defined
intelligent design, and as Dr. Dembski has defined intelligent design. And both of those are basically
religious. They involve the supernatural".
 114.^ "Understanding the Intelligent Design Creationist Movement: Its True Nature and Goals. A
Position Paper" (PDF). Center for Inquiry, Office of Public Policy. May 2007.
http://www.centerforinquiry.net/uploads/attachments/intelligent-design.pdf.
115.^ H. Ross (2002). "More than intelligent design". Facts for Faith. Pasadena, California:
Reasons to Believe. http://www.reasons.org/resources/fff/2002issue10/index.shtml#more_than_id.
Retrieved April 25, 2007.
116.^ Henry M. Morris (1999). "Design is not enough!". Back to Genesis. Santee, California:
Institute for Creation Research. http://www.icr.org/article/859/17/. Retrieved April 25, 2007.
117.^ Carl Wieland (2002). "AiG's views on the Intelligent Design movement". Answers in
Genesis. http://www.answersingenesis.org/docs2002/0830_IDM.asp. Retrieved April 25, 2007.
118.^ "Nearly Two-thirds of U.S. Adults Believe Human Beings Were Created by God". The
Harris Poll #52. Harris Interactive. July 6, 2005.
http://www.harrisinteractive.com/harris_poll/index.asp?PID=581. Retrieved 2007-07-13.
119.^ "Sandia National Laboratories says that the Intelligent Design Network (IDNet-NM/Zogby)
"Lab Poll" is BOGUS!". New Mexicans for Science and Reason. http://www.nmsr.org/id-poll.htm.
Retrieved 2007-07-13.
120.^ Mooney, Chris (September 11, 2003). "Polling for ID". Doubt and About. Committee for
Skeptical Inquiry. http://www.csicop.org/doubtandabout/polling/. Retrieved 2007-02-
16. http://web.archive.org/web/20080327041611/http://csicop.org/doubtandabout/polling/
121.^ David Harris. "'Intelligent Design'-ers launch new assault on curriculum using lies and
deception". Salon.com. Archived from the original on 2003-08-16.
http://web.archive.org/web/20030816135718/http://blogs.salon.com/0001092/2003/07/30.html.
Retrieved 2007-07-13.
 122.^ According to the poll, 18% of the physicians believed that God created humans exactly as
they appear today. Another 42% believed that God initiated and guided an evolutionary process that
has led to current human beings. The poll also found that "an overwhelming majority of Jewish
doctors (83%) and half of Catholic doctors (51%) believe that intelligent design is simply "a religiously
inspired pseudo-science rather than a legitimate scientific speculation". The poll also found that
"more than half of Protestant doctors (63%) believe that intelligent design is a "legitimate scientific
speculation".
"Majority of Physicians Give the Nod to Evolution Over Intelligent Design". Press release.
http://www.hcdi.net/News/PressRelease.cfm?ID=93. Retrieved 2007-10-08.
123.^ Gallup, "Evolution, creationism, intelligent design,". Retrieved 24 August 2010.
124.^ "Expelled, No Intelligence Allowed (2008)". http://www.imdb.com/title/tt1091617/.
125.^ "New Anti-Evolution Film Stirs Controversy".
http://www.livescience.com/strangenews/080404-expelled-movie.html.
126.^ Shaw, Linda (March 31, 2005). "Does Seattle group "teach controversy" or contribute to
it?". Seattle Times. http://seattletimes.nwsource.com/html/localnews/2002225932_design31m.html.
127.^ "NABT's Statement on Teaching Evolution". National Association of Biology Teachers.
Archived from the original on 2006-09-27.
http://web.archive.org/web/20060927160040/http://www.nabt.org/sub/position_statements/evolution.
asp.
128.^ "IAP Statement on the Teaching of Evolution" (PDF). The Interacademy Panel on
International Issues. June 21, 2006.
http://www.interacademies.net/Object.File/Master/6/150/Evolution%20statement.pdf. Retrieved
2008-10-17. Joint statement issued by the national science academies of 67 countries, including the
United Kingdom's Royal Society.
 129.^ From the world's largest general scientific society:
• American Association for the Advancement of Science (February 16, 2006).
"Statement on the Teaching of Evolution" (PDF). Press release.
http://www.aaas.org/news/releases/2006/pdf/0219boardstatement.pdf. Retrieved 2008-10-
17.
• American Association for the Advancement of Science (February 19, 2006). "AAAS
Denounces Anti-Evolution Laws". Press release.
http://www.aaas.org/news/releases/2006/0219boardstatement.shtml. Retrieved 2008-10-17.
130.^ Coultan, Mark (November 27, 2005). "Intelligent design a Trojan horse, says creationist".
Sydney Morning Herald. http://www.smh.com.au/news/world/intelligent-design-a-trojan-horse-says-
creationist/2005/11/26/1132966007431.html. Retrieved 2007-07-29.
131.^ "Intelligent Design: Creationism's Trojan Horse". Americans United for the Separation of
Church and State. February 2005. http://www.au.org/site/PageServer?
pagename=cs_2005_02_special. Retrieved 2007-07-29. [dead link]
132.^ a b Wallis, Claudia (August 7, 2005). "The Evolution Wars". Time Magazine.
http://www.time.com/time/magazine/article/0,9171,1090909,00.html. Retrieved 2007-07-23.
133.^ ""The evolution wars" in Time". National Center for Science Education. August 11, 2005.
http://ncse.com/news/2005/08/evolution-wars-time-00696. Retrieved 2009-11-18.
• Journalism and the Debate Over Origins: Newspaper Coverage of Intelligent Design. Journal of
Media and Religion. 2006;5(1):49–61. doi:10.1207/s15328415jmr0501_3.
• Television wildlife programming as a source of popular scientific information: a case study of
evolution. Public Understanding of Science. 2006;15:131–152. doi:10.1177/0963662506060588.
• Sheppard, Pam (2005-08-10). "Time for evolution wars". Answers in Genesis.
http://www.answersingenesis.org/docs2005/0810time.asp. Retrieved 2009-02-16.
 134.^ a b DeWolf, David K. Intelligent Design Will Survive Kitzmiller v. Dover [PDF]. University of
Montana Law Review. May 4, 2007;68(1).
135.^
• Campbell, John Angus; Bill Marty (2005-08-25). "Does God have a place in class?:
Intelligent design ignites great debate". Calgary Herald. http://www.discovery.org/a/2920.
Retrieved 2009-02-16.
• The Politically Incorrect Guide to Darwinism and Intelligent Design . Regnery
Publishing; 2006. ISBN 1596980133. p. 273.
• The Evolution Wars: Who Is Fighting with Whom about What?. In: Robert B. Stewart.
Intelligent Design: William A. Dembski & Michael Ruse in Dialogue . Minneapolis, MN:
Fortress Press; 2007. ISBN 0800662180.
136.^ Methodological Naturalism and Philosophical Naturalism: Clarifying the Connection. Philo.
2000 [cited 2007-07-27];3(2):7–29.
137.^ Reason in the Balance: The Case Against Naturalism in Science, Law and Education .
InterVarsity Press; 1995. ISBN 0830819290.[Johnson positions himself as a "theistic realist" against
"methodological naturalism".]
138.^ Phillip E. Johnson (August 31, 1996). "Starting a Conversation about Evolution: Johnson,
Phillip". Access Research Network. http://www.arn.org/docs/johnson/ratzsch.htm. Retrieved 2008-
10-18. "My colleagues and I speak of 'theistic realism'—or sometimes, 'mere creation'—as the defining
concept of our [the ID] movement. This means that we affirm that God is objectively real as Creator,
and that the reality of God is tangibly recorded in evidence accessible to science, particularly in
biology."
139.^ See, for instance: Vuletic, Mark I. (February 1997). "Methodological Naturalism and the
Supernatural". Naturalism, Theism and the Scientific Enterprise: An Interdisciplinary Conference .
University of Texas, Austin.
http://www.utexas.edu/cola/depts/philosophy/faculty/koons/ntse/papers/Vuletic.html. Retrieved 2007-
07-27. http://web.archive.org/web/20080114094157/http://www.utexas.edu/cola/depts/philosophy/fac
ulty/koons/ntse/papers/Vuletic.html
140.^ a b Watanabe, Teresa (March 25, 2001). "Enlisting Science to Find the Fingerprints of a
Creator". Los Angeles Times. http://www.discovery.org/scripts/viewDB/index.php?
programs=CSCstories&command=view&id=613. Retrieved 2007-07-22. "[Phillip E. Johnson quoted]:
We are taking an intuition most people have and making it a scientific and academic enterprise ... We
are removing the most important cultural roadblock to accepting the role of God as creator."
141.^ Witnesses For The Prosecution [Reprint by Leadership U.]. World Magazine. November
30, 1996 [cited 2007-07-23];11(28):18.
142.^ Nickson, Elizabeth (January 10, 2003). "Let's Be Intelligent About Darwin". Christianity.ca.
The Evangelical Fellowship of Canada. http://www.christianity.ca/news/social-
issues/2004/03.001.html. Retrieved 2007-07-23. "[Phillip E. Johnson quoted]: Our strategy has been
to change the subject a bit so that we can get the issue of Intelligent Design, which really means the
reality of God, before the academic world and into the schools." [dead
link]http://web.archive.org/web/20070608233455/http://www.christianity.ca/news/social-
issues/2004/03.001.html
143.^ Buell, Jon; Hearn, Virginia, eds (March 1992). "Darwinism: Science or Philosophy" (PDF).
Darwinism: Scientific Inference or Philosophical Preference? (Symposium) . The Foundation for
Thought and Ethics, Dallas Christian Leadership, and the C. S. Lewis Fellowship.
http://ebd10.ebd.csic.es/pdfs/DarwSciOrPhil.pdf. Retrieved 2007-07-23.
144.^ Intelligent design's long march to nowhere. Templeton Foundation, Science & Theology
News; December 5, 2005 [cited 2007-07-23].
 145.^ What is wrong with intelligent design? [PDF]. Quarterly Review of Biology. 2007 [cited
2007-07-23];82(1):3–6. doi:10.1086/511656.
146.^ "What Is Wrong With Intelligent Design?". Science Daily. February 23, 2007.
http://www.sciencedaily.com/releases/2007/02/070222155420.htm. Retrieved 2007-07-23.
147.^ Murray, Michael J. (Forthcoming). "Natural Providence (or Design Trouble)" (PDF).
Franklin & Marshall College. http://edisk.fandm.edu/michael.murray/Providence.pdf. Retrieved 2007-
07-23.
148.^ Dembski, William A.. "What is the position of the NRCSE on the teaching of intelligent
design [ID] as an alternative to neo-Darwinian evolution in Nebraska schools?". Creighton University.
http://puffin.creighton.edu/NRCSE/NRCSEPosReID.html. Retrieved 2007-07-23.
149.^ Schönborn, Cardinal Christoph (October 2, 2005). "Catechetical Lecture at St. Stephan's
Cathedral, Vienna" (Reprint). Bring You To. http://www.bringyou.to/apologetics/p91.htm. Retrieved
2007-07-22. "Purpose and design in the natural world, [has] no difficulty [...] with the theory of
evolution [within] the borders of scientific theory."
150.^ Scott, Eugenie C. (December 7, 2000). "The Creation/Evolution Continuum". National
Center for Science Education. http://ncse.com/creationism/general/creationevolution-continuum.
Retrieved 2009-11-18.
151.^ Resseger, Jan (Chair) (March 2006). "Science, Religion, and the Teaching of Evolution in
Public School Science Classes" (PDF). Committee on Public Education and Literacy. National
Council of Churches. http://www.ncccusa.org/pdfs/evolutionbrochurefinal.pdf. Retrieved 2007-07-17.
152.^ Murphy, George L. (2002). "Intelligent Design as a Theological Problem" (Reprint).
Creighton University. http://puffin.creighton.edu/NRCSE/IDTHG.html. Retrieved 2007-07-21.
 153.^ Sheppard, Pam S. (February 4, 2006). "Intelligent design: is it intelligent; is it Christian?".
Answers in Genesis. http://www.answersingenesis.org/articles/wow/is-idm-christian. Retrieved 2007-
07-21.
154.^ Ross, Hugh. "More Than Intelligent Design". Facts for Faith. Reasons to Believe.
http://www.reasons.org/resources/fff/2002issue10/index.shtml#more_than_id. Retrieved 2007-07-
21.
155.^ Harun Yahya International (2007). "The "Intelligent Design" Distraction". Press release.
http://www.harunyahya.com/new_releases/news/intelligent_design.php. Retrieved 2007-07-20.
156.^ Wieland, Carl (August 30, 2002). "AiG's views on the Intelligent Design Movement".
Answers in Genesis. http://www.answersingenesis.org/docs2002/0830_IDM.asp. Retrieved 2007-07-
20.
157.^ a b Natan Slifkin (2006). The Challenge of Creation (New York: Yashar Books) 288 ff.
158.^ Miller, Kenneth. Debating Design. Cambridge University Press; 2004. The Flagellum
Unspun. p. 95.
159.^ Meyer, Stephen C. (December 1, 2002). "The Scientific Status of Intelligent Design: The
Methodological Equivalence of Naturalistic and Non-Naturalistic Origins Theories". Center for
Science and Culture, Discovery Institute. http://www.discovery.org/scripts/viewDB/index.php?
command=view&id=1780. Retrieved 2007-07-19.
160.^ Wüthrich, Christian (January 11, 2007). "Demarcating science vis-à-vis pseudoscience"
(PDF). Department of Philosophy. University of California at San Diego. Archived from the original on
2007-07-24.
http://web.archive.org/web/20070724203349/http://philosophyfaculty.ucsd.edu/faculty/wuthrich/teach
ing/2007_145/Lecture02.pdf. Retrieved 2007-07-19.
 161.^ Scientific Method in Practice. Cambridge UP; 2003. ISBN 0521017084. Chapters 5–8.
Discusses principles of induction, deduction and probability related to the expectation of consistency,
testability, and multiple observations. Chapter 8 discusses parsimony (Occam's razor)
162.^ Research Methods in Psychology. 8th ed. Wadsworth Publishing; 2005. ISBN
0534609767. Chapter 2. Discusses the scientific method, including the principles of falsifiability,
testability, progressive development of theory, dynamic self-correcting of hypotheses, and
parsimony, or "Occam's razor".
163.^ Kitzmiller v. Dover Area School District, cv 2688 (December 20, 2005)., 4: whether ID is
science. The ruling discusses central aspects of expectations in the scientific community that a
scientific theory be testable, dynamic, correctible, progressive, based upon multiple observations,
and provisional,
164.^ See, e.g., Mark Perakh (2005). "The Dream World of William Dembski's Creationism".
Skeptic. Talk Reason. pp. 54–65. http://talkreason.com/articles/Skeptic_paper.cfm.
165.^ Intelligent design fails to pass Occam's razor. Adding entities (an intelligent agent, a
designer) to the equation is not strictly necessary to explain events. See, e.g., Branden Fitelson, et
al.. How Not to Detect Design–Critical Notice: William A. Dembski The Design Inference. In: Robert T.
Pennock. Intelligent Design Creationism and Its Critics: Philosophical, Theological, and Scientific
Perspectives. MIT Press; 2001. p. 597–616.
166.^ See, e.g., Schneider, Jill E. (2005). "Thoughts on Evolution and Intelligent Design".
Department of Biological Sciences, Lehigh University.
http://www.lehigh.edu/~inbios/schneider/evolution.htm. "Q: Why couldn't intelligent design also be a
scientific theory? A: The idea of intelligent design might or might not be true, but when presented as
a scientific hypothesis, it is not useful because it is based on weak assumptions, lacks supporting
data and terminates further thought."
 167.^ The designer is not falsifiable, since its existence is typically asserted without sufficient
conditions to allow a falsifying observation. The designer being beyond the realm of the observable,
claims about its existence can be neither supported nor undermined by observation, making
intelligent design and the argument from design analytic a posteriori arguments. See, e.g., Kitzmiller
v. Dover Area School District, cv 2688 (December 20, 2005). Ruling, p. 22 and p. 77.
168.^ That intelligent design is not empirically testable stems from the fact that it violates a basic
premise of science, naturalism. See, e.g., Kitzmiller v. Dover Area School District, cv 2688
(December 20, 2005). Ruling, p. 22 and p. 66.
169.^ Intelligent design professes to offer an answer that does not need to be defined or
explained, the intelligent agent, designer. By asserting a conclusion that cannot be accounted for
scientifically, the designer, intelligent design cannot be sustained by any further explanation, and
objections raised to those who accept intelligent design make little headway. Thus intelligent design
is not a provisional assessment of data which can change when new information is discovered. Once
it is claimed that a conclusion that need not be accounted for has been established, there is simply
no possibility of future correction. The idea of the progressive growth of scientific ideas is required to
explain previous data and any previously unexplainable data. See, e.g., the brief explanation in
Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005). p. 66.
170.^ "Nobel Laureates Initiative" (PDF). The Elie Wiesel Foundation for Humanity. September
9, 2005. http://media.ljworld.com/pdf/2005/09/15/nobel_letter.pdf. Retrieved 2007-07-19. The
September 2005 statement by 38 Nobel laureates stated that: "Intelligent design is fundamentally
unscientific; it cannot be tested as scientific theory because its central conclusion is based on belief
in the intervention of a supernatural agent".
171.^ "Intelligent Design is not Science: Scientists and teachers speak out". University of New
South Wales. October 2005.
http://web.archive.org/web/20060614003243/www.science.unsw.edu.au/news/2005/intelligent.html.
Retrieved 2009-01-09. The October 2005 statement, by a coalition representing more than 70,000
Australian scientists and science teachers said: "intelligent design is not science" and called on "all
schools not to teach Intelligent Design (ID) as science, because it fails to qualify on every count as a
scientific theory".
172.^ "Creationism and the Daubert test?". PZ Myers, Pharyngula.org. May 21, 2005.
http://pharyngula.org/index/science/comments/creationism_and_the_daubert_test/.
173.^ Nagel, Thomas. "Public Education and Intelligent Design", Philosophy and Public Affairs,
Vol. 36, no. 2, 2008, pp. 196–197.
174.^ a b c Kitzmiller v. Dover Area School District, cv 2688 (December 20, 2005)., 4. Whether
ID is Science, p. 87
175.^ Free Speech on Evolution Campaign Main Page Discovery Institute, Center for Science
and Culture.
176.^ Hawks, John (August 2005). "The President and the teaching of evolution". John Hawks
Weblog. http://johnhawks.net/weblog/topics/creation/bush_intelligent_design_2005.html. Retrieved
2007-07-19.
177.^ Skeptic: eSkeptic: Thursday, April 17th, 2008
178.^ Goodstein, Laurie (December 4, 2005). "Intelligent Design Might Be Meeting Its Maker".
New York Times. http://www.nytimes.com/2005/12/04/weekinreview/04good.html?
ex=1291352400&en=feb5138e425b9001&ei=5088&partner=rssnyt&emc=rss. Retrieved 2007-07-
19.
179.^ Statement from the Council of the Biological Society of Washington at the Wayback
Machine (archived September 26, 2007).
 180.^ Meyer, S.C.. The origin of biological information and the higher taxonomic categories.
Proceedings of the Biological Society of Washington . 2004 [cited 2007-05-10];117(2):213–239.
181.^ The Sternberg peer review controversy and several similar academic disputes are the
subject of the 2008 documentary "Expelled: No Intelligence Allowed".
182.^ Dembski, William A. (2001). "Is Intelligent Design a Form of Natural Theology?". Design
Inference Website. http://www.designinference.com/documents/2001.03.ID_as_nat_theol.htm.
Retrieved 2007-07-19.
183.^ McMurtie, Beth (December 21, 2001). "Darwinism Under Attack". The Chronicle Of Higher
Education. http://chronicle.com/free/v48/i17/17a00801.htm. Retrieved 2008-12-10.
184.^ Ruling, Kitzmiller v. Dover Area School District 4: whether ID is science
185.^ Simulating evolution by gene duplication of protein features that require multiple amino
acid residues. Protein Science. October 2004 [cited 16 March 2009];13(10):2651–64.
doi:10.1110/ps.04802904. PMID 15340163. PMC 2286568.
186.^ Kitzmiller v. Dover Area School District, 04 cv 2688 (December 20, 2005). 4: whether ID is
science
187.^ Simple evolutionary pathways to complex proteins. Protein Science. September 2005
[cited 16 March 2009];14(9):2217–2225. doi:10.1110/ps.041171805. PMID 16131652.
188.^ Kitzmiller v. Dover Area School District, October 19, 2005, AM session Kitzmiller
Testimony, Behe
189.^ "Peer-Reviewed, Peer-Edited, and other Scientific Publications Supporting the Theory of
Intelligent Design (Annotated)". Discovery Institute. July 2007.
http://www.discovery.org/scripts/viewDB/index.php?command=view&id=2640&program=CSC%20-
%20Scientific%20Research%20and%20Scholarship%20-%20Science. Retrieved 2007-07-17.
 190.^ Is It Science Yet?: Intelligent Design Creationism and the Constitution [PDF]. Washington
University Law Quarterly. 2005 [cited 2007-07-18];83(1). "ID leaders know the benefits of submitting
their work to independent review and have established at least two purportedly "peer-reviewed"
journals for ID articles. However, one has languished for want of material and quietly ceased
publication, while the other has a more overtly philosophical orientation. Both journals employ a weak
standard of "peer review" that amounts to no more than vetting by the editorial board or society
fellows."
191.^ Isaak, Mark (2006). "Index to Creationist Claims". TalkOrigins Archive.
http://www.talkorigins.org/indexcc/CI/CI001_4.html. "With some of the claims for peer review, notably
Campbell and Meyer (2003) and the e-journal PCID, the reviewers are themselves ardent supporters
of intelligent design. The purpose of peer review is to expose errors, weaknesses, and significant
omissions in fact and argument. That purpose is not served if the reviewers are uncritical"
192.^ Dembski, William A. (April 2002). "Detecting Design in the Natural Sciences". Intelligent
Design?. Natural History Magazine. http://www.actionbioscience.org/evolution/nhmag.html.
Retrieved 2007-07-18.
193.^ Shostak, Seth (December 2005). "SETI and Intelligent Design". Space.com.
http://www.space.com/searchforlife/seti_intelligentdesign_051201.html. Retrieved 2007-07-18. "In
fact, the signals actually sought by today's SETI searches are not complex, as the ID advocates
assume. [...] If SETI were to announce that we're not alone because it had detected a signal, it would
be on the basis of artificiality"
194.^ "For human artifacts, we know the designer's identity, human, and the mechanism of
design, as we have experience based upon empirical evidence that humans can make such things,
as well as many other attributes including the designer's abilities, needs, and desires. With ID,
proponents assert that they refuse to propose hypotheses on the designer's identity, do not propose
a mechanism, and the designer, he/she/it/they, has never been seen. In that vein, defense expert
Professor Minnich agreed that in the case of human artifacts and objects, we know the identity and
capacities of the human designer, but we do not know any of those attributes for the designer of
biological life. In addition, Professor Behe agreed that for the design of human artifacts, we know the
designer and its attributes and we have a baseline for human design that does not exist for design of
biological systems. Professor Behe's only response to these seemingly insurmountable points of
disanalogy was that the inference still works in science fiction movies".— Kitzmiller v. Dover Area
School District, cv 2688 (December 20, 2005)., p. 81
195.^ Edis, Taner (March/April 2001). "Darwin in Mind: Intelligent Design Meets Artificial
Intelligence". Skeptical Inquirer Magazine. http://www.csicop.org/si/2001-03/intelligent-design.html.
Retrieved 2007-07-17. http://web.archive.org/web/20011018142820/http://www.csicop.org/si/2001-
03/intelligent-design.html
196.^ "Primer: Intelligent Design Theory in a Nutshell". Intelligent Design and Evolution
Awareness Center. 2007. http://www.ideacenter.org/contentmgr/showdetails.php/id/1136. Retrieved
2007-07-14.
197.^ a b Scott, Eugenie C.; Branch, Glenn (September 2002). ""Intelligent Design" Not
Accepted by Most Scientists". National Center for Science Education.
http://ncse.com/creationism/general/intelligent-design-not-accepted-by-most-scientists. Retrieved
2009-11-18.
198.^ See, for instance: Man Come Of Age: Bonhoeffer's Response To The God-Of-The-Gaps.
Journal of the Evangelical Theological Society. 1971;14:203–220.
199.^ Intelligent Design on Trial: Kitzmiller v. Dover. National Center for Science Education.
October 17th, 2008
200.^ Tammy Kitzmiller, et al. v. Dover Area School District, et al.' , [3] .
 201.^ Powell, Michael (December 21, 2005). "Judge Rules Against 'Intelligent Design'". The
Washington Post. http://pewforum.org/news/display.php?NewsID=5945. Retrieved 2007-09-
03. http://web.archive.org/web/20070928055938/http://pewforum.org/news/display.php?
NewsID=5945
202.^ Crowther, Robert (December 20, 2005). "Dover Intelligent Design Decision Criticized as a
Futile Attempt to Censor Science Education". Evolution News & Views. Discovery Institute.
http://www.evolutionnews.org/2005/12/dover_intelligent_design_decis.html. Retrieved 2007-09-03.
203.^ "Judge rules against 'intelligent design'". Associated Press. MSNBC. December 20, 2005.
http://www.msnbc.msn.com/id/10545387/. Retrieved 2008-10-17.
204.^ Provonsha, Matthew (September 21, 2006). "Godless: The Church of Liberalism".
eSkeptic. http://www.skeptic.com/eskeptic/06-09-21.html. Retrieved 2007-09-03.
205.^ Padian, Kevin; Nick Matzke (January 4, 2006). "Discovery Institute tries to "swift-boat"
Judge Jones". National Center for Science Education.
http://ncse.com/creationism/general/discovery-institute-tries-to-swift-boat-judge-jones. Retrieved
2009-11-18.
206.^ Raffaele, Martha (December 20, 2005). "Intelligent design policy struck down". Dallas
Morning News.
http://www.dallasnews.com/sharedcontent/dws/dn/latestnews/stories/122105dnnatidesign.780fc9a.ht
ml. Retrieved 2007-09-03.
207.^ Articles: Editor's Note: Intelligent Design Articles [PDF]. University of Montana Law
Review. April 10, 2007 [cited 2008-10-16];68(1).
208.^ Irons, Peter. Disaster In Dover: The Trials (And Tribulations) Of Intelligent Design [PDF].
University of Montana Law Review. April 27, 2007 [cited 2008-10-16];68(1).
 209.^ DeWolf, David K; West, John G; Luskin, Casey. Rebuttal to Irons [PDF]. University of
Montana Law Review. April 27, 2007 [cited 2008-10-16];68(1).
210.^ "American Academy of Religion on teaching creationism". National Center for Science
Education. July 23rd, 2010. http://ncse.com/news/2010/07/american-academy-religion-teaching-
creationism-005712. Retrieved 2010-08-09.
211.^ "The dangers of creationism in education". Council of Europe.
http://assembly.coe.int/Main.asp?link=/Documents/WorkingDocs/Doc07/EDOC11297.htm. Retrieved
2007-08-03.
212.^ "NCSE Resource—Council of Europe approves resolution against creationism". National
Center for Science Education. October 4, 2007. http://ncse.com/news/2007/10/council-europe-
approves-resolution-against-creationism-001200. Retrieved 2009-11-18.
• "Council of Europe firmly opposes creationism in school" . Reuters. 2007-10-04.
http://uk.reuters.com/article/scienceNewsMolt/idUKL0417855220071004. Retrieved 2007-
10-05.
213.^ "WIRED Magazine response". Illustra Media.
http://www.illustramedia.com/ID01WiredMagPage.htm. Retrieved 2007-07-13. "It's also important
that you read a well developed rebuttal to Wired's misleading accusations. Links to both the article
and a response by the Discovery Institute (our partners in the production of Unlocking the Mystery of
Life and The Privileged Planet)"
214.^ Meyer, Stephen C.; Allen, W. Peter (July 15, 2004). "Unlocking the Mystery of Life".
Center for Science and Culture, Discovery Institute.
http://www.discovery.org/scripts/viewDB/index.php?command=view&id=2116&program=CSC%20-
%20Video%20and%20Curriculum%20-%20Multimedia. Retrieved 2007-07-13.
 215.^ Randerson, James (November 27, 2006). "Revealed: rise of creationism in UK schools".
London: Guardian. http://www.guardian.co.uk/science/2006/nov/27/controversiesinscience.religion.
Retrieved 2008-10-17.
216.^ "'Design' attack on school science". BBC News. September 29, 2006.
http://news.bbc.co.uk/1/hi/education/5392096.stm. Retrieved 2007-07-13.
217.^ "Written Answers". Daily Hansard. Parliament of the United Kingdom. November 1, 2006.
http://www.publications.parliament.uk/pa/cm200506/cmhansrd/cm061101/text/61101w0010.htm#061
1021004183. Retrieved 2007-07-13.
218.^ "Schools: Intelligent Design". Daily Hansard. Parliament of the United Kingdom.
December 18, 2006.
http://www.publications.parliament.uk/pa/ld200607/ldhansrd/text/61218w0006.htm. Retrieved 2007-
07-13.
219.^ "NCSE Resource—Guidance on creationism for British teachers". NCSE. September 25,
2007. http://ncse.com/news/2007/02/guidance-creationism-british-teachers-001170. Retrieved 2009-
11-18.
220.^ "How can we answer questions about creation and origins?" (PDF). Qualifications and
Curriculum Authority for England. 2006. http://www.qca.org.uk/libraryAssets/media/qca-06-
2728_y9_science_religion_master.pdf. Retrieved 2007-10-01.
221.^ Prime Minister's Office (21 June 2007). "NoCreSciEd - epetition reply". London: Her
Majesty's Stationery Office. http://www.number10.gov.uk/Page12021. Retrieved 2007-07-13.
222.^ Lengagne, Guy (June 8, 2007). "The dangers of creationism in education". Committee on
Culture, Science and Education, Socialist Group, Parliamentary Assembly . Council of Europe, Doc.
11297. http://www.assembly.coe.int/Main.asp?
link=/Documents/WorkingDocs/Doc07/EDOC11297.htm. Retrieved 2008-10-17.
 223.^ Henry, Lesley-Anne (September 26, 2007). "Tussle of Biblical proportions over
creationism in Ulster classrooms". Belfast Telegraph.
http://www.belfasttelegraph.co.uk/news/education/tussle-of-biblical-proportions-over-creationism-in-
ulster-classrooms-13479246.html. Retrieved 2007-10-01.
*"Viewpoint: The world, according to Lisburn folk". Belfast Telegraph. September 27, 2007.
http://www.belfasttelegraph.co.uk/opinion/viewpoint-the-world-according-to-lisburn-folk-
13479536.html. Retrieved 2007-10-01.
224.^ "Dup Call For Schools To Teach Creation Passed By Council—Lisburn Today". Ulster Star.
September 26, 2007. http://www.lisburntoday.co.uk/news/Creation-Motion-Passed-By-
Council.3233193.jp. Retrieved 2008-10-17.
225.^ Enserink, Martin. Evolution Politics: Is Holland Becoming the Kansas of Europe?. Science.
June 3, 2005;308(5727):1394b. doi:10.1126/science.308.5727.1394b. PMID 15933170.
226.^ "Cabinet ministers announced (update 2)". DutchNews.nl. February 13, 2007.
http://www.dutchnews.nl/news/archives/print/001501.php. Retrieved 2008-05-31.
227.^ De Morgen, May 23, 2005
228.^ a b Edis, Taner (January 2008). "The History of Science Society : The Society". Islamic
Creationism: A Short History. History of Science Society. pp. Newsletter.
http://www.hssonline.org/publications/Newsletter2008/NewsletterJanuary2008Creationism.html.
Retrieved 2008-02-23.
229.^ Edis, Taner. "Cloning Creationism in Turkey". National Center for Science Education.
http://ncse.com/rncse/19/6/cloning-creationism-turkey. Retrieved 2009-11-18.
230.^ Jones, Dorian L (March 12, 2008). "Turkey's survival of the fittest". ISN Security Watch.
http://www.isn.ethz.ch/isn/Current-Affairs/Security-Watch/Detail/?id=54183&lng=en. Retrieved 2008-
03-13.
 231.^ Smith, Deborah (October 21, 2005). "Intelligent design not science: experts". Sydney
Morning Herald. http://www.smh.com.au/news/national/intelligent-design-not-science-
experts/2005/10/20/1129775902661.html. Retrieved 2007-07-13.

[edit] Further reading

• Gross PR; Forrest B. Creationism's Trojan Horse: the Wedge of Intelligent Design . Oxford
[Oxfordshire]: Oxford University Press; 2004. ISBN 0-19-515742-7. Preview at Google Books
• Humes E. Monkey Girl: Evolution, Education, Religion, and the Battle for America's Soul .
New York, N.Y: ECCO; 2007. ISBN 0-06-088548-3.
• Edis T; Young M. Why Intelligent Design Fails: A Scientific Critique of the New Creationism .
New Brunswick, N.J: Rutgers University Press; 2006. ISBN 0-8135-3872-6. Preview at Google Books
• Why Darwin Matters: The Case Against Intelligent Design . New York: Henry Holt; 2007.
ISBN 978-0-8050-8306-4.
• Slack G. The Battle over the Meaning of Everything: Evolution, Intelligent Design, and a
School Board in Dover, PA. San Francisco: Jossey-Bass; 2007. ISBN 0-7879-8786-7. Preview at
Google Books
ID perspectives
• Access Research Network
• Design Inference: The website of William A. Dembski
• Discovery Institute, Center for Science and Culture (Hub of the intelligent design movement)
• EvolutionNews.org Discovery Institute website tracking media coverage of intelligent design.
• ID The Future A multiple contributor weblog by Discovery Institute fellows.
 • International Society for Complexity, Information, and Design (ISCID)
• Uncommon Descent William Dembski's blog
Non-ID perspectives
• Scientific American - 15 Answers to Creationist Questions
• ACLU site on Intelligent Design
• Intelligent Design?
http://web.archive.org/web/20070620122151/http://www.naturalhistorymag.com/darwinanddesign.ht
ml special feature in the Natural History Magazine
• Internet Encyclopedia of Philosophy: Design Arguments for the Existence of God
• National Center for Science Education What Is Intelligent Design Creationism?
• Resolution from the American Association for the Advancement of Science
• Science and Creationism: A View from the National Academy of Sciences Second Edition
(1999)
• Talk Origins Archive (Archive of the UseNet discussion group talk.origins)
• 139 page in-depth analysis of intelligent design, irreducible complexity, and the book Of
Pandas and People by the Kitzmiller v. Dover Area School District judge
• Kitzmiller: An Intelligent Ruling on 'Intelligent Design' , JURIST
• ID and Creationism
• The Design Argument Elliott Sober, 2004.
• Short discussion on Natural Knowledge and Natural Design as a contrast to Intelligent
Designhttp://web.archive.org/web/20080307082606/http://richarddawkins.net/article,129,Natural-
Knowledge-and-Natural-Design,Richard-Dawkins by Richard Dawkins
Media articles
• Judgment Day: Intelligent Design on Trial A PBS-NOVA documentary on the Dover, PA
Intelligent Design trial in 2005.(PBS)
• Discovery's Creation An overview of the origin of the intelligent design movement. ( Seattle
Weekly)
• Intelligent Design vs. Evolution debate between paleontologist Peter Ward and Stephen
Meyer co-founder of the Discovery Institute
• Intelligent Design Deja Vu What would "intelligent design" science classes look like? All we
have to do is look inside some 19th century textbooks. (The Washington Post)
• How the media have covered ID[dead link] (Columbia Journalism Review)
• Banned in biology class: intelligent design (Christian Science Monitor)
• Devolution (The New Yorker) The Evolution Debate
• (The New York Times)
• Debating Evolution in the Classroom (NPR)
• Darwin Victorious (TIME)
• Intelligent Design: Scientific Inquiry or Religious Indoctrination? (Justice Talking)
• Intelligent Judging—Evolution in the Classroom and the Courtroom (New England Journal of
Medicine)
 [show]

Links to related articles

 [show]
v•d•e
Creationism topics

Types Young Earth Creationism · Old Earth creationism · Progressive creationism

design

Genesis
interpretations Genesis creation narrative · Framework interpretation · Day-Age theory

Related concepts Creation myth · Omphalos hypothesis · Specified complexity · Irreducible complexity
Theistic realism · Intelligent designer

Pseudoscience Creation science · Creationist cosmologies · Creation biology · Created kinds

canopy · Modern geocentrism

Miscellanea Creation-evolution controversy · History of creationism · Creation and evolution in pu

Quotes about creation and evolution · Teach the Controversy

[show]
v•d•e
Pseudoscience
 Retrieved from "http://en.wikipedia.org/wiki/Intelligent_design"

Categories: Intelligent design | Creationist objections to evolution | Denialism | Pseudoscience |

Arguments for the existence of God
 W000

Dessein intelligent
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Le dessein intelligent (Intelligent Design en anglais[1]) est la croyance selon laquelle « certaines
observations de l'univers et du monde du vivant sont mieux expliquées par une cause intelligente que par
des processus aléatoires tels que la sélection naturelle »[2]. Cette thèse a été développée par le Discovery
Institute, un cercle de réflexion conservateur chrétien américain. Le dessein intelligent est présenté comme
une théorie scientifique par ses promoteurs[3], mais dans le monde scientifique, il est considéré comme
relevant de la pseudo-science, tant par des arguments aussi bien internes à la biologie (les promoteurs du
dessein intelligent apparaissant aux biologistes comme ne tenant pas compte de nombreuses observations)
qu'épistémologiques (en particulier le critère de réfutabilité de Karl Popper).
 La plupart des commentateurs et des scientifiques y voient une résurgence du créationnisme[4],
dissimulée sous une apparence de scientificité ; le biologiste britannique Richard Dawkins le désigne même
sous le nom de « créationnisme affublé d'un costume bon marché ». Le dessein intelligent est désormais
classé aux États-Unis dans les théories néo-créationnistes, en particulier suite à la publication du Wedge
document (voir objectifs et stratégie). D'un point de vue idéologique, les deux thèses sont apparentées
(intervention d'une puissance supérieure).
Sommaire
[masquer]
• 1 Concept
• 2 Objectifs et stratégie
• 3 Caractère scientifique
• 4 Récentes tentatives scientifiques
• 5 Un puissant lobbying
• 6 En Europe
• 6.1 En Belgique
• 6.2 En Suisse
• 6.3 En France
• 7 Parodies
• 8 Notes et références
• 9 Voir aussi
• 9.1 Articles connexes
• 9.2 Bibliographie
• 9.3 Liens externes
Concept [modifier]
Les défenseurs de ce concept, dans le domaine de la biologie et de la biochimie, affirment que la
théorie scientifique traditionnelle de l'évolution par voie de sélection naturelle ne suffit pas pour rendre
compte de l'origine, de la complexité et de la diversité de la vie. En particulier, les partisans de ce concept
estiment qu'il existe des exemples de complexité irréductible qui ne peuvent être expliqués par l'évolution, et
plaident donc pour la théorie du dessein intelligent.

Objectifs et stratégie [modifier]

Articles détaillés : Intelligent design movement et Stratégie du coin.
Les objectifs du dessein intelligent ont été clairement définis par le think tank à l'origine du
mouvement, le Discovery Institute, dans un document à usage interne The Wedge. Des fuites permettront
finalement sa diffusion en 1999.
Les objectifs du Discovery Institute et du dessein intelligent sont de nature politique et religieuse et
ont un lien sans équivoque avec le dessein intelligent et le fondamentalisme religieux ; ils se retrouvent de
manière chiffrée en termes de retombées médiatiques et législatives avec des délais pour les atteindre [5] :
• Objectifs principaux :
• Vaincre le matérialisme scientifique et ses héritages moraux, culturels et
scientifiques ;
• Remplacer les explications matérialistes par la compréhension théistique que la
nature et l’être humain sont créés par Dieu.
• Objectifs sur cinq ans :
 • Voir la théorie du dessein intelligent devenir une alternative acceptée dans les
sciences, et des recherches scientifiques menées depuis la perspective de la théorie du
dessein ;
• Assister au commencement de l’influence de la théorie du dessein dans des sphères
autres que la science naturelle ;
• Voir de nouveaux débats majeurs dans l’éducation, les sujets relatifs à la vie, la
responsabilité pénale et personnelle poussées au front de l’agenda national.
• Objectifs sur vingt ans :
• Voir la théorie du dessein intelligent comme la perspective dominante dans la
science ;
• Voir des applications de la théorie du dessein dans des champs spécifiques incluant
la biologie moléculaire, la biochimie, la paléontologie, la physique et la cosmologie dans les
sciences naturelles ; la psychologie, l’éthique, la politique, la théologie, la philosophie, et les
matières littéraires ; voir son influence dans les arts.
Le principe est de faire passer la doctrine religieuse pour une science et de semer la confusion dans
les cercles scientifiques, avant de rayonner dans toutes les sphères de la société grâce en particulier à une
campagne de publicité et de façonnage d'opinion (« publicity and opinion making »).
L'exposition au grand public des visées fondamentalistes du dessein intelligent a porté un coup
majeur à ses promoteurs qui, faute de pouvoir nier l'existence du document, en ont proposé une relecture
édulcorée sur leur site où le Discovery Institute se défend entre autres de vouloir l'établissement d'une
théocratie[6].
Caractère scientifique [modifier]
L'Académie nationale des sciences des États-Unis et le Centre national pour l'éducation scientifique
des États-Unis (National Center for Science Education) ont décrit le dessein intelligent comme étant de la
pseudo-science. Les critiques affirment que le dessein intelligent est une tentative pour réexprimer les
dogmes religieux sous forme pseudo-scientifique, afin de forcer les institutions scolaires à enseigner la
théorie du créationnisme. Le chef astronome du Vatican, le révérend George Coyne, a affirmé que le dessein
intelligent « n'est pas de la science, même s'il en a la prétention »[7].
Alors que le modèle scientifique de l'évolution est corroboré par des faits observables et
reproductibles, comme le principe des mutations, du flux génétique, de la dérive génétique, de la sélection
naturelle et de la spéciation, les critiques démontrent que l'hypothèse du dessein intelligent repose sur des
éléments qui ne peuvent être reproduits ni observés, et ne répond donc pas au principe de réfutabilité de
Karl Popper : une théorie ne peut être qualifiée de scientifique que si elle permet des prédictions pouvant
être invalidées par l'expérimentation.
En introduisant une explication externe, le dessein intelligent ne respecte pas non plus un autre
principe scientifique, celui du Rasoir d'Occam, car il crée une entité supplémentaire pour expliquer un
phénomène sans que cela apporte de lumière supplémentaire en l'état.
L'hypothèse du dessein intelligent ne fournit par ailleurs aucune explication sur les anomalies
manifestes de la nature : point aveugle de l'œil humain (et plus généralement des vertébrés), chiasma
optique, risque de fausse couche, grossesse extra-utérine, pouce du panda, etc...
Cette hypothèse ne fait pas davantage l'unanimité dans les milieux religieux, les libéraux considérant
que la démarche scientifique n'est pas censée être mise sur le même plan que le dogme, et en sens inverse
certains dogmatiques considérant qu'une Vérité révélée ne requiert pas de preuves d'ordre scientifique.
 Enfin, la démarche adoptée par les tenants du dessein intelligent s'oppose à la démarche
scientifique en ce qu'ils ne tentent pas de démontrer que leurs arguments sont valables mais demandent à
leurs détracteurs de prouver qu'ils ne le sont pas.

Récentes tentatives scientifiques [modifier]

Article détaillé : Néo-créationnisme.
Ces critiques assez solides ont provoqué des dissensions dans les rangs mêmes des tenants du
dessein intelligent, dont une frange érige cette notion en principe théorique au sein des raisonnements
scientifiques plutôt qu'en une science en elle-même. C'est ainsi que l'on voit poindre, depuis quelques
années, des tentatives plus rigoureuses – qu'on évoque souvent sous le nom de « créationnisme vieille
terre » (Old-Earth Creationism), par opposition au « créationnisme terre jeune » (Young-Earth Creationism) –
et plus séparées des raisonnements dogmatiques (auxquels on préfère un enseignement des « implications
théologiques » du modèle de la création « à côté » de ce modèle), des tentatives qui se veulent
prétendument réfutables et testables – scientifiques, donc –, et tentent de démontrer une certaine intégrité
scientifique.
Le courant, porté par des érudits rationalistes, mais exclusivement chrétiens (puisqu'il prétend
pouvoir appuyer la véracité de la Bible), est notamment représenté aux États-Unis et au Canada par
l'astrophysicien Hugh Ross, le biochimiste Fazale Rana, mais également par le théologien et exégète Walter
Kaiser Jr et le philosophe, épistémologue et théologien Kenneth Samples. Leur approche répond (quoique
partiellement) aux avis des autorités judiciaires et académiques qualifiées par eux de « non sectaires », et
donc dignes de foi. Préférant du coup distinguer leur modèle qui se veut scientifique, des implications
théologiques de ce modèle, avec pour visée de promouvoir à la fois, et de manière séparée, une démarche
scientifique et un enseignement religieux.
 Cet extrait d’un texte par Hugh Ross[8] indique d'ailleurs l'aspect novateur et quelque peu dissident
que ce mouvement veut donner au concept du dessein intelligent (toutes les parenthèses sont ajoutées) :
« Dans les échelons les plus hauts de la recherche et de l'érudition, la fragilité des théories
naturalistes est de plus en plus librement admise. Même si les tenants [du dessein intelligent]
ne font rien pour exposer les inadéquations et les incohérences de l'explication [des théories
naturalistes] sur le cosmos et sur la vie, le naturalisme peut s'autodétruire.

Gagner l'argument du dessein sans identifier l'auteur du dessein offre, au mieux, un modèle des
origines brouillon. Un tel modèle n'a qu'un impact positif léger, si tant est qu'il en ait un, sur la
communauté des scientifiques et des autres érudits. Un tel modèle ne s'offre pas à la
vérification, ni ne peut faire de prédictions spécifiques et crédibles. De chaque côté, les érudits,
et en particulier les scientifiques, seraient réticents à admettre la viabilité du concept et à lui
accorder une attention sérieuse. Cette approche ne leur offre pas, non plus, de direction
spirituelle.

[…]

L'expérience me persuade du fait que le moment est idéal pour une approche directe, un saut
unique dans la bataille des origines. Introduire un modèle de la création bibliquement fondé et
scientifiquement vérifiable représente un tel saut . Il allie un dynamisme à la fois scientifique et
spirituel. Il bâtit la confiance, stimule la discussion, relâche une tension inutile au sujet d'un
agenda religieux caché, et dirige l'attention de manière rapide et fructueuse vers la mise à
l'épreuve et les prédictions.
 […]
Une discussion et une critique honnêtes des divers modèles des origines, y compris des divers
modèles chrétiens des origines, peuvent avoir un impact positif sur la poursuite de l'effort
scientifique dans son ensemble. Les dogmes retranchés et le « politiquement correct » ont
depuis de trop nombreuses années entravé le progrès vers un corps de savoir. […]

Il y a ici une opportunité d'exemplifier la liberté qui existe en Christ. La vérité ne constitue pas
une menace pour le chrétien. La vérité dans le domaine scientifique, qui peut être directement
ou indirectement mise à l'épreuve, sera toujours cohérente avec la vérité dans le domaine
spirituel. Et, malgré les protestations de toutes parts, la vérité dans la nature doit être reliée à
quelque chose, ou Quelqu'Un, au-delà du monde naturel – le quelque chose ou Quelqu'Un
responsable de l'existence et des caractéristiques de la nature. »

— Dr Hugh Ross, More Than Intelligent Design

Travaillant depuis une quinzaine d'années, ce courant ne rencontre pas la même opposition érudite
ni la même focalisation médiatique que le créationnisme terre jeune, puisqu'il est d'une part plutôt épars, ne
jouissant que beaucoup plus faiblement d'une structure de lobby (des scientifiques soutiennent cette vision
des choses sans être engagés dans un mouvement structuré), et d'autre part parce qu'il s'oppose clairement,
que ce soit sur des bases religieuses exégétiques ou scientifiques empiriques, à des conceptions
(pseudo)scientifiques phares des milieux protestants fondamentalistes (suprématie de la Bible sur la
science, non-valeur des méthodes scientifiques de datation, inexistence du Big Bang, Terre vieille de 6 000
ans, génération spontanée et simultanée des animaux et des êtres humains ensemble le même jour (de 24
heures), Déluge historique et mondial, etc.) en se souciant plus d'une harmonie avec les procédures et le
monde scientifiques. Il faut croire que ces traits, à des degrés divers, expliquent probablement le succès
grandissant que le mouvement remporte chez les scientifiques, les étudiants et dans les communautés
chrétiennes, s'essaimant jusqu'en Amérique du Sud, en Europe orientale, en Afrique du Sud et en Australie
et Nouvelle-Zélande.

Un puissant lobbying [modifier]

Dans les années 1990 et 2000, suite à un important lobbying et alors que le créationnisme était
depuis 1987 retiré officiellement de tous les manuels scolaires après une longue bataille juridique, de
nombreux États américains (Pennsylvanie, Kansas, Géorgie, Mississippi, etc.) sont revenus sur la question
et ont tenté de faire en sorte que les théories issues du darwinisme soient présentées aux écoliers comme
de simples théories concurrentes, mais en rien supérieures, au dogme créationniste, cette démarche a été
soutenue notamment par George W. Bush. Certaines de ces démarches ont été renversées au niveau des
conseils d'État sur l'éducation.
Article détaillé : Kitzmiller v. Dover Area School.
À Dover, en Pennsylvanie, le bureau de l'éducation locale a décidé en octobre 2004 de dire à tous
les élèves de 14 ans que la théorie de l'évolution n'est qu'une simple théorie en proposant qu'un texte leur
soit lu avant le début du cours de biologie abordant le sujet. Un livre de vulgarisation du dessein intelligent,
Of Pandas and People (Des pandas et des Hommes), est même proposé à ceux désirant approfondir la
question. Cependant le tribunal fédéral de Harrisburg en Pennsylvanie a conclu le 20 décembre 2005 que
l'enseignement du dessein intelligent comme une alternative à la théorie de l'évolution de Darwin dans les
classes de science des écoles est anticonstitutionnel. Pour rendre ce jugement, le juge John Jones s'est
basé sur le fait que le dessein intelligent n'est qu'une forme déguisée de croyance religieuse et l'enseigner
enfreint la séparation de Cultes et de l'État et l'interdiction de promouvoir une religion quelconque dans
l'éducation publique aux États-Unis[9]. Il a été très critique sur le comportement du bureau d'éducation de
Dover en déclarant que « les citoyens de la région de Dover sont mal servis par les membres de leur bureau
d'éducation ayant voté en faveur du dessein intelligent ». Il a également accusé certains parents du conseil
scolaire de Dover d'avoir menti et déclare à ce sujet qu'« il est ironique que plusieurs de ces individus, qui
affichent ouvertement et fièrement leurs convictions religieuses en public, aient menti systématiquement
pour déguiser leur véritable but. » Suite à ce jugement, le juge John E. Jones III a reçu plusieurs menaces de
mort de tenants du dessein intelligent[10].
En Europe, en mai 2005, la ministre néerlandaise de l'éducation, Maria van der Hoeven, a tenu des
propos à ce sujet en invoquant elle aussi la théorie du dessein intelligent qui cherche d'abord à établir
« scientifiquement » le fait que la nature semble être « pensée » avant de se hasarder à sous-entendre par
qui elle l'a été. Cependant, elle n'a pas été suivie par le reste de son gouvernement.
Le dessein intelligent et le créationnisme sont des théories qui connaissent un certain succès auprès
des protestants, mais relativement peu auprès des catholiques pour qui l'Ancien Testament ne doit pas se
lire au premier degré et où le pape Jean-Paul II, en 1996, a admis que la théorie de l'évolution était « plus
qu’une hypothèse ». Son successeur Benoît XVI semble en revanche promouvoir le retour de la religion au
cœur de la démarche scientifique[11].

En Europe [modifier]
Plusieurs établissement scolaires belges[12], français[13] et suisses[14] ont reçu en 2007 un livre
s'intitulant l'Atlas de la création[15]. Son auteur Harun Yahya (de son vrai nom Adnan Oktar) est un turc
musulman qui a écrit nombre de livres sur le sujet et conteste l'évolution et toute idéologie s'opposant à
« Dieu ». De nombreuses références au Coran et l'affirmation fausse, répétée à longueur de pages, qu'il
n'existerait aucune preuve scientifique de l'évolution servent d'argumentation. Pour appuyer sa thèse, il
utilise nombre de photos de fossiles et d'animaux actuels ressemblant aux restes fossilisés, mis côte à côte.
Il affirme qu'il s'agit d'animaux identiques et que par conséquent, il n'y a pas eu d'évolution. Notons
cependant que Harun Yahya a aussi accusé le dessein intelligent d'être un instrument de Satan[16].
Le sénateur français Guy Lengagne (PS) a vu son rapport Les dangers du créationnisme dans
l'éducation retiré au dernier moment de l'agenda de la réunion du Conseil de l'Europe en juin 2007, sous la
pression du parlementaire belge ultraconservateur Luc Van den Brande, président du Conseil[17]. Ses
travaux, qui s'inscrivaient dans la continuité de ceux d'Andrew McIntosh, ont finalement fait l'objet d'une
résolution intitulée : « Dangers du créationnisme dans l’éducation ». Celle-ci, adoptée le 4 octobre 2007,
marque la position du Conseil de l'Europe vis-à-vis du dessein intelligent de la façon suivante :
« Le créationnisme présente de multiples facettes contradictoires. L’ intelligent design (dessein
intelligent), dernière version plus nuancée du créationnisme, ne nie pas une certaine évolution.
Cependant l’intelligent design, présenté de manière plus subtile, voudrait faire passer son
approche comme scientifique, et c’est là que réside le danger[18]. »

En Belgique [modifier]
En Belgique, une enquête publiée en 2008 a montré que 20 % des Flamands croyaient en la théorie
du créationnisme[12]. Des résultats similaires avaient été trouvés en 2007 pour l'ensemble de la population
belge[19].

En Suisse [modifier]
En Suisse, on estime qu'en 2006, 28% des Suisses ne croyaient pas à l'évolution, taux parmi les
plus élevé en Europe occidentale [19],[20].
En France [modifier]
En raison de la tradition laïque de la France, on n'y trouve pas de chercheurs soutenant clairement le
dessein intelligent ou proposant des théories ouvertement appuyées sur une évolution guidée par une
puissance divine. En revanche, il existe différents courants que les biologistes considèrent comme relevant
de la même mouvance ou de la même démarche. Pour les biologistes les plus engagés, ces penseurs sont
même des néocréationnistes non avoués[21]. Guillaume Lecointre, ichtyologue, Professeur au Muséum
national d'histoire naturelle, parle de « créationnisme doux » pour qualifier ce mouvement[22]. La même
expression a été employée par le théologien dominicain Jacques Arnould lors d'une émission sur Canal
Académie portant sur les différentes formes de créationnismes en France et en Europe[23].
Jean Staune est l'un des principaux porte-parole d'une mouvance non-darwinienne ouvrant des
perspectives cohérentes avec les traditions monothéistes tout en réfutant le dessein intelligent[24][25].
L'hypothèse de l' Inside Story, une théorie du paléontologue français Jean Chaline qui reprend la découverte
des origines embryonnaires d'Homo par la paléontologue française Anne Dambricourt-Malassé, reste
l'exemple le plus marquant d'une controverse scientifique autour d'une théorie considérée comme
créationniste par la communauté scientifique. Chercheur au CNRS, A. Dambricourt-Malassé soumet un
article à La Recherche (publié en avril 1996[réf. nécessaire]) apportant un éclairage original aux troubles de
la posture et de l'occlusion. Les résultats exposés dans cet article seront controversés, car basé sur une
hypothèse évolutionniste ne faisant pas intervenir la sélection naturelle. En 2005, sa position est reprise par
la chaine de télévision Arte, dans le documentaire de Thomas Johnson, «Homo sapiens, une nouvelle
histoire», vivement critiqué par la communauté scientifique. Les partisans de l'hypothèse de l' Inside Story ont
parfois dénoncé la récupération de cette découverte par des formes de néocréationnismes comme la thèse
d'une programmation William Demsky[26].
Parodies [modifier]
Le pastafarisme est une parodie de religion[27] créée par Bobby Henderson, pour protester contre la
décision du Comité d'Éducation de l'État du Kansas, soutenue par le président des États-Unis George W.
Bush et le sénateur Bill Frist[28], de permettre au dessein intelligent d'être enseigné dans les cours de
science au même titre que la théorie de l'évolution.

Notes et références [modifier]

1. ↑ La traduction en français de design par dessein est devenue usuelle dans ce cadre. Une
traduction par conception serait plus exacte et permettrait de traduire intelligent designer par
concepteur intelligent plutôt que par cause intelligente
2. ↑ Voir site du Discovery Institute, un des promoteurs du dessein Intelligent, Questions About
Intelligent Design [archive]
3. ↑ (en) Intelligent Design as Science [archive] sur TalkOrigins. Consulté le 22 août 2010
4. ↑ (en) Intelligent Design as creationism [archive] sur TalkOrigins. Consulté le 22 août
2010
5. ↑ (fr) The Discovery Institute, « The Wedge [archive] », p. 4. Consulté le 22 janvier 2008
6. ↑ (fr) The Discovery Institute, « The “Wedge Document”: “So What?” [archive] », The
Discovery Institute. Consulté le 22 janvier 2008
7. ↑ (en) Propos du chef astronome du Vatican sur le dessein Intelligent [archive]
8. ↑ (en) More Than Intelligent Design de Hugh Ross [archive]
9. ↑ Ruling, Kitzmiller v. Dover Area School, Cas No. 04cv2688. 20 December 2005
10.↑ Le juge John E. Jones III menacé de mort par les tenants du dessein Intelligent [archive]
 11.↑ Ciel & espace, novembre 2006, « Le pape et la science » par Agnès Lenoire
12.↑ a et b (fr) Jean-Pierre Stroobants, « En Belgique, le combat contre le créationnisme
s'organise [archive] », 7 février 2008, Le Monde. Mis en ligne le 7 février 2008, consulté le
8 février 2008
13.↑ (fr) Marc Mennssier, « Offensive du créationnisme islamique en France [archive] »,
15 octobre 2007, Le Figaro. Mis en ligne le 15 octobre 2007, consulté le 8 février 2008
14.↑ Rachad Armanios, « La propagande créationniste débarque en Suisse romande », dans
Le Courrier du 28/03/2007, [lire en ligne [archive]]
15.↑ (fr) Harun Yahya, « L'Atlas de la création [archive] ». Consulté le 8 février 2008
16.↑ The "Intelligent Design" Distraction [archive], Harun Yahya, Harun Yahya International ©
2007
17.↑ mms://coenews.coe.int/vod/070625_w04_w.wmv (Conférence de presse de Guy
Lengagne au sujet de son rapport retiré)
18.↑ Assemblée parlementaire du Conseil de l’Europe, « Dangers du créationnisme dans
l’éducation [archive] » sur Assemblée parlementaire du Conseil de l’Europe [archive], 4 octobre 2007
19.↑ a et b Jon D. Miller, Eugenie C. Scott et Shinji Okamoto, « Public Acceptance of
Evolution », dans Science, vol. 313, no 5788, 2006, p. 765–766 [ texte intégral [archive], lien
DOI [archive] (pages consultées le 14 janvier 2009) ]
20.↑ Tribune de Genève, « Un Suisse sur trois renie ses origines animales [archive] », 5
octobre 2006. Consulté le 14 janvier 2009
21.↑ Le jeu de masques du néocréationnisme français [archive]
22.↑ Guillaume Lecointre répond à Jean Staune au sujet de l’UIP et du sens des actions à
mener contre elle [archive]
 23.↑ Emission de Radio Canal Académie du 27 janvier 2007 [archive]
24.↑ Les mécanismes de l'évolution [archive]
25.↑ Le matérialisme méthodologique [archive]
26.↑ Anne Dambricourt-Malassé déclare qu'« en tant que paléontologue, [elle] ne peut adhérer
à un tel mouvement de pensée »; L'os de la discorde, entretien avec Anne Dambricourt, Historia
d'août 2006.
27.↑ (fr) Guy Lengagne, « Les dangers du créationnisme dans l’éducation [archive] »,
8 juin 2007, Assemblée parlementaire du Conseil de l'Europe. Consulté le 1er août 2008
Le paragraphe #52 de l'exposé des motifs est consacré au Pastafarisme.
28.↑ (fr) Bush: Intelligent Design Should Be Taught [archive], 2 août 2005, Associated
Press. Consulté le 7 février 2008

Voir aussi [modifier]

Articles connexes [modifier]
• Pastafarisme
• Créationnisme
• Pseudoscience
• Relation entre science et religion
• Les dangers du créationnisme dans l'éducation
• Holism and evolution
• Néo-créationnisme
Bibliographie [modifier]
• Dialogues sur la religion naturelle, David Hume
• Cyrille Baudouin et Olivier Brosseau, Les Créationnismes, une menace pour la société
française ?, Éditions Syllepse, Paris, 2008 (ISBN 978-2-84950-16-72) [présentation en ligne]
• Rouvière, Jean-Marc (2006), Brèves méditations sur la création du monde, L'Harmattan.
• Gould, S. J. (2000), Et Dieu dit : Que Darwin soit ! : Science et religion, enfin la paix ?,
préface de Dominique Lecourt, Éditions du Seuil.
• Lecourt, D. (1992, 3e éd. « Quadrige » 2007), L'Amérique entre la Bible et Darwin : Suivi de
Intelligent design : science, morale et politique, PUF.
• Lecourt, D. (dir) (1999, 4e réed. « Quadrige » 2006), Dictionnaire d’histoire et philosophie
des sciences, PUF.

Liens externes [modifier]

• (fr) Du créationnisme au design intelligent, série d'articles (1999-2005) sur le site de
l'Agence Science-Presse
• (fr) Inquiétante offensive des créationnistes américains , sur le site de Le Monde.fr
• (fr) Évolution et créationnismes, sur le site du CNRS
• (fr) Évolution contre création en procès aux États-Unis, sur le site du journal Libération
• (fr) Jacques Debelmas, Créationnisme et Intelligent Design, 02/03/2009
• (fr) Science, religion, société, site d'analyses et de réflexions
 • (fr) À propos de la Genèse et de ses contradictions, voir les pages 18 à 25 du magazine
Allez savoir ! (N°43) de l'Université de Lausanne (UNIL)

• Portail des religions et croyances

Ce document provient de « http://fr.wikipedia.org/wiki/Dessein_intelligent ».

Catégories : Croyance | Pseudo-science | Créationnisme | [+]

 W000

Irreducible complexity
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This article covers irreducible complexity as used by those who argue for intelligent design.
For information on irreducible complexity as used in Systems Theory, see Irreducible complexity
(Emergence).
Part of a series of articles on

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Irreducible complexity (IC) is a nonscientific argument by proponents of intelligent design that certain
biological systems are too complex to have evolved from simpler, or "less complete" predecessors, through
natural selection acting upon a series of advantageous naturally-occurring, chance mutations.[1] The
argument is central to intelligent design, and is rejected by the scientific community,[2] which overwhelmingly
regards intelligent design as pseudoscience.[3] Irreducible complexity is one of two main arguments intended
to support intelligent design, the other being specified complexity.[4]
Biochemistry professor Michael Behe, the originator of the term irreducible complexity, defines an
irreducibly complex system as one "composed of several well-matched, interacting parts that contribute to
the basic function, wherein the removal of any one of the parts causes the system to effectively cease
functioning".[5] These examples are said to demonstrate that modern biological forms could not have
evolved naturally. Evolutionary biologists have shown that such systems can in fact evolve,[6] and Behe's
examples are considered to constitute an argument from ignorance.[7]
In the 2005 Kitzmiller v. Dover Area School District trial, Behe gave testimony on the subject of
irreducible complexity. The court found that "Professor Behe's claim for irreducible complexity has been
refuted in peer-reviewed research papers and has been rejected by the scientific community at large."[2]
Nonetheless, irreducible complexity continues to be cited as an important argument by creationists,
particularly intelligent design proponents.
Contents
[hide]
• 1 Definitions
• 2 History
• 2.1 Forerunners
• 2.1.1 Up to the 18th century
• 2.1.2 19th century
• 2.1.3 20th century
• 2.2 Origins
• 2.3 The mousetrap analogy
• 2.4 Consequences of irreducible complexity
• 3 Stated examples
• 3.1 Blood clotting cascade
• 3.2 Eye
• 3.3 Flagella
• 4 Response of the scientific community
• 4.1 Reducibility of "irreducible" systems
• 4.2 Gradual adaptation to new functions
• 4.3 Falsifiability and experimental evidence
• 4.4 Argument from ignorance
• 5 Irreducible complexity in the Dover trial
[edit] Definitions
The term "irreducible complexity" was coined by Behe, who defined it as applying to:
A single system which is composed of several interacting parts that contribute to the basic
function, and where the removal of any one of the parts causes the system to effectively cease
functioning. (Darwin's Black Box p39 in the 2006 edition)

Supporters of intelligent design use this term to refer to biological systems and organs that they
believe could not have come about by any series of small changes. They argue that anything less than the
complete form of such a system or organ would not work at all, or would in fact be a detriment to the
organism, and would therefore never survive the process of natural selection. Although they accept that
some complex systems and organs can be explained by evolution, they claim that organs and biological
features which are irreducibly complex cannot be explained by current models, and that an intelligent
designer must have created life or guided its evolution. Accordingly, the debate on irreducible complexity
concerns two questions: whether irreducible complexity can be found in nature, and what significance it
would have if it did exist in nature.
A second definition given by Behe (his "evolutionary definition") is as follows:
An irreducibly complex evolutionary pathway is one that contains one or more unselected steps
(that is, one or more necessary-but-unselected mutations). The degree of irreducible complexity
is the number of unselected steps in the pathway.

Intelligent design advocate William Dembski gives this definition:

 A system performing a given basic function is irreducibly complex if it includes a set of well-
matched, mutually interacting, nonarbitrarily individuated parts such that each part in the set is
indispensable to maintaining the system's basic, and therefore original, function. The set of
these indispensable parts is known as the irreducible core of the system.[8]

[edit] History
[edit] Forerunners
The argument from irreducible complexity is a descendant of the teleological argument for God (the
argument from design or from complexity). This states that because certain things in nature are very
complicated, they must have been designed. William Paley famously argued, in his 1802 watchmaker
analogy, that complexity in nature implies a God for the same reason that the existence of a watch implies
the existence of a watchmaker.[9] This argument has a long history, and can be traced back at least as far as
Cicero's De natura deorum ii.34.[10][11]

[edit] Up to the 18th century

Galen (1st and 2nd centuries AD) wrote about the large number of parts of the body and their
relationships, which observation was cited as evidence for creation.[12] The idea that specifically the
interdependence between parts would have implications for the origins of living things was raised by writers
starting with Pierre Gassendi in the mid 17th century[13] and John Wilkins, who wrote (citing Galen), "Now to
imagine, that all these things, according to their several kinds, could be brought into this regular frame and
order, to which such an infinite number of Intentions are required, without the contrivance of some wise
Agent, must needs be irrational in the highest degree."[14] In the late 17th century, Thomas Burnet referred
to "a multitude of pieces aptly joyn’d" to argue against the eternity of life.[15] In the early 18th century,
Nicolas Malebranche[16] wrote "An organized body contains an infinity of parts that mutually depend upon
one another in relation to particular ends, all of which must be actually formed in order to work as a whole,"
arguing in favor of preformation, rather than epigenesis, of the individual;[17] and a similar argument about
the origins of the individual was made by other 18th century students of natural history.[18] In his 1790 book,
The Critique of Judgment, Kant argues that "we cannot conceive how a whole that comes into being only
gradually from its parts can nevertheless be the cause of the properties of those parts"[19]

[edit] 19th century

As we transition to the 19th century, we find references which relate to evolution.
Chapter XV of Paley's Natural Theology discusses at length what he called "relations" of parts of
living things as an indication of their design.[9]
Georges Cuvier applied his principle of the correlation of parts to describe an animal from
fragmentary remains. For Cuvier, this was related to another principle of his, the conditions of existence,
which excluded the possibility of transmutation of species.[20]
While he did not originate the term, Charles Darwin identified the argument as a possible way to
falsify a prediction of the theory of evolution at the outset. In The Origin of Species, he wrote, "If it could be
demonstrated that any complex organ existed, which could not possibly have been formed by numerous,
successive, slight modifications, my theory would absolutely break down. But I can find out no such
case."[21] Darwin's theory of evolution challenges the teleological argument by postulating an alternative
explanation to that of an intelligent designer—namely, evolution by natural selection. By showing how simple
unintelligent forces can ratchet up designs of extraordinary complexity without invoking outside design,
Darwin showed that an intelligent designer was not the necessary conclusion to draw from complexity in
nature. The argument from irreducible complexity attempts to demonstrate that certain biological features
cannot be purely the product of Darwinian evolution.
In the late 19th century, in a dispute between supporters of the adequacy of natural selection and
those who held for inheritance of acquired characters, one of the arguments made repeatedly by Herbert
Spencer, and followed by others, depended on what Spencer referred to as co-adaptation of co-operative
parts, as in: "We come now to Professor Weismann's endeavour to disprove my second thesis — that it is
impossible to explain by natural selection alone the co-adaptation of co-operative parts. It is thirty years since
this was set forth in "The Principles of Biology." In §166, I instanced the enormous horns of the extinct Irish
elk, and contended that in this and in kindred cases, where for the efficient use of some one enlarged part
many other parts have to be simultaneously enlarged, it is out of the question to suppose that they can have
all spontaneously varied in the required proportions."[22][23] The history of this concept in the dispute has
been characterized: "An older and more religious tradition of idealist thinkers were committed to the
explanation of complex adaptive contrivances by intelligent design. ... Another line of thinkers, unified by the
recurrent publications of Herbert Spencer, also saw coadaptation as a composed, irreducible whole, but
sought to explain it by the inheritance of acquired characteristics."[24]

[edit] 20th century

Hermann Muller, in the early 20th century, discussed a concept similar to irreducible complexity.
However, far from seeing this as a problem for evolution, he described the "interlocking" of biological features
as a consequence to be expected of evolution, which would lead to irreversibility of some evolutionary
changes.[25] He wrote, "Being thus finally woven, as it were, into the most intimate fabric of the organism,
the once novel character can no longer be withdrawn with impunity, and may have become vitally
necessary."[26]
In 1974, Young Earth Creationist Henry M. Morris introduced a similar concept in his book Scientific
Creationism in which he wrote; "This issue can actually be attacked quantitatively, using simple principles of
mathematical probability. The problem is simply whether a complex system, in which many components
function unitedly together, and in which each component is uniquely necessary to the efficient functioning of
the whole, could ever arise by random processes."[27]
A book-length study of a concept similar to irreducible complexity, explained by gradual, step-wise,
non-teleological evolution, was published in 1975 by Thomas H. Frazzetta. "A complex adaptation is one
constructed of several components that must blend together operationally to make the adaptation "work". It is
analogous to a machine whose performance depends upon careful cooperation among its parts. In the case
of the machine, no single part can greatly be altered without changing the performance of the entire
machine." The machine that he chose as an analog is the Peaucellier machine, and one biological system
given extended description was the jaw apparatus of a python. The conclusion of this investigation, rather
than that evolution of a complex adaptation was impossible, "awed by the adaptations of living things, to be
stunned by their complexity and suitability", was "to accept the inescapable but not humiliating fact that much
of mankind can be seen in a tree or a lizard."[28]
In 1981, Ariel Roth, in defense of the creation science position in the trial McLean v. Arkansas, said
of "complex integrated structures" that "This system would not be functional until all the parts were there ...
How did these parts survive during evolution ...?"[29]
In 1985 Cairns-Smith wrote of "interlocking", "How can a complex collaboration between components
evolve in small steps?" and used the analogy of the scaffolding called centering used to build an arch then
removed afterwards: "Surely there was 'scaffolding'. Before the multitudinous components of present
biochemistry could come to lean together they had to lean on something else."[30] However, neither Muller or
Cairns-Smith claimed that their ideas were evidence of something supernatural.[31]
An essay in support of creationism published in 1994 referred to bacterial flagella as showing
"multiple, integrated components", where "nothing about them works unless every one of their complexly
fashioned and integrated components are in place" and asked the reader to "imagine the effects of natural
selection on those organisms that fortuitously evolved the flagella ... without the concommitant [ sic] control
mechanisms".[32][33]
An early concept of irreducibly complex systems comes from Ludwig von Bertalanffy, a 20th-century
Austrian biologist.[34] He believed that complex systems must be examined as complete, irreducible systems
in order to fully understand how they work. He extended his work on biological complexity into a general
theory of systems in a book titled General Systems Theory. After James Watson and Francis Crick published
the structure of DNA in the early 1950s, General Systems Theory lost many of its adherents in the physical
and biological sciences.[35] However, Systems theory remained popular in the social sciences long after its
demise in the physical and biological sciences.
[edit] Origins

Michael Behe's controversial book Darwin's Black Box popularized the concept of irreducible
complexity.
Michael Behe developed his ideas on the concept around 1992, in the early days of the 'wedge
movement', and first presented his ideas about "irreducible complexity" in June 1993 when the "Johnson-
Behe cadre of scholars" met at Pajaro Dunes in California.[36] He set out his ideas in the second edition of
Of Pandas and People published in 1993, extensively revising Chapter 6 Biochemical Similarities with new
sections on the complex mechanism of blood clotting and on the origin of proteins.[37]
He first used the term "irreducible complexity" in his 1996 book Darwin's Black Box, to refer to certain
complex biochemical cellular systems. He posits that evolutionary mechanisms cannot explain the
development of such "irreducibly complex" systems. Notably, Behe credits philosopher William Paley for the
original concept, not von Bertalanffy, and suggests that his application of the concept to biological systems is
entirely original. Intelligent design advocates argue that irreducibly complex systems must have been
deliberately engineered by some form of intelligence.
In 2001, Michael Behe wrote: "[T]here is an asymmetry between my current definition of irreducible
complexity and the task facing natural selection. I hope to repair this defect in future work." Behe specifically
explained that the "current definition puts the focus on removing a part from an already functioning system",
but the "difficult task facing Darwinian evolution, however, would not be to remove parts from sophisticated
pre-existing systems; it would be to bring together components to make a new system in the first place".[38]
In the 2005 Kitzmiller v. Dover Area School District trial, Behe testified under oath that he "did not judge [the
asymmetry] serious enough to [have revised the book] yet."[39]
Behe additionally testified that the presence of irreducible complexity in organisms would not rule out
the involvement of evolutionary mechanisms in the development of organic life. He further testified that he
knew of no earlier "peer reviewed articles in scientific journals discussing the intelligent design of the blood
clotting cascade," but that there were "probably a large number of peer reviewed articles in science journals
that demonstrate that the blood clotting system is indeed a purposeful arrangement of parts of great
complexity and sophistication."[40] (The judge ruled that "intelligent design is not science and is essentially
religious in nature".)[41]
 According to the theory of evolution, genetic variations occur without specific design or intent. The
environment "selects" the variants that have the highest fitness, which are then passed on to the next
generation of organisms. Change occurs by the gradual operation of natural forces over time, perhaps
slowly, perhaps more quickly (see punctuated equilibrium). This process is able to adapt complex structures
from simpler beginnings, or convert complex structures from one function to another (see spandrel). Most
intelligent design advocates accept that evolution occurs through mutation and natural selection at the "micro
level", such as changing the relative frequency of various beak lengths in finches, but assert that it cannot
account for irreducible complexity, because none of the parts of an irreducible system would be functional or
advantageous until the entire system is in place.

[edit] The mousetrap analogy

Michael Behe believes that many aspects of life show evidence of design, using the mousetrap in an
analogy disputed by others.[42]
 Behe uses the mousetrap as an illustrative example of this concept. A mousetrap consists of five
interacting pieces—the base, the catch, the spring, the hammer and the hold-down bar. All of these must be in
place for the mousetrap to work, as the removal of any one piece destroys the function of the mousetrap.
Likewise, he asserts that biological systems require multiple parts working together in order to function.
Intelligent design advocates claim that natural selection could not create from scratch those systems for
which science is currently unable to find a viable evolutionary pathway of successive, slight modifications,
because the selectable function is only present when all parts are assembled.
In his 2008 book Only A Theory, biologist Kenneth R. Miller challenges Behe's claim that the
mousetrap is irreducibly complex. Miller observes that various subsets of the five components can be
devised to form cooperative units, ones that have different functions from the mousetrap and so, in biological
terms, could form functional spandrels before being adapted to the new function of catching mice. In an
example taken from his high school experience, Miller recalls that one of his classmates
...struck upon the brilliant idea of using an old, broken mousetrap as a spitball catapult, and it
worked brilliantly....It had worked perfectly as something other than a mousetrap....my rowdy
friend had pulled a couple of parts --probably the hold-down bar and catch-- off the trap to make
it easier to conceal and more effective as a catapult...[leaving] the base, the spring, and the
hammer. Not much of a mousetrap, but a helluva spitball launcher....I realized why [Behe's]
mousetrap analogy had bothered me. It was wrong. The mousetrap is not irreducibly complex
after all.[43]

Other systems identified by Miller that include mousetrap components include the following:[43]
• use the spitball launcher as a tie clip (same three-part system with different function)
 • remove the spring from the spitball launcher/tie clip to create a two-part key chain (base +
hammer)
• glue the spitball launcher/tie clip to a sheet of wood to create a clipboard (launcher + glue +
wood)
• remove the hold-down bar for use as a toothpick (single element system)
Behe has responded, "What Miller actually means is that if you take away some components and
then go on to, say, twist a couple of metal pieces in just the right way and add a few staples in the correct
positions, you can construct a new kind of working trap, which may superficially resemble the starting trap.
That, however, is intelligent design. Neither Miller nor anyone else has shown that the mousetrap I pictured in
my book can be constructed by a series of small changes, one at a time, as Darwinian evolution would have
to do." [44]

[edit] Consequences of irreducible complexity

Behe's original examples of irreducibly complex mechanisms included the bacterial flagellum of E.
coli, the blood clotting cascade, cilia, and the adaptive immune system.
Behe argues that organs and biological features which are irreducibly complex cannot be wholly
explained by current models of evolution. In explicating his definition of "irreducible complexity" he notes that:
An irreducibly complex system cannot be produced directly (that is, by continuously improving
the initial function, which continues to work by the same mechanism) by slight, successive
modifications of a precursor system, because any precursor to an irreducibly complex system
that is missing a part is by definition nonfunctional.
 Irreducible complexity is not an argument that evolution does not occur, but rather an argument that it
is "incomplete". In the last chapter of Darwin's Black Box, Behe goes on to explain his view that irreducible
complexity is evidence for intelligent design. Mainstream critics, however, argue that irreducible complexity,
as defined by Behe, can be generated by known evolutionary mechanisms. Behe's claim that no scientific
literature adequately modeled the origins of biochemical systems through evolutionary mechanisms has
been challenged by TalkOrigins.[45][46] The judge in the Dover trial wrote "By defining irreducible complexity
in the way that he has, Professor Behe attempts to exclude the phenomenon of exaptation by definitional fiat,
ignoring as he does so abundant evidence which refutes his argument. Notably, the NAS has rejected
Professor Behe’s claim for irreducible complexity..."[47]

[edit] Stated examples

Behe and others have suggested a number of biological features that they believe may be irreducibly
complex.

[edit] Blood clotting cascade

The blood clotting or coagulation cascade in vertebrates is a complex biological pathway which is
given as an example of apparent irreducible complexity.[48]
The irreducible complexity argument assumes that the necessary parts of a system have always
been necessary, and therefore could not have been added sequentially. However, in evolution, natural
selection can lead to complex biochemical systems being built up from simpler systems, or to existing
functional systems being recombined as a new system with a different function.[47] For example, one of the
clotting factors that Behe listed as a part of the clotting cascade was later found to be absent in whales,
demonstrating that it is not essential for a clotting system.[49] Many purportedly irreducible structures can be
found in other organisms as much simpler systems that utilize fewer parts. These systems, in turn, may have
had even simpler precursors that are now extinct. Behe has responded to critics of his clotting cascade
arguments by suggesting that homology is evidence for evolution, but not for natural selection.[50]
The "improbability argument" also misrepresents natural selection. It is correct to say that a set of
simultaneous mutations that form a complex protein structure is so unlikely as to be unfeasible, but that is not
what Darwin advocated. His explanation is based on small accumulated changes that take place without a
final goal. Each step must be advantageous in its own right, although biologists may not yet understand the
reason behind all of them—for example, jawless fish accomplish blood clotting with just six proteins instead of
the full 10.[51]

[edit] Eye
Main article: Evolution of the eye
 Often used as an example of irreducible complexity.
(a) A pigment spot
(b) A simple pigment cup
(c) The simple optic cup found in abalone
(d) The complex lensed eye of the marine snail and the octopus
The eye is a famous example of a supposedly irreducibly complex structure, due to its many
elaborate and interlocking parts, seemingly all dependent upon one another. It is frequently cited by
intelligent design and creationism advocates as an example of irreducible complexity. Behe used the
"development of the eye problem" as evidence for intelligent design in Darwin's Black Box. Although Behe
acknowledged that the evolution of the larger anatomical features of the eye have been well-explained, he
claimed that the complexity of the minute biochemical reactions required at a molecular level for light
sensitivity still defies explanation. Creationist Jonathan Sarfati has described the eye as evolutionary
biologists' "greatest challenge as an example of superb 'irreducible complexity' in God's creation", specifically
pointing to the supposed "vast complexity" required for transparency.[52]
In an often misquoted[53] passage from On the Origin of Species, Charles Darwin appears to
acknowledge the eye's development as a difficulty for his theory. However, the quote in context shows that
Darwin actually had a very good understanding of the evolution of the eye. He notes that "to suppose that the
eye ... could have been formed by natural selection, seems, I freely confess, absurd in the highest possible
degree". Yet this observation was merely a rhetorical device for Darwin. He goes on to explain that if gradual
evolution of the eye could be shown to be possible, "the difficulty of believing that a perfect and complex eye
could be formed by natural selection ... can hardly be considered real". He then proceeded to roughly map
out a likely course for evolution using examples of gradually more complex eyes of various species.[54]
 The eyes of vertebrates (left) and invertebrates such as the octopus (right) developed independently:
vertebrates evolved an inverted retina with a blind spot over their optic disc, whereas octopuses avoided this
with a non-inverted retina.
Since Darwin's day, the eye's ancestry has become much better understood. Although learning about
the construction of ancient eyes through fossil evidence is problematic due to the soft tissues leaving no
imprint or remains, genetic and comparative anatomical evidence has increasingly supported the idea of a
common ancestry for all eyes.[55][56][57]
Current evidence does suggest possible evolutionary lineages for the origins of the anatomical
features of the eye. One likely chain of development is that the eyes originated as simple patches of
photoreceptor cells that could detect the presence or absence of light, but not its direction. When, via random
mutation across the population, the photosensitive cells happened to have developed on a small depression,
it endowed the organism with a better sense of the light's source. This small change gave the organism an
advantage over those without the mutation. This genetic trait would then be "selected for" as those with the
trait would have an increased chance of survival, and therefore progeny, over those without the trait.
Individuals with deeper depressions would be able to discern changes in light over a wider field than those
individuals with shallower depressions. As ever deeper depressions were advantageous to the organism,
gradually, this depression would become a pit into which light would strike certain cells depending on its
angle. The organism slowly gained increasingly precise visual information. And again, this gradual process
continued as individuals having a slightly shrunken aperture of the eye had an advantage over those without
the mutation as an aperture increases how collimated the light is at any one specific group of photoreceptors.
As this trait developed, the eye became effectively a pinhole camera which allowed the organism to dimly
make out shapes—the nautilus is a modern example of an animal with such an eye. Finally, via this same
selection process, a protective layer of transparent cells over the aperture was differentiated into a crude
lens, and the interior of the eye was filled with humours to assist in focusing images.[58][59][60] In this way,
eyes are recognized by modern biologists as actually a relatively unambiguous and simple structure to
evolve, and many of the major developments of the eye's evolution are believed to have taken place over
only a few million years, during the Cambrian explosion.[61]
Behe maintains that the complexity of light sensitivity at the molecular level and the minute
biochemical reactions required for those first "simple patches of photoreceptor[s]" still defies explanation.
Other intelligent design proponents have pointed to the difficulty of the entire visual system evolving rather
than the eye alone.[62]

[edit] Flagella
Main article: Evolution of flagella
 The bacterial flagellum is frequently invoked as an example of irreducible complexity.
The flagella of certain bacteria constitute a molecular motor requiring the interaction of about 40
complex protein parts. Behe asserts that the absence of any one of these proteins causes the flagella to fail
to function, and that the flagellum "engine" is irreducibly complex as in his view if we try to reduce its
complexity by positing an earlier and simpler stage of its evolutionary development, we get an organism
which functions improperly.
Scientists regard this argument as having been disproved in the light of research dating back to 1996
as well as more recent findings.[63] They point out that the basal body of the flagella has been found to be
similar to the Type III secretion system (TTSS), a needle-like structure that pathogenic germs such as
Salmonella and Yersinia pestis use to inject toxins into living eucaryote cells. The needle's base has ten
elements in common with the flagellum, but it is missing forty of the proteins that make a flagellum work.[64]
Thus, this system negates the claim that taking away any of the flagellum's parts would render it useless. On
this basis, Kenneth Miller notes that, "The parts of this supposedly irreducibly complex system actually have
functions of their own."[65][66]

[edit] Response of the scientific community

Like intelligent design, the concept it seeks to support, irreducible complexity has failed to gain any
notable acceptance within the scientific community. One science writer called it a "full-blown intellectual
surrender strategy."[67]

[edit] Reducibility of "irreducible" systems

Potentially viable evolutionary pathways have been proposed for allegedly irreducibly complex
systems such as blood clotting, the immune system[68] and the flagellum,[69][70] which were the three
examples Behe used. Even his example of a mousetrap was shown to be reducible by John H. McDonald.
[42] If irreducible complexity is an insurmountable obstacle to evolution, it should not be possible to conceive
of such pathways.[71]
Niall Shanks and Karl H. Joplin, both of East Tennessee State University, have shown that systems
satisfying Behe's characterization of irreducible biochemical complexity can arise naturally and
spontaneously as the result of self-organizing chemical processes.[72] They also assert that what evolved
biochemical and molecular systems actually exhibit is "redundant complexity"—a kind of complexity that is the
product of an evolved biochemical process. They claim that Behe overestimated the significance of
irreducible complexity because of his simple, linear view of biochemical reactions, resulting in his taking
snapshots of selective features of biological systems, structures and processes, while ignoring the redundant
complexity of the context in which those features are naturally embedded. They also criticized his over-
reliance of overly simplistic metaphors, such as his mousetrap. In addition, research published in the peer-
reviewed journal Nature has shown that computer simulations of evolution demonstrate that it is possible for
irreducible complexity to evolve naturally.[73]
It is illustrative to compare a mousetrap with a cat, in this context. Both normally function so as to
control the mouse population. The cat has many parts that can be removed leaving it still functional; for
example, its tail can be bobbed, or it can lose an ear in a fight. Comparing the cat and the mousetrap, then,
one sees that the mousetrap (which is not alive) offers better evidence, in terms of irreducible complexity, for
intelligent design than the cat. Even looking at the mousetrap analogy, several critics have described ways in
which the parts of the mousetrap could have independent uses or could develop in stages, demonstrating
that it is not irreducibly complex.[42][43]
Moreover, even cases where removing a certain component in an organic system will cause the
system to fail do not demonstrate that the system couldn't have been formed in a step-by-step, evolutionary
process. By analogy, stone arches are irreducibly complex—if you remove any stone the arch will collapse—
yet we build them easily enough, one stone at a time, by building over centering that is removed afterward.
Similarly, naturally occurring arches of stone are formed by weathering away bits of stone from a large
concretion that has formed previously. Evolution can act to simplify as well as to complicate. This raises the
possibility that seemingly irreducibly complex biological features may have been achieved with a period of
increasing complexity, followed by a period of simplification.
In April 2006 a team led by Joe Thornton, assistant professor of biology at the University of Oregon's
Center for Ecology and Evolutionary Biology, using techniques for resurrecting ancient genes, scientists for
the first time reconstructed the evolution of an apparently irreducibly complex molecular system. The
research was published in the April 7 issue of Science.[6][74]
It may be that irreducible complexity does not actually exist in nature, and that the examples given by
Behe and others are not in fact irreducibly complex, but can be explained in terms of simpler precursors.
There has also been a theory that challenges irreducible complexity called facilitated variation. The theory
has been presented in 2005 by Marc W. Kirschner, a professor and chair of Department of Systems Biology
at Harvard Medical School, and John C. Gerhart, a professor in Molecular and Cell Biology, University of
California, Berkeley. In their theory, they describe how certain mutation and changes can cause apparent
irreducible complexity. Thus, seemingly irreducibly complex structures are merely "very complex", or they are
simply misunderstood or misrepresented.

[edit] Gradual adaptation to new functions

Main article: Exaptation
The precursors of complex systems, when they are not useful in themselves, may be useful to
perform other, unrelated functions. Evolutionary biologists argue that evolution often works in this kind of
blind, haphazard manner in which the function of an early form is not necessarily the same as the function of
the later form. The term used for this process is "exaptation". The mammalian middle ear (derived from a
jawbone) and the panda's thumb (derived from a wrist bone spur) are considered classic examples. A 2006
article in Nature demonstrates intermediate states leading toward the development of the ear in a Devonian
fish (about 360 million years ago).[75] Furthermore, recent research shows that viruses play a heretofore
unexpectedly great role in evolution by mixing and matching genes from various hosts.[ citation needed]
 Arguments for irreducibility often assume that things started out the same way they ended up—as we
see them now. However, that may not necessarily be the case. In the Dover trial an expert witness for the
plaintiffs, Ken Miller, demonstrated this possibility using Behe's mousetrap analogy. By removing several
parts, Miller made the object unusable as a mousetrap, but he pointed out that it was now a perfectly
functional, if unstylish, tie clip.[43][76]

[edit] Falsifiability and experimental evidence

Some critics, such as Jerry Coyne (professor of evolutionary biology at the University of Chicago)
and Eugenie Scott (a physical anthropologist and executive director of the National Center for Science
Education) have argued that the concept of irreducible complexity, and more generally, the theory of
intelligent design is not falsifiable, and therefore, not scientific.
Behe argues that the theory that irreducibly complex systems could not have been evolved can be
falsified by an experiment where such systems are evolved. For example, he posits taking bacteria with no
flagellum and imposing a selective pressure for mobility. If, after a few thousand generations, the bacteria
evolved the bacterial flagellum, then Behe believes that this would refute his theory.[ citation needed]
Other critics take a different approach, pointing to experimental evidence that they believe falsifies
the argument for Intelligent Design from irreducible complexity. For example, Kenneth Miller cites the lab
work of Barry G. Hall on E. coli, which he asserts is evidence that "Behe is wrong."[77]
Other evidence that irreducible complexity is not a problem for evolution comes from the field of
computer science, where computer analogues of the processes of evolution are routinely used to
automatically design complex solutions to problems. The results of such Genetic Algorithms are frequently
irreducibly complex since the process, like evolution, both removes non-essential components over time as
well as adding new components. The removal of unused components with no essential function, like the
natural process where rock underneath a natural arch is removed, can produce irreducibly complex
structures without requiring the intervention of a designer. Researchers applying these algorithms are
automatically producing human competitive designs—but no human designer is required.[78]

[edit] Argument from ignorance

Intelligent design proponents attribute to an intelligent designer those biological structures they
believe are irreducibly complex and where they say a natural explanation is insufficient to account for them.
[79] However, critics view irreducible complexity as a special case of the "complexity indicates design" claim,
and thus see it as an argument from ignorance and God of the gaps argument.[7]
Eugenie Scott, along with Glenn Branch and other critics, has argued that many points raised by
intelligent design proponents are arguments from ignorance.[80] Behe has been accused of using an
"argument by lack of imagination", and Behe himself acknowledges that a failure of current science to explain
how an "irreducibly complex" organism did or could evolve does not automatically prove the impossibility of
such an evolution.
Irreducible complexity is at its core an argument against evolution. If truly irreducible systems are
found, the argument goes, then intelligent design must be the correct explanation for their existence.
However, this conclusion is based on the assumption that current evolutionary theory and intelligent design
are the only two valid models to explain life, a false dilemma.[81][82]
[edit] Irreducible complexity in the Dover trial
While testifying at the Kitzmiller v. Dover Area School District trial Behe conceded that there are no
peer-reviewed papers supporting his claims that complex molecular systems, like the bacterial flagellum, the
blood-clotting cascade, and the immune system, were intelligently designed nor are there any peer-reviewed
articles supporting his argument that certain complex molecular structures are "irreducibly complex."[83]
In the final ruling of Kitzmiller v. Dover Area School District, Judge Jones specifically singled out
Behe and irreducible complexity:[83]
• "Professor Behe admitted in "Reply to My Critics" that there was a defect in his view of
irreducible complexity because, while it purports to be a challenge to natural selection, it does not
actually address "the task facing natural selection." and that "Professor Behe wrote that he hoped to
"repair this defect in future work..." (Page 73)
• "As expert testimony revealed, the qualification on what is meant by "irreducible complexity"
renders it meaningless as a criticism of evolution. (3:40 (Miller)). In fact, the theory of evolution
proffers exaptation as a well-recognized, well-documented explanation for how systems with multiple
parts could have evolved through natural means." (Page 74)
• "By defining irreducible complexity in the way that he has, Professor Behe attempts to
exclude the phenomenon of exaptation by definitional fiat, ignoring as he does so abundant evidence
which refutes his argument. Notably, the NAS has rejected Professor Behe’s claim for irreducible
complexity..." (Page 75)
• "As irreducible complexity is only a negative argument against evolution, it is refutable and
accordingly testable, unlike ID [Intelligent Design], by showing that there are intermediate structures
with selectable functions that could have evolved into the allegedly irreducibly complex systems.
(2:15-16 (Miller)). Importantly, however, the fact that the negative argument of irreducible complexity
is testable does not make testable the argument for ID. (2:15 (Miller); 5:39 (Pennock)). Professor
Behe has applied the concept of irreducible complexity to only a few select systems: (1) the bacterial
flagellum; (2) the blood-clotting cascade; and (3) the immune system. Contrary to Professor Behe’s
assertions with respect to these few biochemical systems among the myriad existing in nature,
however, Dr. Miller presented evidence, based upon peer-reviewed studies, that they are not in fact
irreducibly complex." (Page 76)
• "...on cross-examination, Professor Behe was questioned concerning his 1996 claim that
science would never find an evolutionary explanation for the immune system. He was presented with
fifty-eight peer-reviewed publications, nine books, and several immunology textbook chapters about
the evolution of the immune system; however, he simply insisted that this was still not sufficient
evidence of evolution, and that it was not "good enough." (23:19 (Behe))." (Page 78)
• "We therefore find that Professor Behe’s claim for irreducible complexity has been refuted in
peer-reviewed research papers and has been rejected by the scientific community at large. (17:45-46
(Padian); 3:99 (Miller)). Additionally, even if irreducible complexity had not been rejected, it still does
not support ID as it is merely a test for evolution, not design. (2:15, 2:35-40 (Miller); 28:63-66
(Fuller)). We will now consider the purportedly “positive argument” for design encompassed in the
phrase used numerous times by Professors Behe and Minnich throughout their expert testimony,
which is the “purposeful arrangement of parts.” Professor Behe summarized the argument as follows:
We infer design when we see parts that appear to be arranged for a purpose. The strength of the
inference is quantitative; the more parts that are arranged, the more intricately they interact, the
stronger is our confidence in design. The appearance of design in aspects of biology is
overwhelming. Since nothing other than an intelligent cause has been demonstrated to be able to
yield such a strong appearance of design, Darwinian claims notwithstanding, the conclusion that the
design seen in life is real design is rationally justified. (18:90-91, 18:109-10 (Behe); 37:50 (Minnich)).
As previously indicated, this argument is merely a restatement of the Reverend William Paley’s
argument applied at the cell level. Minnich, Behe, and Paley reach the same conclusion, that
complex organisms must have been designed using the same reasoning, except that Professors
Behe and Minnich refuse to identify the designer, whereas Paley inferred from the presence of
design that it was God. (1:6- 7 (Miller); 38:44, 57 (Minnich)). Expert testimony revealed that this
inductive argument is not scientific and as admitted by Professor Behe, can never be ruled out. (2:40
(Miller); 22:101 (Behe); 3:99 (Miller))." (Pages 79–80)

[edit] Notes and references

1. ^ Forrest, Barbara (May,2007) (PDF). Understanding the Intelligent Design Creationist
Movement: Its True Nature and Goals. A Position Paper from the Center for Inquiry, Office of Public
Policy. Washington, D.C.: Center for Inquiry, Inc..
http://www.centerforinquiry.net/uploads/attachments/intelligent-design.pdf. Retrieved 2007-08-22. .
2. ^ a b "We therefore find that Professor Behe’s claim for irreducible complexity has been
refuted in peer-reviewed research papers and has been rejected by the scientific community at
large." Ruling, Judge John E. Jones III, Kitzmiller v. Dover Area School District
3. ^ "True in this latest creationist variant, advocates of so-called intelligent design ... use more
slick, pseudoscientific language. They talk about things like 'irreducible complexity'" —
Shulman, Seth (2006). Undermining science: suppression and distortion in the Bush Administration .
Berkeley: University of California Press. p. 13. ISBN 0-520-24702-7. "for most members of the
mainstream scientific community, ID is not a scientific theory, but a creationist pseudoscience."
David Mu (Fall 2005). "Trojan Horse or Legitimate Science: Deconstructing the Debate over
Intelligent Design". Harvard Science Review 19 (1).
http://www.hcs.harvard.edu/~hsr/fall2005/mu.pdf.
Perakh M (2005 Summer). "Why Intelligent Design Isn't Intelligent — Review of: Unintelligent Design".
Cell Biol Educ. 4 (2): 121–2. doi:10.1187/cbe.05-02-0071.
Mark D. Decker. College of Biological Sciences, General Biology Program, University of Minnesota
Frequently Asked Questions About the Texas Science Textbook Adoption Controversy "The
Discovery Institute and ID proponents have a number of goals that they hope to achieve using
disingenuous and mendacious methods of marketing, publicity, and political persuasion. They do not
practice real science because that takes too long, but mainly because this method requires that one
have actual evidence and logical reasons for one's conclusions, and the ID proponents just don't
have those. If they had such resources, they would use them, and not the disreputable methods they
actually use."
4. ^ Ker Than (September 23, 2005). "Why scientists dismiss 'intelligent design' - LiveScience".
msnbc.com. http://www.msnbc.msn.com/id/9452500/ns/technology_and_science-science/. Retrieved
2010-05-17.
5. ^ Darwin's Black Box: The Biochemical Challenge to Evolution , Michael Behe, 1996, quoted
in Irreducible Complexity and Michael Behe (retrieved 8 January 2006)
6. ^ a b Bridgham JT, Carroll SM, Thornton JW (April 2006). "Evolution of hormone-receptor
complexity by molecular exploitation". Science 312 (5770): 97–101. doi:10.1126/science.1123348.
PMID 16601189.
7. ^ a b Index to Creationist Claims. Mark Isaak. The Talk.Origins Archive. "Irreducible
complexity and complex specified information are special cases of the "complexity indicates design"
claim; they are also arguments from incredulity." [1] "The argument from incredulity creates a god of
the gaps." [2]
 8. ^ No Free Lunch: Why Specified Complexity Cannot Be Purchased without Intelligence. by
William Dembski pp. 285
9. ^ a b William Paley:Natural Theology; or, Evidences of the Existence and Attributes of the
Deity. Collected from the Appearances of Nature 12th edition, 1809
10.^ On the Nature of the Gods, translated by Francis Brooks, London: Methuen, 1896.
11.^ See Henry Hallam's Introduction to the Literature of Europe part iii chapter iii paragraph 26
footnote u
12.^ De Formatione Foetus=The Construction of the Embryo, chapter 11 in Galen: Selected
Works, translated by P. N. Singer, The World's Classics, Oxford, Oxford University Press, 1997 ISBN
978-019-282450-9. One 18th-century reference to Galen is David Hume Dialogues Concerning
Natural Religion, 1779, Part 12, § 3, page 215. Also see Galen's De Usu Partium=On the Usefulness
of the Parts of the Body, translated and edited by Margaret Tallmadge May, Ithaca, New York,
Cornell University Press, 1968, especially book XVII. For a relevant discussion of Galen and other
ancients see pages 121-122, Goodman, Lenn Evan (2010). Creation and evolution. Milton Park,
Abingdon, Oxon and New York: Routledge. ISBN 978-0-415-91380-5.
13.^ De Generatione Animalium, chapter III. Partial translation in: Howard B. Adelmann,
Marcello Malpighi and the Evolution of Embryology Ithaca, New York, Cornell University Press, 1966,
volume 2, pages 811-812.
14.^ John Wilkins,Of the Principles and Duties of Natural Religion , London, 1675, book I,
chapter 6, page 82.
15.^ The Sacred Theory of the Earth, 2nd edition, London: Walter Kettilby, 1691. Book I Chapter
IV page 43
 16.^ De la recherche de la verité 6.2.4, 6th edition, 1712. English translation The Search after
Truth, translated and edited by Thomas M. Lennon and Paul J. Olscamp, Cambridge University
Press, 1997, ISBN 0-521-58004-8. Second paragraph from the end of the chapter, on page 465.
17.^ Pages 202-203 of Andrew Pyle (2006). "Malebranche on Animal Generation: Preexistence
and the Microscope". in Smith JH. The problem of animal generation in early modern philosophy .
Cambridge, UK: Cambridge University Press. pp. 194–214. ISBN 0-521-84077-5.
18.^ The Chicken or the Egg
19.^ This is Guyer's exposition on page 22 of Paul Guyer (1992). "Introduction". in Paul Guyer.
The Cambridge Companion to Kant. Cambridge: Cambridge University Press. pp. 1–25. ISBN 978-0-
521-36768-4. . Guyer goes on to add this parenthetical comment: "(here is where the theory of
natural selection removes the difficulty)". See Kant's discussion in section IX of the "First
Introduction" to the Critique of Judgment and in §§61, 64 (where he uses the expression
wechselsweise abhängt="reciprocally dependent"), and 66 of "Part Two, First Division".
20.^ See especially chapters VI and VII of William Coleman (1964). Georges Cuvier, Zoologist:
A Study in the History of Evolution Theory. Cambridge, Massachusetts: Harvard University Press.
See also the discussion of these principles in the Wikipedia article on Cuvier.
21.^ Darwin, Charles (1859). On the Origin of Species. London: John Murray. page 189,
Chapter VI
22.^ Page 594 in: Herbert Spencer (October 1894). "Weismannism Once More". The
Contemporary Review 66: 592–608. Another essay of his treating this concept is: Herbert Spencer
(1893). "The Inadequacy of "Natural Selection"". The Contemporary Review 63: 153–166. (Part I:
February) and pages 439-456 (Part II: March). These essays were reprinted in Herbert Spencer
(1891). The Works of Herbert Spencer. 17. London: Williams and Norgate. (also Osnabrück: Otto
Zeller, 1967). See also part III, Chapter XII, §166, pages 449-457 in: Herbert Spencer (1864).
Principles of Biology. I. London: Williams and Norgate. And: Herbert Spencer (1886). "The Factors
of Organic Evolution". Nineteenth Century 19: 570–589. (Part I: April) and pages 749-770 (Part II:
May). "Factors" was reprinted in pages 389-466 of Herbert Spencer (1891). The Works of Herbert
Spencer. 13. London: Williams and Norgate. (also Osnabrück: Otto Zeller, 1967) = volume 1
ofEssays: Scientific, Political, and Speculative .
23.^ One example of a response by was in Section III(γ) pages 32-42 of August Weismann
(1909). "The Selection theory". in Albert Charles Seward. Darwin and Modern Science: Essays in
Commemoration of the Centenary of the Birth of Charles Darwin and of the Fiftieth Anniversay of the
Publication of The Origin of Species. Cambridge: Cambridge University Press. pp. 19–65. See also
Chapter VII, §12(1), pages 237-238 in: J. Arthur Thomson (1908). Heredity. London: John Murray.
Both of these referred to what has become known as the Baldwin effect. An analysis of both sides of
the issue is: George John Romanes (1895). "III: Characters as Hereditary and Acquired (continued)".
Darwin and After Darwin: Post-Darwinian Questions, Heredity, Utility . II. London: Longman, Green.
pp. 60–102.
24.^ Pages 67-68 in: Mark Ridley (March,1982). "Coadapatation and the Inadequacy of Natural
Selection". British Journal for the History of Science 15 (1): 45–68.
doi:10.1017/S0007087400018938.
25.^ Muller HJ (1918). "Genetic variability, twin hybrids and constant hybrids, in a case of
balanced lethal factors". Genetics 3 (5): 422–99. PMID 17245914. PMC 1200446.
http://www.genetics.org/cgi/reprint/3/5/422. , especially pages 463–4.
26.^ Muller HJ (1939). "Reversibility in evolution considered from the standpoint of genetics".
Biological Reviews of the Cambridge Philosophical Society 4 (3): 261–80, quotation from 272.
27.^ Morris, Henry (1974). Scientific creationism (2nd ed.). San Diego, Calif: Creation-Life
Publishers. pp. 59. ISBN 0-89051-003-2.
 28.^ T. H. Frazzetta, Complex Adaptations in Evolving Populations, Sunderland,
Massachusetts: Sinauer Associates, 1975. ISBN 0-87893-194-5. Referencing pages 3, 4-7, 7-20,
and xi, respectively.
29.^ Keough, Mark J.; Geisler, Norman L. (1982). The Creator in the courtroom "Scopes II": the
1981 Arkansas creation-evolution trial. Milford, Mich: Mott Media. p. 146. ISBN 0-88062-020-X.
http://www.antievolution.org/projects/mclean/new_site/docs/geislerbook.htm#Chapter%20Seven.
30.^ Cairns-Smith, A. G. (1986). Seven clues to the origin of life: a scientific detective story .
Cambridge, UK: Cambridge University Press. pp. 39, 58–64. ISBN 0-521-27522-9.
31.^ Mark Perakh (2008). "Bacteria Flagella Look Like Man-made Machines". Skeptic (U.S.
magazine). http://www.skeptic.com/the_magazine/. Retrieved 2008-12-06.
32.^ Lumsden RD (June 1994). "Not So Blind A Watchmaker". Creation Research Society
Quarterly 31 (1): 13–22, quotations from 13, 20.
33.^ Scott EC, Matzke NJ (May 2007). "Biological design in science classrooms". Proc Natl
Acad Sci USA. 104 Suppl 1: 8669–76, See page 8672. doi:10.1073/pnas.0701505104.
PMID 17494747.
34.^ Ludwig von Bertalanffy (1952). Problems of Life: An Evaluation of Modern Biological and
Scientific Thought, pg 148 ISBN 1-131-79242-4
35.^ Monod, Jacques (1972). Chance and necessity: an essay on the natural philosophy of
modern biology. New York: Vintage Books. ISBN 0-394-71825-9.
36.^ Barbara Forrest, The Wedge at Work. Talk Reason.
Forrest B (2001). "1: The Wedge at Work: How Intelligent Design Creationism is Wedging its way into
the Cultural and Academic Mainstream". in Pennock RT. Intelligent design creationism and its critics:
philosophical, theological, and scientific perspectives . Cambridge, Mass: MIT Press. pp. 5–54.
ISBN 0-262-66124-1.
 37.^ The New Pandas: Has Creationist Scholarship Improved? Comments on 1993 Revisions
by Frank J. Sonleitner (1994)
Introduction: Of Pandas and People, the foundational work of the 'Intelligent Design' movement by
Nick Matzke 2004,
Design on Trial in Dover, Pennsylvania by Nicholas J Matzke, NCSE Public Information Project
Specialist
38.^ Behe MJ (November 2001). "Reply to My Critics: A Response to Reviews of Darwin's Black
Box: The Biochemical Challenge to Evolution". Biology and Philosophy 16 (5): 685–709.
doi:10.1023/A:1012268700496. http://friends-of-wisdom.com/readings/Behe2001.pdf.
39.^ Behe's testimony in Kitzmiller v. Dover
40.^ Behe, Michael 2005 Kitzmiller v. Dover Area School District 4: whether ID is science (p. 88)
41.^ Kitzmiller v. Dover Area School District 6: Conclusion, section H
42.^ a b c A reducibly complex mousetrap (graphics-intensive, requires JavaScript)
43.^ a b c d Miller, Kenneth R. (2008). Only A Theory. New York: Viking Penguin. pp. 54–55.
ISBN 978-0-670-01883-3.
44.^ Behe, Michael "Comments on Ken Miller's Reply to My Essays"
45.^ Claim CA350: Professional literature is silent on the subject of the evolution of biochemical
systems TalkOrigins Archive.
46.^ Behe, Michael J. (1996) [1996]. Darwin's black box: the biochemical challenge to evolution .
New York, NY: Free Press. pp. 72. ISBN 0684827549. ""Yet here again the evolutionary literature is
totally missing. No scientist has ever published a model to account for the gradual evolution of this
extraordinary molecular machine.""
47.^ a b Ruling, Kitzmiller v. Dover Area School District, December 2005. Page 74.
48.^ Action, George "Behe and the Blood Clotting Cascade"
 49.^ Semba U, Shibuya Y, Okabe H, Yamamoto T (1998). "Whale Hageman factor (factor XII):
prevented production due to pseudogene conversion". Thromb Res 90 (1): 31–7. doi:10.1016/S0049-
3848(97)00307-1. PMID 9678675.
50.^ Behe, Michael [http://www.arn.org/docs/behe/mb_indefenseofbloodclottingcascade.htm "In
Defense of the Irreducibility of the Blood Clotting Cascade: Response to Russell Doolittle, Ken Miller
and Keith Robison"]
51.^ Creationism special: A sceptic's guide to intelligent design, New Scientist, 9 July 2005
52.^ Sarfati, Jonathan (2000). Argument: 'Irreducible complexity', from Refuting Evolution
(Answers in Genesis).
53.^ CA113.1: Evolution of the eye
54.^ Darwin, Charles (1859). On the Origin of Species. London: John Murray. pages 186ff,
Chapter VI
55.^ Halder G, Callaerts P, Gehring WJ (October 1995). "New perspectives on eye evolution".
Curr Opin Genet Dev. 5 (5): 602–9. doi:10.1016/0959-437X(95)80029-8. PMID 8664548.
http://linkinghub.elsevier.com/retrieve/pii/0959-437X(95)80029-8.
56.^ Halder G, Callaerts P, Gehring WJ (March 1995). "Induction of ectopic eyes by targeted
expression of the eyeless gene in Drosophila". Science 267 (5205): 1788–92.
doi:10.1126/science.7892602. PMID 7892602. http://www.sciencemag.org/cgi/pmidlookup?
view=long&pmid=7892602.
57.^ Tomarev SI, Callaerts P, Kos L, et al. (March 1997). "Squid Pax-6 and eye development".
Proc Natl Acad Sci USA. 94 (6): 2421–6. doi:10.1073/pnas.94.6.2421. PMID 9122210. PMC 20103.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=9122210.
58.^ Fernald, Russell D. (2001). The Evolution of Eyes: Why Do We See What We See? Karger
Gazette 64: "The Eye in Focus".
 59.^ Fernald RD (1988). "Aquatic Adaptations in Fish Eyes". in Atema J. Sensory biology of
aquatic animals. Berlin: Springer-Verlag. ISBN 0-387-96373-1.
60.^ Fernald RD (1997). "The evolution of eyes". Brain Behav Evol. 50 (4): 253–9.
doi:10.1159/000113339. PMID 9310200.
61.^ Conway-Morris S (1999). The Crucible of Creation: The Burgess Shale and the Rise of
Animals. Oxford [Oxfordshire]: Oxford University Press. ISBN 0-19-286202-2.
62.^ Benjamin Wiker & Jonathan Witt (2006). A Meaningful World. p. 44.
63.^ Miller, Kenneth R. The Flagellum Unspun: The Collapse of "Irreducible Complexity" with
reply here
64.^ Kenneth Miller's The Collapse of Intelligent Design: Section 5 Bacterial Flagellum (Case
Western Reserve University, 2006 January 3)
65.^ Unlocking cell secrets bolsters evolutionists (Chicago Tribune, 2006 February 13)
66.^ Evolution in (Brownian) space: a model for the origin of the bacterial flagellum (Talk Design,
2006 September)
67.^ Mirsky, Steve Sticker Shock: In the beginning was the cautionary advisory Scientific
American, February 2005
68.^ Matt Inlay, 2002. "Evolving Immunity." In TalkDesign.org.
69.^ Nicholas J. Matzke, 2003. "Evolution in (Brownian) space: a model for the origin of the
bacterial flagellum."
70.^ Pallen MJ, Matzke NJ (October 2006). "From The Origin of Species to the origin of bacterial
flagella". Nat Rev Microbiol. 4 (10): 784–90. doi:10.1038/nrmicro1493. PMID 16953248.
http://www.pandasthumb.org/archives/2006/09/flagellum_evolu.html.
71.^ Pigliucci, Massimo [3] Collaboration Sept. 2001
 72.^ Shanks, Niall; Joplin, Karl H. (1999). "Redundant Complexity: A Critical Analysis of
Intelligent Design in Biochemistry". Philosophy of Science (The University of Chicago Press) 66 (2,
June): 268–282. doi:10.1086/392687. http://www.jstor.org/pss/188646.
73.^ Lenski RE, Ofria C, Pennock RT, Adami C (2003). "The evolutionary origin of complex
features". Nature 423 (6936): 139–44. doi:10.1038/nature01568. PMID 12736677.
74.^ Press release University of Oregon, April 4, 2006.
75.^ M. Brazeau and P. Ahlberg (January 19, 2006). "Tetrapod-like middle ear architecture in a
Devonian fish". Nature 439 (7074): 318–21. doi:10.1038/nature04196. PMID 16421569.
76.^ "NOVA : Transcripts : Judgment Day: Intelligent Design on Trial Chapter 8". PBS.
November 13, 2007. http://www.pbs.org/wgbh/nova/transcripts/3416_id_08.html. Retrieved 2008-12-
17.
77.^ Miller K (1999). Finding Darwin's God: a scientist's search for common ground between
God and evolution. New York: Cliff Street Books. ISBN 0060930497.
78.^ 36 Human-Competitive Results Produced by Genetic Programming
79.^ Michael Behe. Evidence for Intelligent Design from Biochemistry. 1996.
80.^ Eugenie C. Scott and Glenn Branch, "Intelligent Design" Not Accepted by Most Scientists,
National Center for Science Education website, September 10, 2002.
81.^ IC and Evolution makes the point that: if "irreducible complexity" is tautologically redefined
to allow a valid argument that intelligent design is the correct explanation for life then there is no such
thing as "irreducible complexity" in the mechanisms of life; while, if we use the unmodified original
definition then "irreducible complexity" has nothing whatever to do with evolution.
82.^ The Court in Dover noted that this implicit assumption of the defendant school board
created a "flawed and illogical contrived dualism" (Opinion p. 64).
83.^ a b Memorandum Opinion, Judge John E. Jones III, Kitzmiller v. Dover Area School District
[edit] Additional references
• Behe, Michael (1996). Darwin's Black Box. New York: The Free Press. ISBN 0-684-83493-6
• Denton, Michael (1986). Evolution: A Theory in Crisis. Bethesda, Md: Adler & Adler. ISBN 0-
917561-05-8.
• Macnab RM (2004). "Type III flagellar protein export and flagellar assembly". Biochim
Biophys Acta 1694 (1-3): 207–17. doi:10.1016/j.bbamcr.2004.04.005. PMID 15546667.
• Ruben, J.A.; Jones, T.D.; Geist, N.R.; & Hillenius, W.J. (November 14, 1997). "Lung
Structure and Ventilation in Theropod Dinosaurs and Early Birds". Science 278 (5341): 1267–70.
doi:10.1126/science.278.5341.1267.
• Sunderland, Luther D. (March 1976). Miraculous Design in Woodpeckers. Creation
Research Society Quarterly.
• Testing Darwin Discover Magazine Vol. 26 No. 02 | February 2005

[edit] External links

Supportive
• Michael J. Behe home page
• About Irreducible Complexity Discovery Institute
• How to Explain Irreducible Complexity -- A Lab Manual Discovery Institute
• Institute for Creation Research (pdf)
• Irreducible Complexity Revisited (pdf)
• Behe's Reply to his Critics (pdf)
 • Response to Avida computer simulations
Critical
• Behe, Biochemistry, and the Invisible Hand
• Facilitated Variation
• Darwin vs. Intelligent Design (again), by H. Allen Orr (review of Darwin's Black Box)
• Talk.origins archive (see talk.origins)
• Irreducible Complexity and Michael Behe: Do Biochemical Machines Show Intelligent
Design?
• The Mullerian Two-Step: Add a part, make it necessary by Douglas Theobald
• Darwin's Black Box: Irreducible Complexity or Irreproducible Irreducibility? by Keith
Robinson
• Is the Complement System Irreducibly Complex? by Mike Coon
• Genetic Algorithms (Genetic algorithms have produced irreducibly complex solutions
to real problems.)
• Discussion of the Bombardier Beetle at Talk.origins
• TalkDesign.org (sister site to talk.origins archive on intelligent design)
• Irreducible complexity demystified by Pete Dunkelberg.
• Professor Kenneth R. Miller's textbook website
• A Darwinian explanation of the blood clotting cascade
• "The Flagellum Unspun: The Collapse of "Irreducible Complexity" by Professor Miller
• A rigorous mathematical analysis of the concept of irreducible complexity by Mishkin Berteig.
 • PDF. 139 page in-depth analysis of Intelligent Design, Irreducible Complexity, and the book
"Of Pandas and People" by a judge and based on expert testimony
• Devolution: Why intelligent design isn't (The New Yorker)
• Unlocking cell secrets bolsters evolutionists (Chicago Tribune)
• Does irreducible complexity imply Intelligent Design? by Mark Perakh
• Evolution of the Eye (Video) Zoologist Dan-Erik Nilsson demonstrates eye evolution through
intermediate stages with working model. (PBS)
• Himma, Kenneth Einar. Design Arguments for the Existence of God. Internet Encyclopedia of
Philosophy: 2. Contemporary Versions of the Design Argument, a. The Argument from Irreducible
Biochemical Complexity

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Genesis Genesis creation narrative · Framework interpretation · Day-Age theory ·

interpretations Gap theory · Biblical literalism

Related concepts Creation myth · Omphalos hypothesis · Specified complexity ·

Irreducible complexity · Dating Creation · Theistic realism · Intelligent designer
 Pseudoscience Creation science · Creationist cosmologies · Creation biology · Created
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Retrieved from "http://en.wikipedia.org/wiki/Irreducible_complexity"

Categories: Creationist objections to evolution | Intelligent design | Intelligent design movement |

Biological systems | Complex systems theory
 W000

Complexité irréductible
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Cet article traite de la complexité irréductible telle qu'elle est utilisée par les partisans du dessein
intelligent ((en) intelligent design). Pour la complexité irréductible dans le cadre de la théorie des
systèmes, voir l'article Émergence.
La complexité irréductible est la thèse selon laquelle certains systèmes biologiques sont trop
complexes pour être le résultat de l'évolution de précurseurs plus simples ou « moins complets », du fait de
mutations au hasard et de la sélection naturelle. Le terme a été inventé et défini en 1996 par le professeur de
biochimie Michael Behe, un système de complexité irréductible étant, « composé de plusieurs parties
ajustées et interagissantes, qui contribuent chacune à sa fonction élémentaire, alors que l'absence d'une
quelconque de ces parties empêche le fonctionnement du système »[1]. Les exemples cités par Behe, la
coagulation en cascade, le moteur (ou corps basal) des flagelles cellulaires et le système immunitaire, ne
pourraient donc être le résultat de l'évolution naturelle : tout système précurseur au système complet ne
fonctionnerait pas, et ne constituerait donc pas un avantage sélectif.
De façon plus générale, cet argument est utilisé par les partisans du créationnisme et du dessein
intelligent pour réfuter la théorie scientifique actuelle de l'évolution et prouver l'implication d'une cause divine
ou intelligente dans la création de la vie. Ces thèses sont anciennes et reprennent l'argument téléologique de
l'analogie du grand horloger. En dehors des systèmes biochimiques présentés par Behe, un exemple très
couramment avancé de système trop complexe pour être le résultat de l'évolution est l'œil.
La thèse de la complexité irréductible est rejetée par une très large majorité de la communauté
scientifique[2] ; elle est souvent considérée comme pseudoscientifique[3]. Des travaux scientifiques ont
montré que les exemples présentés par Behe ne répondaient pas à sa définition, et des précurseurs ont été
identifiés pour certains d'entre eux. Les critiques considèrent que la thèse de la complexité irréductible est
fondée sur une incompréhension du fonctionnement de ces systèmes biochimiques, et une méconnaissance
des mécanismes de l'évolution (en particulier l'exaptation). Elle est également considérée comme un
excellent exemple d'argumentum ad ignorandam (argument d'ignorance, sophisme par lequel on déclare
fausse une proposition qui n'a pas été démontrée vraie).
Bien qu'elle aie été rejetée en tant que théorie scientifique lors du procès de Dover[4], à l'issue
duquel la cour a jugé que « La thèse du Professeur Behe sur la complexité irréductible a été réfutée par des
articles scientifiques publiés dans des revues à comité de lecture, et a été rejetée par la communauté
scientifique dans son ensemble » procès de Dover (p. 64)[2], le concept de complexité irréductible reste un
argument courant pour les partisans du dessein intelligent et d'autres créationnistes.
Sommaire
[masquer]
• 1 Histoire du concept
• 1.1 La preuve téléologique et le grand horloger
• 1.2 La complexité irréductible
• 1.2.1 Définitions
• 1.2.2 Explications et implications : le
dessein intelligent
• 2 Exemples présentés
• 2.1 L'œil
• 2.2 Exemples en biochimie cellulaire
• 2.2.1 La tapette à souris
• 2.2.2 Le flagelle
• 2.2.3 La coagulation sanguine
• 3 Réponse des évolutionnistes
• 3.1 Réductibilité des « systèmes irréductibles »
• 3.2 Évolution des systèmes irréductibles
• 3.3 Les échafaudages de l'évolution
• 3.3.1 Adaptation progressive à de nouvelles
fonctions : l'exaptation
• 3.3.2 Sophismes
Histoire du concept [modifier]
La preuve téléologique et le grand horloger [modifier]
L'argument de complexité irréductible est un descendant de la preuve téléologique de l'existence de
Dieu. C'est l'argument fameux du grand horloger que la complexité de la nature implique l'existence de Dieu
de la même façon que l'existence d'une horloge implique celle d'un horloger.
Cet argument a une longue histoire qui remonte au moins au Ier siècle av. J.-C. avec De natura
deorum (ii.34) de Cicéron : « Un cadran solaire ou une clepsydre donnent l'heure du fait de leur conception
et non par hasard. Comment pouvez-vous donc imaginer que l'univers, comme un tout, est sans but et sans
intelligence, quand il contient tout, y compris ces objets et leurs artisans ?»
Le déiste Voltaire le reprend dans un distique célèbre de sa satire Les cabales (1772) :
« L'univers m'embarrasse, et je ne puis songer
Que cette horloge existe et n'ait point d'horloger. »

Citation à mettre en perspective avec ce propos de jeunesse dans le Traité de métaphysique (1734) :
« Ainsi, quand je vois les ressorts du corps humain, je conclus qu’un être intelligent a arrangé
ces organes pour être reçus et nourris neuf mois dans la matrice; que les yeux sont donnés
pour voir, les mains pour prendre, etc. Mais de ce seul argument je ne peux conclure autre
chose, sinon qu’il est probable qu’un être intelligent et supérieur a préparé et façonné la matière
avec habileté; mais je ne peux conclure de cela seul que cet être ait fait la matière avec rien, et
qu’il soit infini en tout sens. J’ai beau chercher dans mon esprit la connexion de ces idées: « Il
 est probable que je suis l’ouvrage d’un être plus puissant que moi, donc cet être existe de toute
éternité, donc il a créé tout, donc il est infini, etc. ». Je ne vois pas la chaîne qui mène droit à
cette conclusion ; je vois seulement qu’il y a quelque chose de plus puissant que moi, et rien de
plus. »

Les implications de la complexité des organismes vivants et des interactions entre leurs éléments ont
été discutées par différents auteurs et savants. Au début du XVIIe siècle, Nicolas Malebranche[5] utilise cette
idée en faveur de la préformation (théorie qui voit l'embryon comme un être vivant « miniature » où tous les
organes sont déjà présents), plutôt que de l'épigénèse (la complexité apparaît au fur et à mesure du
développement). Dans une application différente, au début du XIXe siècle Georges Cuvier utilisa le concept
de « corrélation des parties » dans la reconstitution de l'anatomie d'animaux à partir de restes épars[6],[7].
Dans le monde anglo-saxon, l'argument du grand horloger est développé et popularisé par le
révérend William Paley dans sa Théologie naturelle (1802). Prenant l'analogie d'une montre trouvée par
hasard, il conclut que la structure complexe des êtres vivants et l'adaptation remarquable des plantes et des
animaux sont le fait d'un concepteur intelligent. Le monde est donc une création de Dieu, et montre la nature
de ce créateur. Dieu a conçu avec attention « même le plus humble et le plus insignifiant des organismes ».
Charles Darwin rejette la preuve téléologique en proposant une autre explication à la complexité et la
diversité du vivant : l'évolution par la sélection naturelle. Sans le nommer ainsi, il identifie l'argument de la
complexité irréductible comme un moyen possible de réfuter les résultats de sa théorie de l'évolution. Dans
L'Origine des espèces[8], il écrit : « Si on pouvait démontrer qu'il existe un organe complexe, qui ne pourrait
avoir été formé par de nombreuses petites modifications successives, ma théorie s'effondrerait
complètement. Mais je n'en trouve aucun exemple ».
 Le généticien Hermann Muller expose, en 1939, un concept similaire à la complexité irréductible,
mais pas de façon problématique vis-à-vis de l'évolution. Au contraire, il présente l'« intrication » des
éléments biologiques comme une conséquence attendue de l'évolution, qui conduit à l'irréversibilité de
certains changements évolutifs[9] : Ayant été ainsi entretissée au sein de la trame la plus intime de
l'organisme, la caractéristique précédemment nouvelle ne peut plus être retirée impunément, et peut être
devenue nécessaire et vitale[10].
En 1952, le biologiste Ludwig von Bertalanffy propose un concept précurseur de la complexité
irréductible[11] : les systèmes organiques complexes doivent être étudiés en tant que systèmes complets
irréductibles pour pouvoir comprendre leur fonctionnement. Il étend ses travaux sur la complexité biologique
à une théorie générale des systèmes. Après la découverte de la structure de l'ADN par James Watson et
Francis Crick au début des années 1950, la théorie générale des systèmes perd la plupart de ses adhérents
en sciences physiques et biologie. Ce triomphe du point de vue mécaniste en biochimie est exposé dans Le
Hasard et la Nécessité de Jacques Monod[12]. Cependant, la théorie des systèmes reste utilisée dans les
sciences sociales.

La complexité irréductible [modifier]

Dans son livre Darwin's Black Box[1], le biochimiste Michael Behe, reprenant la thèse de William
Paley, définit et applique le terme « complexité irréductible » à certains systèmes complexes en biologie
cellulaire. Il cherche à montrer que les mécanismes de l'évolution ne peuvent expliquer le développement de
ces systèmes « irréductiblement complexes ».
 Définitions [modifier]
Le terme « complexité irréductible » a été défini par Behe :
« Un système composé de plusieurs parties en interaction, qui contribuent chacune à sa
fonction élémentaire, et dont l'absence d'une quelconque de ces parties empêche le
fonctionnement du système Behe (Darwin's Black Box[1], p. 9). »

Behe a donné une seconde définition (« définition évolutionniste ») :

« Une lignée évolutive est de complexité irréductible si elle contient au moins une étape
insélectionnable (c'est-à-dire au moins une mutation nécessaire mais insélectionnable – au sens
de la sélection naturelle). Le degré de complexité irréductible est le nombre d'étapes
insélectionnables dans la lignée. »

Le partisan du dessein intelligent William Dembski donne cette définition :

« Un système effectuant une fonction basique donnée est de complexité irréductible, s'il
comprend un ensemble d'éléments individualisés de façon non-arbitraire, ajustés et
interagissant mutuellement, tel que chaque élément de l'ensemble est indispensable à la
fonction basique, et donc original. L'ensemble de ces éléments indispensables constitue le
noyau irréductible du système[13]. »
 Explications et implications : le dessein intelligent [modifier]
Les partisans du dessein intelligent utilisent ce concept et les exemples de systèmes de « complexité
irréductible », pour en conclure que le monde vivant est mieux expliqué par l'intervention d'une cause
intelligente que par les mécanismes de la théorie de l'évolution.
Selon la théorie de l'évolution, les variations génétiques adviennent au hasard. L'environnement
« sélectionne » les variations les mieux adaptées, qui sont ensuite transmises aux générations suivantes.
Les changements au cours du temps se font du fait de l'action graduelle des forces naturelles, de façon
parfois lente ou parfois rapide (voir équilibre ponctué). Ce processus permet l'adaptation de structures
complexes depuis des formes plus simples, ou de convertir des structures complexes d'une fonction à une
autre.
Behe et la plupart des partisans du dessein intelligent ne remettent pas en cause tout le rôle des
mécanismes évolutifs dans le dévoppement de la vie organique à l'échelle de la microévolution (comme les
variations de longueur des becs des pinsons de Darwin). Mais ils jugent qu'ils ne peuvent rendre compte de
la complexité irréductible, car aucun des éléments d'un système irréductible ne serait fonctionnel ou
avantageux avant que le système complet ne soit en place : « un système de complexité irréductible ne peut
pas être produit directement (c'est-à-dire, par une amélioration continue de la fonction initiale, qui garde le
même mécanisme) par de petites modifications successives d'un système précurseur, car tout précurseur
d'un système de complexité irréductible auquel il manque un élément est par définition non-fonctionnel. »
(Behe)
La complexité irréductible n'est donc pas un argument que l'évolution n'existe pas, mais plutôt qu'elle
est « incomplète ». Dans le dernier chapitre de Darwin's Black Box[1], Behe conclut que la complexité
irréductible est une preuve du dessein intelligent.
Exemples présentés [modifier]
Behe et d'autres ont présenté un certain nombre d'exemples de systèmes biologiques qu'il pensent
être de complexité irréductible. Les exemples donnés ne sont pas, selon la communauté scientifique dans
son ensemble, probants.
L'œil [modifier]

Souvent présenté comme un exemple de complexité irréductible, l'œil humain peut être expliqué par
l'évolution.
 L'œil est un exemple fameux de structure présentée comme de « complexité irréductible », du fait de
des nombreux éléments intriqués et sophistiqués, dépendant apparemment tous les uns des autres. Il est
fréquemment cité par les partisans du dessein intelligent et du créationnisme.
Dans un passage fréquemment cité de L'Origine des espèces[8], Charles Darwin reconnaît lui-même
que le développement de l'œil est une difficulté pour sa théorie, notant que « supposer que l'œil [...] peut
avoir été formé par sélection naturelle semble, je le confesse volontiers, absurde au plus haut degré ».
Cependant il continue en notant que si « la difficulté de croire que l'œil complet et parfait peut être formé par
le mécanisme de la sélection naturelle, bien qu'insurmontable par notre imagination, ne peut être considérée
comme réelle », et il propose un schéma grossier de lignée évolutive possible, à partir d'exemples de plus en
plus complexes d'yeux de différentes espèces (Charles Darwin, L'Origine des espèces, p. 186 et suiv[8]).
 Les yeux des vertébrés (à gauche) et des invertébrés comme la pieuvre (à droite) ont évolué
indépendamment : les vertébrés ont développé une rétine 1 inversée avec un point aveugle 4 au niveau du
nerf optique 3, alors que les pieuvres l'ont évité grâce à une rétine non-inversée (les terminaisons nerveuses
2 sont à l'extérieur).
Depuis Darwin l'évolution de l'œil est bien mieux comprise. Bien que l'observation et l'analyse des
ancêtres de l'œil dans les fossiles soit problématique, du fait que les tissus mous ne laissent pas
d'empreintes ou de restes, la génétique et l'anatomie comparative vont dans le sens d'un ancêtre commun
pour tous les yeux[14],[15],[16].
Les éléments actuels permettent de proposer des lignées évolutives possibles aboutissant aux
caractéristiques anatomiques de l'œil. Un schéma évolutif possible est le suivant :
• les yeux se sont développés à partir de simples petites surfaces de cellules photoréceptrices
pouvant détecter la présence ou l'absence de lumière, mais pas sa direction.
• En développant une petite dépression de ces cellules photosensibles, l'organisme obtient
une meilleure perception de la source lumineuse, en permettant à la lumière de ne frapper que
certaines cellules, en fonction de l'angle.
• Cette dépression se creusant, la précision de l'observation s'améliore.
• L'ouverture de l'œil se rétrécit alors pour augmenter cette précision, ce qui transforme l'œil
en une chambre noire, permettant à l'organisme de percevoir les formes — le nautile est un exemple
actuel d'animal ayant un tel œil. Le compromis nécessaire entre la précision ainsi obtenue, et la
baisse du flux de lumière captée limite les possibilités de développement dans cette direction.
• Mais ces exigences deviennent compatibles si la couche protectrice de cellules
transparentes couvrant l'ouverture vient former une lentille grossière, et l'intérieur de l'œil se remplit
 d'humeur qui introduit une focalisation[17],[18],[19]. De cette façon, l'œil est en fait considéré par les
biologistes modernes comme une structure à l'évolution relativement simple et sans mystère, et la
plupart de ses évolutions majeures ont eu lieu en seulement quelques millions d'années, durant
l'explosion cambrienne[20]. Ceci est d'autant plus crédible que plusieurs dispositifs en gros
équivalents existent dans la nature : l'œil des vertébrés, celui des pieuvres et celui des mouches,
avec des dispositions totalement différentes.

La tapette à souris, exemple illustratif de « complexité irréductible » utilisé par Michael Behe[21]
Selon Behe, si l'évolution des grandes caractéristiques anatomiques de l'œil a été bien expliquée, la
complexité du détail des réactions biochimiques nécessaires à l'échelle moléculaire pour la sensibilité à la
lumière, défie encore les explications. Le créationniste Jonathan Sarfati décrit l'œil comme « le plus grand
défi [des évolutionnistes] en tant que superbe exemple de complexité irréductible dans la création divine »,
mettant particulièrement en avant la « grande complexité » de la transparence de la cornée[22].
Exemples en biochimie cellulaire [modifier]
Michael Behe a proposé un certain nombre d'exemples de systèmes biochimiques complexes de
« complexité irréductible » : le flagelle des cellules, la coagulation sanguine, et le système immunitaire.

La tapette à souris [modifier]

Behe utilise la tapette à souris comme exemple illustratif de ce concept. Une tapette à souris est
constituée de plusieurs pièces - la base, le déclencheur, le ressort, l'arceau. Toutes ces pièces doivent être
en place pour que la tapette fonctionne, et la suppression de n'importe laquelle entraîne la perte de cette
fonctionnalité.
 La coagulation en cascade : HWMK = High molecular weight kininogen, PK = Prékallikréine, TFPI =
Tissue factor pathway inhibitor. Flèches noires = conversion/activation du facteur. Flèches rouges = action
sur les inhibiteurs. Flèches bleues = réactions catalysées par le facteur activé. Flèches grises = diverses
fonctions de la thrombine.
De la même façon, les systèmes biologiques nécessitent plusieurs éléments travaillant ensemble
pour fonctionner. Selon lui, il n'est pas possible de trouver une succession de petites évolutions viables, car
l'avantage sélectif de la fonction n'est présent que quand tous les éléments sont assemblés.

Le flagelle [modifier]
Le flagelle, comme les cils de certaines cellules et bactéries, constitue un moteur moléculaire rotatif
qui assure leur mobilité. Dans le cas des procaryotes (comme la bactérie E. coli), leur fonctionnement
nécessite l'interaction d'un quarantaine de protéines complexes, et l'absence d'une seule de ces protéines
empêche le flagelle de fonctionner.

La coagulation sanguine [modifier]

La coagulation sanguine chez les vertébrés se fait par une cascade complexe de processus
biochimiques présentée par Behe comme un exemple de complexité irréductible[23].
Il en est de même des anticorps du système immunitaire, qui présentent à la fois une substance
marqueur et une substance tueur, indispensables l'une et l'autre au fonctionnement.
Réponse des évolutionnistes [modifier]
De même que le dessein intelligent, concept qu'elle est censée soutenir, la complexité irréductible a
été largement rejetée par la communauté scientifique. Les critiques jugent qu'une « complexité irréductible »,
telle qu'elle est définie par Behe, peut être générée par des mécanismes évolutifs connus, comme
l'exaptation. Des chemins évolutifs plausibles ont été proposés pour les trois exemples présentés par Behe
comme étant de complexité irréductible : la coagulation, le système immunitaire[24] et le flagelle[25],[26]. Il
en est de même l'exemple illustratif de la tapette à souris : le professeur de biologie John McDonald a montré
comment on pouvait la considérer comme « facilement réductible », jusqu'à un seul élément[21].

Réductibilité des « systèmes irréductibles » [modifier]

La thèse de la complexité irréductible implique que les éléments nécessaires d'un système ont
toujours été nécessaires, et ne peuvent donc avoir été ajoutés les uns après les autres. Cependant, dans
l'évolution, un élément qui est seulement avantageux au début peut devenir nécessaire ensuite. Par
exemple, on a découvert par la suite qu'un des facteurs de la coagulation, cité par Behe comme un élément
de la cascade de coagulation, était absent chez les baleines, et n'est donc pas indispensable pour la
coagulation[27]. On a montré récemment que le corps basal du flagelle est similaire au système de sécrétion
de type III (TTSS), une structure en forme d'aiguille que des germes pathogènes tels que la salmonelle et
Yersinia pestis utilise pour injecter des toxines dans les cellules eucaryotes. La base de l'aiguille a de
nombreux éléments communs avec le flagelle, mais sans la plupart des protéines qui font fonctionner le
flagelle. Ainsi, retirer des éléments du flagelle ne le rend pas nécessairement inutile. Ainsi, selon Miller, « les
éléments de ce système réputé de complexité irréductible ont en fait des fonctions qui leurs sont
propres. »[28],[29],[30]. Cela est vrai pour la majeure partie de la structure du flagelle ; sur 42 protéines qu'on
y trouve, 40 ont été observées dans différents canaux biologiques[31].
Évolution des systèmes irréductibles [modifier]
Différents mécanismes évolutifs permettent d'expliquer la formation de systèmes de complexité
apparemment irréductible.
Niall Shanks et Karl H. Joplin, de East Tennessee State University, ont montré que des systèmes
satisfaisants à la définition de Behe de la complexité biochimique irréductible peuvent apparaître
naturellement et spontanément du fait de processus chimiques auto-organisés[32],[33]. Ils affirment
également que les systèmes biochimiques et moléculaires évolués présentent en fait une « complexité
redondante » – ils jugent que Behe surestime l'importance de la complexité irréductible à cause d'une vision
simpliste et linéaire des réactions biochimiques, prenant des instantanés des caractérististiques de
systèmes, structures et processus biologiques et ignorant la complexité redondante du contexte dans lequel
s'insèrent ces caractéristiques. En outre, des simulations informatiques ont montré qu'il était possible pour
des systèmes de complexité irréductible d'évoluer naturellement[34].
Le professeur Marc W. Kirschner, directeur du département de biologie des systèmes de la Harvard
Medical School, et John C. Gerhart, professeur en biologie moléculaire et cellulaire à University of California,
Berkeley ont proposé en 2005 la théorie de la variation facilitée, qui explique comment certaines mutations
ou changement peuvent engendrer une apparente complexité irréductible.

Les échafaudages de l'évolution [modifier]

Le fait que retirer un élément d'un système organique entraîne le non-fonctionnement ne prouve pas
que le système ne peut avoir été formé par un processus évolutif progressif. En 1985, Graham Cairns-Smith
se demande à propos de l'« intrication » des systèmes biologiques « Comment une collaboration complexe
entre des éléments peut-elle évoluer par petites étapes ? », et répond en utilisant l'analogie de l'échafaudage
dans la construction d'une arche de pierre ; si on retire n'importe quelle pierre d'une arche, elle s'effondre (en
ce sens elle est de « complexité irréductible ») ; pourtant elle peut être construite sans problème, une pierre
après l'autre, à l'aide d'un échafaudage (ou cintre) que l'on retire après : « Clairement il y a eu des
échafaudages. Avant que les multiples éléments de la biochimie actuelle ne puissent s'appuyer les uns sur
les autres, ils ont dû s'appuyer sur quelque chose d'autre »[35].
L'évolution n'agit pas nécessairement linéairement vers la complexification ; elle peut simplifier tout
autant que complexifier. De ce fait des systèmes biologiques apparemment de complexité irréductible
peuvent être le résultat d'une phase de complexification suivie d'une phase de simplification.
Pour l'exemple des anticorps, présenté par Behe, on a la substance « marqueur » et la substance
« tueur », qui, pour la première, repère et marque les « envahisseurs », et, pour la seconde, tue ce qui a été
marqué. Selon Behe, le marqueur et le tueur sont, par eux-mêmes, inutiles, et doivent donc avoir été créés
en même temps. Le tueur ne peut tuer ce qu'il ne peut trouver, et le marqueur ne peut tuer même s'il trouve
une cible.
Cependant, avec un remplacement progressif, un marqueur différent peut avoir commencé comme
tueur et marqueur à la fois. Puis un aide-tueur rejoint cette armée du fait de ses particularités intéressantes.
Ce second tueur dépend toujours du premier pour trouver la cible. Il est possible qu'au cours du temps le
premier tueur-marqueur se spécialise en marquage et soit remplacé par une substance similaire uniquement
marqueur (éventuellement plus adaptée que la première à double usage).
Ainsi chaque étape apporte un avantage sélectif, et le résultat final est une paire interdépendante qui
ne ressemble pas à la substance initiale. Cet exemple peut être représenté ainsi :
A = tueur-marqueur original
 K = deuxième tueur
M = marqueur de remplacement
1. A
2. A K
3. A M K
4. M K
Ce qu'on observe aujourd'hui est « M K ». Selon les opposants au concept de complexité, Behe se
trompe en postulant que si la structure actuelle est M K, elle a dû commencer soit par M soit par K.

Adaptation progressive à de nouvelles fonctions : l'exaptation [modifier]

Les arguments pour l'irréductibilité présupposent généralement que les choses ont commencé de la
même façon qu'elles ont fini – c'est-à-dire comme on les observe actuellement. Mais ce n'est pas
nécessairement le cas.
Les précurseurs de systèmes complexes quand ils ne sont pas utiles en eux-mêmes, peuvent être
utiles à d'autres fonctions sans rapport. L'évolution se fait de façon aveugle et désordonnée, dans laquelle la
fonction d'une forme initiale n'est pas nécessairement la même que celle de la forme finale : c'est le
phénomène d'exaptation (par opposition à l'adaptation). L'enclume dans l'oreille des mammifères (dérivée de
l'os carré dans la mâchoire), et le pouce du panda (dérivé d'une excroissance osseuse du poignet) sont des
exemples classiques.
 Sophismes [modifier]
Les critiques[36] voient la complexité irréductible comme un cas particulier de la thèse « la
complexité est la preuve d'un dessein » (argument du grand Horloger), et la voient donc comme un argument
d'ignorance (sophisme dans lequel on considère comme faux ce qui n'a pas été prouvé), un argument du
« dieu bouche-trous » (« God of the gaps ») et un argument de manque d'imagination[37],[38].
Les critiques considèrent également que l'idée que la complexité irréductible, en tant
fondamentalement qu'argument contre la théorie de l'évolution, impliquerait celle du dessein intelligent
comme un faux dilemme, supposant qu'il n'y a que deux modèles valides[39].
Behe lui même reconnaît que même si les scientifiques n'imaginent pas actuellement comment un
système a évolué vers un état de complexité irréductible, ceci ne prouve pas qu'une évolution n'ait pas pu le
produire. Le scientifique Steve Mirsky l'a décrit comme une « stratégie de reddition intellectuelle totale »[40].

La complexité irréductible au procès de Dover[2] [modifier]

Le prononcé du jugement précise que « le dessein intelligent n'est pas de la science et est
essentiellement de nature religieuse »[4].
Lors de ses témoignages au procès, Behe a admis qu'aucun article publié dans une revue
scientifique à comité de lecture ne soutenait sa thèse selon laquelle des systèmes moléculaires complexes,
comme le flagelle bactérien, la coagulation en cascade et le système immunitaire sont le résultat d'une
cause intelligente, ni que certaines structures moléculaires complexes sont de complexité irréductible[2].
Dans les attendus du jugement, le juge Jones a spécifiquement cité Behe et la complexité irréductible
(Memorandum Opinion)[2] :
 • « Le professeur Behe a admis dans « Reply to My Critics » (Réponse à mes critiques) qu'il y
avait un défaut dans sa thèse de la complexité irréductible, car elle se présente comme un défi à la
sélection naturelle, alors qu'en fait elle ne cherche pas à répondre à l'objectif auquel doit faire face la
sélection naturelle ». Le Professeur Behe a écrit qu'« il espérait résoudre ce défaut dans ses travaux
futurs » (p. 73)
• « Comme le témoignage d'expert l'a mis en évidence, la définition de la « complexité
irréductible » la rend sans signification en tant que critique de l'évolution (3:40 (Miller)). En fait, la
théorie de l'évolution propose l'exaptation comme une explication largement reconnue et bien
documentée de la façon dont les systèmes composés de plusieurs éléments peuvent avoir évolué de
façon naturelle. »(p. 74)
• « Par sa définition de la complexité irréductible, le professeur Behe exclut a priori le
phénomène d'exaptation, ignorant de ce fait de nombreux éléments qui réfutent sa thèse. En
particulier, l'académie des sciences des États-Unis a rejeté la thèse de la complexité irréductible du
Professeur Behe... » (p. 75)
• « Comme la complexité irréductible est uniquement un argument négatif contre l'évolution, il
est réfutable et de ce fait vérifiable, à la différence du dessein intelligent, en montrant des structures
intermédiaires avec des fonctions sélectionnables qui peuvent avoir évolué vers les systèmes de
prétendue complexité irréductible (2:15-16 (Miller)). Cependant, il faut souligner que le fait que
l'argument négatif de la complexité irréductible soit vérifiable ne rend pas vérifiable la thèse du
dessein intelligent (2:15 (Miller); 5:39 (Pennock)). Le professeur Behe n'a appliqué le concept de
complexité irréductible qu'à quelques systèmes choisis : (1) the flagelle bactérien ; (2) la coagulation
en cascade ; et (3) le système immunitaire. Contrairement aux assertions du professeur Behe sur
ces quelques systèmes biochimiques parmi la myriade existant dans la nature, le Dr. Miller a
présenté des travaux publiés dans des revues à comité de lecture, montrant qu'ils ne sont pas en fait
de complexité irréductible. » (p. 76)
• « ...lors des débats contradictoires, le professeur Behe a été interrogé au sujet de ses
déclarations de 1996 qu'une explication évolutionniste du système immunitaire ne serait jamais
trouvée. Il lui a été présenté 58 publications dans des revues à comité de lecture, 9 livres, et
plusieurs chapitres d'ouvrages d'immunologie au sujet de l'évolution du système immunitaire ;
cependant, il a simplement insisté sur le fait que cela ne constituait pas une preuve suffisante de
l'évolution, et que cela n'était pas « assez bon » (23:19 (Behe)). » (p. 78)
• « Nous concluons donc que la thèse du professeur Behe sur la complexité irréductible a été
réfutée dans des publications scientifiques à comité de lecture et rejetée par la communauté
scientifique dans son ensemble (17:45-46 (Padian); 3:99 (Miller)). En outre, même si la complexité
irréductible n'avait pas été rejetée, elle ne soutiendrait pas la thèse du Dessein intelligent, car elle
constitue seulement un test de l'évolution, non d'un « dessein ». (2:15, 2:35-40 (Miller); 28:63-66
(Fuller)).
Considérons maintenant le prétendu « argument en faveur » d'un dessein, impliqué dans
l'expression utilisée à de nombreuses reprises par les professeurs Behe et Minnich lors de leur
témoignage d'expert comme : « un arrangement d'éléments dans un but ». Le professeur Behe a
résumé l'argument comme suit : Nous concluons à un dessein quand nous observons des éléments
qui semblent avoir été arrangés dans un but. La force de la conclusion est quantitative ; plus le
nombre d'éléments est important, plus ils interagissent de façon intriquée, plus notre confiance dans
un dessein est grande. L'évidence d'un dessein dans certains domaines de la biologie est indéniable.
Comme rien, sinon une cause intelligente, n'a été démontré capable de conduire à une telle
évidence de dessein, en dépit des thèses darwiniennes, la conclusion que la structure observée
dans le vivant est un réel dessein est rationnellement justifiée (18:90-91, 18:109-10 (Behe); 37:50
 (Minnich)). Comme indiqué précédemment, cet argument est simplement une reprise de l'argument
du révérend William Paley appliqué à l'échelle cellulaire. Minnich, Behe, et Paley arrivent à la même
conclusion, à savoir que les organismes compliqués doivent avoir été conçus, en utilisant le même
raisonnement, avec la différence que les professeurs Behe et Minnich refusent d'identifier le
concepteur, alors que Paley déduit de l'existence d'une conception qu'il s'agit de Dieu (1:6- 7 (Miller);
38:44, 57 (Minnich)). Le témoignage d'experts a révélé que cet argument inductif n'est pas
scientifique, et comme l'a admis le professeur Behe, ne pourra jamais être exclu (2:40 (Miller);
22:101 (Behe); 3:99 (Miller)). » (p. 79-80)

Notes et références [modifier]

1. ↑ a, b, c et d Michael Behe, « Darwin's Black Box : The Biochemical Challenge to Evolution »,
(1996), cité dans Irreducible Complexity and Michael Behe [archive] (retrieved 8 January 2006)
2. ↑ a, b, c, d et e Juge John E. Jones III : « Attendus, Kitzmiller v. Dover Area School District »
3. ↑ For most members of the mainstream scientific community, ID is not a scientific theory, but
a creationist pseudoscience. David Mu, « Trojan Horse or Legitimate Science: Deconstructing the
Debate over Intelligent Design » [archive]. Harvard Science Review, Vol. 19, Issue 1, Fall 2005. –
Mark D. Decker « Why Intelligent Design Isn't Intelligent » Review of: Unintelligent Design, by Mark
Perakh [archive] . College of Biological Sciences, General Biology Program, University of Minnesota .
4. ↑ a et b Juge John E. Jones III : « Prononcé, Kitzmiller v. Dover Area School District »
5. ↑ Nicolas Malebranche, « De la recherche de la vérité », 6.2.4, 6° édition, (1712)
6. ↑ Andrew Pyle, « Malebranche on Animal Generation: Preexistence and the Microscope »,
dans Justin E.H. Smith, (éd.) « The Problem of Animal Generation in Early Modern Philosophy »
(2006) (ISBN 0-521-84077-5), p. 202-203
 7. ↑ Tom Scharle, (en) The chicken and the egg [archive]
8. ↑ a, b et c Charles Darwin, L'Origine des espèces. John Murray, London, (1859). (en) p.
189, chap. VI [archive]
9. ↑ Hermann J. Muller: « Genetic variability, twin hybrids and constant hybrids, in a case of
balanced lethal factors » , Genetics, 1918, 3, 422-499, (en) spécialement p. 463-464. [archive]
10.↑ Biological Reviews of the Cambridge Philosophical Society , 4(3) 1939, 261-280, p. 272.
11.↑ Ludwig von Bertalanffy (1952). Problems of Life: An Evaluation of Modern Biological and
Scientific Thought, p. 148 ISBN 1-131-79242-4
12.↑ Jacques Monod, Le hasard et la nécessité : Essai sur la philosophie naturelle de la
biologie moderne. Paris, Le Seuil, 1970. ISBN 2020006189
13.↑ William Dembski, No Free Lunch, 285
14.↑ G. Halder, P. Callaerts, et W.J. Gehring, (1995). « New perspectives on eye evolution. »
Curr. Opin. Genet. Dev. 5 (p. 602–609).
15.↑ G. Halder, P. Callaerts, et W.J. Gehring, (1995). « Induction of ectopic eyes by targeted
expression of the eyeless gene in Drosophila ». Science 267 (p. 1788–1792).
16.↑ S.I. Tomarev, P. Callaerts, L. Kos, R. Zinovieva, G. Halder, W. Gehring, et J. Piatigorsky,
« Squid Pax-6 and eye development. » Proc. Natl. Acad. Sci. USA, 94 (1997) (p. 2421–2426).
17.↑ Russell D. Fernald, (2001). « The Evolution of Eyes: Why Do We See What We
See? » [archive] Karger Gazette 64: « The Eye in Focus ».
18.↑ Russell D. Fernald, Aquatic Adaptations in Fish Eyes. New York, Springer, (1998).
19.↑ Russell D. Fernald, (1997). " The evolution of eyes." [archive] Brain Behav Evol. 50 (pp.
253–259).
20.↑ S. Conway-Morris, (1998). The Crucible of Creation. Oxford: Oxford University Press.
 21.↑ a et b A reducibly complex mousetrap [archive] (graphics-intensive, requires JavaScript)
22.↑ Jonathan Sarfati « Argument: 'Irreducible complexity' » [archive], from Refuting Evolution
(Answers in Genesis (2000). ).
23.↑ George Action, « Behe and the Blood Clotting Cascade » [archive]
24.↑ Matt Inlay, 2002. « Evolving Immunity [archive]. » In TalkDesign.org.
25.↑ Nicholas J. Matzke, 2003. « Evolution in (Brownian) space: a model for the origin of the
bacterial flagellum [archive]. »
26.↑ Mark Pallen et Nicholas J. Matzke, 2006, « From The Origin of Species to the origin of
bacterial flagella [archive]. » Nature Reviews Microbiology, (10), 784-790.
27.↑ U. Semba, Y. Shibuya, H. Okabe, T. Yamamoto, « Whale Hageman factor (factor XII):
prevented production due to pseudogene conversion », dans Thromb Res, vol. 90, no 1, 1998, p. 31-
7
28.↑ Kenneth R. Miller, The Flagellum Unspun: The Collapse of « Irreducible
Complexity » [archive] with reply here [archive]
29.↑ Jeremy Manier, Unlocking cell secrets bolsters evolutionists [archive] (Chicago Tribune,
2006 Feb 13)
30.↑ Kenneth R. Miller, The Flagellum Unspun [archive]
31.↑ Ode to the Flagellum [archive] {{en]}} Vidéo montrant la construction évolutive du flagelle
selon Nicholas J. Matzke
32.↑ Niall Shanks, Redundant Complexity:A Critical Analysis of Intelligent Design in
Biochemistry [archive]
33.↑ Niall Shanks and Karl H. Joplin. Redundant Complexity:A Critical Analysis of Intelligent
Design in Biochemistry. [archive] East Tennessee State University.
 34.↑ Lenski RE, C. Ofria, R.T. Pennock, C. Adami, « The evolutionary origin of complex
features », dans Nature, vol. 423, no 6936, 2003, p. 139-44
35.↑ A. G. Cairns-Smith, Seven Clues to the Origin of Life: A Scientific Detective Story ,
Cambridge: Cambridge University Press, 1985. ISBN 0-521-27522-9, p. 39, 58-64.
36.↑ Par exemple Eugenie Scott et Glenn Branch
37.↑ Mark Isaak. Index to Creationist Claims. The Talk. Origins Archive. « Irreducible
complexity and complex specified information are special cases of the complexity indicates design »
claim; they are also arguments from « incredulity. » [1] [archive] « The argument from incredulity
creates a god of the gaps. » [2] [archive]
38.↑ Eugenie C. Scott et Glenn Branch « Intelligent Design » Not Accepted by Most
Scientists [archive], National Center for Science Education website, September 10, 2002.
39.↑ Pete Dunkelberg IC Demystified [archive] makes the point that : if « irreducible
complexity » is tautologically redefined to allow a valid argument that intelligent design is the correct
explanation for life then there is no such thing as  irreducible complexity » in the mechanisms of life;
while, if we use the unmodified original definition then « irreducible complexity » has nothing
whatever to do with evolution.
40.↑ Steve Mirsky, Sticker Shock: In the beginning was the cautionary advisory [archive]
Scientific American, Feb 2005

Bibliographie [modifier]
• Behe, Michael (1996). Darwin's Black Box. New York: The Free Press. ISBN 0-684-83493-6
• Denton, Michael (1986). Evolution: A Theory in Crisis. Adler & Adler.
• Denton, Michael (1997). L'évolution a-t-elle un sens ?. Fayard. ISBN {978-}2-213-59748-5
 • Macnab RM, « Type III flagellar protein export and flagellar assembly », dans Biochim
Biophys Acta, vol. 1694, no 1-3, 2004, p. 207-17
• Ruben, J.A.; Jones, T.D.; Geist, N.R.; & Hillenius, W.J., « Lung Structure and Ventilation in
Theropod Dinosaurs and Early Birds », dans Science, vol. 278, no 5341, 14 novembre 1997, p. 1267–
1270
• Sunderland, Luther D. (March 1976). Miraculous Design in Woodpeckers. Creation
Research Society Quarterly.
• Testing Darwin Discover Magazine Vol. 26 No. 02, février 2005

Annexes [modifier]
Articles connexes [modifier]
• Arbre phylogénétique
• Classification phylogénétique
• Évolution (biologie)
• Fixisme
• Histoire du transformisme
• Historique des critiques des théories de l'évolution
• Holism and Evolution
• Origines de la vie
 • Portail de l’origine et de l’évolution du vivant

• Portail des religions et croyances

Ce document provient de « http://fr.wikipedia.org/wiki/Complexit%C3%A9_irr%C3%A9ductible ».

Catégorie : Créationnisme | [+]

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Specified complexity
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Specified complexity is an argument proposed by William Dembski and used by him and others to
promote intelligent design. According to Dembski, the concept is intended to formalize a property that singles
out patterns that are both specified and complex. Dembski states that specified complexity is a reliable
marker of design by an intelligent agent, a central tenet to intelligent design which Dembski argues for in
opposition to modern evolutionary theory. The concept of specified complexity is widely regarded as
mathematically unsound and has not been the basis for further independent work in information theory,
complexity theory, or biology.[1][2][3] Specified complexity is one of the two main arguments used by
intelligent design proponents, the other being irreducible complexity.
In Dembski's terminology, a specified pattern is one that admits short descriptions, whereas a
complex pattern is one that is unlikely to occur by chance. Dembski argues that it is impossible for specified
complexity to exist in patterns displayed by configurations formed by unguided processes. Therefore,
Dembski argues, the fact that specified complex patterns can be found in living things indicates some kind of
guidance in their formation, which is indicative of intelligence. Dembski further argues that one can rigorously
show by applying no free lunch theorems the inability of evolutionary algorithms to select or generate
configurations of high specified complexity.
In intelligent design literature, an intelligent agent is one that chooses between different possibilities
and has, by supernatural means and methods, caused life to arise.[4] Specified complexity is what Dembski
terms an "explanatory filter" which can recognize design by detecting complex specified information (CSI).
The filter is based on the premise that the categories of regularity, chance, and design are, according to
Dembski, "mutually exclusive and exhaustive." Complex specified information detects design because it
detects what characterizes intelligent agency; it detects the actualization of one among many competing
possibilities.
A study by Wesley Elsberry and Jeffrey Shallit states that "Dembski's work is riddled with
inconsistencies, equivocation, flawed use of mathematics, poor scholarship, and misrepresentation of others'
results".[5] Another objection concerns Dembski's calculation of probabilities. According to Martin Nowak, a
Harvard professor of mathematics and evolutionary biology "We cannot calculate the probability that an eye
came about. We don't have the information to make the calculation".[6] Critics also reject applying specified
complexity to infer design as an argument from ignorance.

Contents
[hide]
• 1 Definition
• 1.1 Orgel's original use
• 1.2 Dembski's definition
• 2 Law of conservation of information
• 3 Specificity
• 3.1 Dembski's explanation of specified complexity
• 3.2 Calculation of specified complexity
• 4 Criticisms
• 5 See also
• 6 Notes and references
• 7 References
• 8 External links
[edit] Definition
[edit] Orgel's original use
The term "specified complexity" was originally coined by origin of life researcher Leslie Orgel to
denote what distinguishes living things from non-living things:
In brief, living organisms are distinguished by their specified complexity. Crystals are usually
taken as the prototypes of simple well-specified structures, because they consist of a very large
number of identical molecules packed together in a uniform way. Lumps of granite or random
mixtures of polymers are examples of structures that are complex but not specified. The crystals
fail to qualify as living because they lack complexity; the mixtures of polymers fail to qualify
because they lack specificity.[7]

The term was later employed by physicist Paul Davies in a similar manner:
Living organisms are mysterious not for their complexity per se, but for their tightly specified
complexity[8]

[edit] Dembski's definition

For Dembski, specified complexity is a property which can be observed in living things. However,
whereas Orgel used the term for that which, in Darwinian theory, is understood to be created through
evolution, Dembski uses it for that which he says cannot be created through "undirected" evolution—and
concludes that it allows one to infer intelligent design. While Orgel employed the concept in a qualitative way,
Dembski's use is intended to be quantitative. Dembski's use of the concept dates to his 1998 monograph
The Design Inference. Specified complexity is fundamental to his approach to intelligent design, and each of
his subsequent books has also dealt significantly with the concept. He has stated that, in his opinion, "if there
is a way to detect design, specified complexity is it."[9]
Dembski asserts that specified complexity is present in a configuration when it can be described by a
pattern that displays a large amount of independently specified information and is also complex, which he
defines as having a low probability of occurrence. He provides the following examples to demonstrate the
concept: "A single letter of the alphabet is specified without being complex. A long sentence of random letters
is complex without being specified. A Shakespearean sonnet is both complex and specified." [10]
In his earlier papers Dembski defined complex specified information (CSI) as being present in a
specified event whose probability did not exceed 1 in 10 150, which he calls the universal probability bound. In
that context, "specified" meant what in later work he called "pre-specified", that is specified before any
information about the outcome is known. The value of the universal probability bound corresponds to the
inverse of the upper limit of "the total number of [possible] specified events throughout cosmic history," as
calculated by Dembski. [11] Anything below this bound has CSI. The terms "specified complexity" and
"complex specified information" are used interchangeably. In more recent papers Dembski has redefined the
universal probability bound, with reference to another number, corresponding to the total number of bit
operations that could possibly have been performed in the entire history of the universe.
Dembski asserts that CSI exists in numerous features of living things, such as DNA and other
functional biological molecules, and argues that it cannot be generated by the only known natural
mechanisms of physical law and chance, or by their combination. He argues that this is so because laws can
only shift around or lose information, but do not produce it, and chance can produce complex unspecified
information, or simple specified information, but not CSI; he provides a mathematical analysis that he claims
demonstrates that law and chance working together cannot generate CSI, either. Moreover, he claims that
CSI is holistic, with the whole being greater than the sum of the parts, and that this decisively eliminates
Darwinian evolution as a possible means of its creation. Dembski maintains that by process of elimination,
CSI is best explained as being due to intelligence, and is therefore a reliable indicator of design.

[edit] Law of conservation of information

Dembski formulates and proposes a law of conservation of information as follows:
This strong proscriptive claim, that natural causes can only transmit CSI but never originate it, I
call the Law of Conservation of Information. Immediate corollaries of the proposed law are the
following:

1. The specified complexity in a closed system of natural causes remains constant or

decreases.
2. The specified complexity cannot be generated spontaneously, originate endogenously
or organize itself (as these terms are used in origins-of-life research).
3. The specified complexity in a closed system of natural causes either has been in the
system eternally or was at some point added exogenously (implying that the system,
though now closed, was not always closed).
4. In particular any closed system of natural causes that is also of finite duration received
whatever specified complexity it contains before it became a closed system.[12]

Dembski notes that the term "Law of Conservation of Information" was previously used by Peter
Medawar in his book The Limits of Science (1984) "to describe the weaker claim that deterministic laws
cannot produce novel information."[13] The actual validity and utility of Dembski's proposed law are
uncertain; it is neither widely used by the scientific community nor cited in mainstream scientific literature. A
2002 essay by Erik Tellgren provided a mathematical rebuttal of Dembski’s law and concludes that it is
"mathematically unsubstantiated." [14]

[edit] Specificity
In a more recent paper,[15] Dembski provides an account which he claims is simpler and adheres
more closely to the theory of statistical hypothesis testing as formulated by Ronald Fisher. In general terms,
Dembski proposes to view design inference as a statistical test to reject a chance hypothesis P on a space of
outcomes Ω.
Dembski's proposed test is based on the Kolmogorov complexity of a pattern T that is exhibited by an
event E that has occurred. Mathematically, E is a subset of Ω, the pattern T specifies a set of outcomes in Ω
and E is a subset of T. Quoting Dembski[16]
Thus, the event E might be a die toss that lands six and T might be the composite event
consisting of all die tosses that land on an even face.

Kolmogorov complexity provides a measure of the computational resources needed to specify a

pattern (such as a DNA sequence or a sequence of alphabetic characters).[17] Given a pattern T, the
number of other patterns may have Kolmogorov complexity no larger than that of T is denoted by φ(T). The
number φ(T) thus provides a ranking of patterns from the simplest to the most complex. For example, for a
pattern T which describes the bacterial flagellum, Dembski claims to obtain the upper bound φ(T) ≤ 1020.
Dembski defines specified complexity of the pattern T under the chance hypothesis P as
 where P(T) is the probability of observing the pattern T, R is the number of "replicational resources"
available "to witnessing agents". R corresponds roughly to repeated attempts to create and discern a pattern.
Dembski then asserts that R can be bounded by 10120. This number is supposedly justified by a result of
Seth Lloyd[18] in which he determines that the number of elementary logic operations that can have
performed in the universe over its entire history cannot exceed 10 120 operations on 1090 bits.
Dembski's main claim is that the following test can be used to infer design for a configuration: There
is a target pattern T that applies to the configuration and whose specified complexity exceeds 1. This
condition can be restated as the inequality

[edit] Dembski's explanation of specified complexity

Dembski's expression σ is unrelated to any known concept in information theory, though he claims
he can justify its relevance as follows: An intelligent agent S witnesses an event E and assigns it to some
reference class of events Ω and within this reference class considers it as satisfying a specification T. Now
consider the quantity φ(T) × P(T) (where P is the "chance" hypothesis):
 Possible targets with complexity ranking and probability not exceeding those of attained target T.
Probability of set-theoretic union does not exceed φ(T) × P(T)
Think of S as trying to determine whether an archer, who has just shot an arrow at a large wall,
happened to hit a tiny target on that wall by chance. The arrow, let us say, is indeed sticking
squarely in this tiny target. The problem, however, is that there are lots of other tiny targets on
the wall. Once all those other targets are factored in, is it still unlikely that the archer could have
hit any of them by chance?
 In addition, we need to factor in what I call the replicational resources associated with T, that is,
all the opportunities to bring about an event of T's descriptive complexity and improbability by
multiple agents witnessing multiple events.

According to Dembski, the number of such "replicational resources" can be bounded by "the maximal
number of bit operations that the known, observable universe could have performed throughout its entire
multi-billion year history", which according to Lloyd is 10 120.
However, according to Elsberry and Shallit, "[specified complexity] has not been defined formally in
any reputable peer-reviewed mathematical journal, nor (to the best of our knowledge) adopted by any
researcher in information theory."[19]

[edit] Calculation of specified complexity

Thus far, Dembski's only attempt at calculating the specified complexity of a naturally occurring
biological structure is in his book No Free Lunch, for the bacterial flagellum of E. coli. This structure can be
described by the pattern "bidirectional rotary motor-driven propeller". Dembski estimates that there are at
most 1020 patterns described by four basic concepts or fewer, and so his test for design will apply if

However, Dembski says that the precise calculation of the relevant probability "has yet to be done",
although he also claims that some methods for calculating these probabilities "are now in place".
 These methods assume that all of the constituent parts of the flagellum must have been generated
completely at random, a scenario that biologists do not seriously consider. He justifies this approach by
appealing to Michael Behe's concept of "irreducible complexity" (IC), which leads him to assume that the
flagellum could not come about by any gradual or step-wise process. The validity of Dembski's particular
calculation is thus wholly dependent on Behe's IC concept, and therefore susceptible to its criticisms, of
which there are many.
To arrive at the ranking upper bound of 1020 patterns, Dembski considers a specification pattern for
the flagellum defined by the (natural language) predicate "bidirectional rotary motor-driven propeller", which
he regards as being determined by four independently chosen basic concepts. He furthermore assumes that
English has the capability to express at most 105 basic concepts (an upper bound on the size of a dictionary).
Dembski then claims that we can obtain the rough upper bound of

for the set of patterns described by four basic concepts or fewer.

From the standpoint of Kolmogorov complexity theory, this calculation is problematic. Quoting
Ellsberry and Shallit[20] "Natural language specification without restriction, as Dembski tacitly permits,
seems problematic. For one thing, it results in the Berry paradox". These authors add: "We have no objection
to natural language specifications per se, provided there is some evident way to translate them to Dembski's
formal framework. But what, precisely, is the space of events Ω here?"
[edit] Criticisms
The soundness of Dembski's concept of specified complexity and the validity of arguments based on
this concept are widely disputed. A frequent criticism (see Elsberry and Shallit) is that Dembski has used the
terms "complexity", "information" and "improbability" interchangeably. These numbers measure properties of
things of different types: Complexity measures how hard it is to describe an object (such as a bitstring),
information measures how close to uniform a random probability distribution is and improbability measures
how unlikely an event is given a probability distribution.
When Dembski's mathematical claims on specific complexity are interpreted to make them
meaningful and conform to minimal standards of mathematical usage, they usually turn out to be false.
Dembski often sidesteps these criticisms by responding that he is not "in the business of offering a strict
mathematical proof for the inability of material mechanisms to generate specified complexity".[21] Yet on
page 150 of No Free Lunch he claims he can prove his thesis mathematically: "In this section I will present
an in-principle mathematical argument for why natural causes are incapable of generating complex specified
information." Others have pointed out that a crucial calculation on page 297 of No Free Lunch is off by a
factor of approximately 1065.[22]
Dembski's calculations show how a simple smooth function cannot gain information. He therefore
concludes that there must be a designer to obtain CSI. However, natural selection has a branching mapping
from one to many (replication) followed by pruning mapping of the many back down to a few (selection).
When information is replicated, some copies can be differently modified while others remain the same,
allowing information to increase. These increasing and reductional mappings were not modeled by Dembski.
In other words, Dembski's calculations do not model birth and death. This basic flaw in his modeling renders
all of Dembski's subsequent calculations and reasoning in No Free Lunch irrelevant because his basic model
does not reflect reality. Since the basis of No Free Lunch relies on this flawed argument, the entire thesis of
the book collapses.[23]
According to Martin Nowak, a Harvard professor of mathematics and evolutionary biology "We
cannot calculate the probability that an eye came about. We don't have the information to make the
calculation".[6]
Dembski's critics note that specified complexity, as originally defined by Leslie Orgel, is precisely
what Darwinian evolution is supposed to create. Critics maintain that Dembski uses "complex" as most
people would use "absurdly improbable". They also claim that his argument is a tautology: CSI cannot occur
naturally because Dembski has defined it thus. They argue that to successfully demonstrate the existence of
CSI, it would be necessary to show that some biological feature undoubtedly has an extremely low probability
of occurring by any natural means whatsoever, something which Dembski and others have almost never
attempted to do. Such calculations depend on the accurate assessment of numerous contributing
probabilities, the determination of which is often necessarily subjective. Hence, CSI can at most provide a
"very high probability", but not absolute certainty.
Another criticism refers to the problem of "arbitrary but specific outcomes". For example, if a coin is
tossed randomly 1000 times, the probability of any particular outcome occurring is roughly one in 10 300. For
any particular specific outcome of the coin-tossing process, the a priori probability that this pattern occurred
is thus one in 10300, which is astronomically smaller than Dembski's universal probability bound of one in
10150. Yet we know that the post hoc probability of its happening is exactly one, since we observed it
happening. This is similar to the observation that it is unlikely that any given person will win a lottery, but,
eventually, a lottery will have a winner; to argue that it is very unlikely that any one player would win is not the
same as proving that there is the same chance that no one will win. Similarly, it has been argued that "a
space of possibilities is merely being explored, and we, as pattern-seeking animals, are merely imposing
patterns, and therefore targets, after the fact."[24]
Apart from such theoretical considerations, critics cite reports of evidence of the kind of evolutionary
"spontanteous generation" that Dembski claims is too improbable to occur naturally. For example, in 1982,
B.G. Hall published research demonstrating that after removing a gene that allows sugar digestion in certain
bacteria, those bacteria, when grown in media rich in sugar, rapidly evolve new sugar-digesting enzymes to
replace those removed.[25] Another widely cited example is the discovery of nylon eating bacteria that
produce enzymes only useful for digesting synthetic materials that did not exist prior to the invention of nylon
in 1935.
Other commentators have noted that evolution through selection is frequently used to design certain
electronic, aeronautic and automotive systems which are considered problems too complex for human
"intelligent designers"[26]. This strongly contradicts the argument that an intelligent designer is required for
the most complex systems. Such evolutionary techniques can lead to designs that are difficult to understand
or evaluate since no human understands which trade-offs were made in the evolutionary process, something
which mimics our poor understanding of biological systems.
Dembski's book No Free Lunch was criticised for not addressing the work of researchers who use
computer simulations to investigate artificial life. According to Jeffrey Shallit:
The field of artificial life evidently poses a significant challenge to Dembski's claims about the
failure of evolutionary algorithms to generate complexity. Indeed, artificial life researchers
regularly find their simulations of evolution producing the sorts of novelties and increased
complexity that Dembski claims are impossible.[22]
[edit] See also
• Evolution
• Intelligent design
• List of topics characterized as pseudoscience
• Texas sharpshooter fallacy

[edit] Notes and references

1. ^ Rich Baldwin (2005). "Information Theory and Creationism: William Dembski". TalkOrigins
Archive. http://www.talkorigins.org/faqs/information/dembski.html. Retrieved 2010-05-10.
2. ^ Mark Perakh, (2005). Dembski "displaces Darwinism" mathematically -- or does he?
3. ^ Jason Rosenhouse, (2001). How Anti-Evolutionists Abuse Mathematics The Mathematical
Intelligencer, Vol. 23, No. 4, Fall 2001, pp. 3-8.
4. ^ "no intelligent agent who is strictly physical could have presided over the origin of the
universe or the origin of life." - Dembski The Act of Creation: Bridging Transcendence and
Immanence Dembski has also said that "intelligent design should be understood as the evidence that
God has placed in nature to show that the physical world is the product of intelligence and not simply
the result of mindless material forces" Why President Bush Got It Right about Intelligent Design
William A. Dembski DesignInference.com, August 4 2005. Dembski has also said that "Intelligent
design is just the Logos theology of John's Gospel restated in the idiom of information theory." Signs
of Intelligence A Primer on the Discernment of Intelligent Design. William A. Dembski. Touchstone
Journal, Volume 12, Issue 4, July/August 1999.
5. ^ Wesley Elsberry and Jeffrey Shallit, (2003). Information Theory, Evolutionary Computation,
and Dembski’s “Complex Specified Information”
 6. ^ a b Martin Nowak (2005). Time Magazine, 15 August 2005, page 32
7. ^ Leslie Orgel (1973). The Origins of Life, p. 189.
8. ^ Paul Davies (1999). The Fifth Miracle p. 112
9. ^ William A. Dembski (2002). No Free Lunch, p. 19.
10.^ William A. Dembski (1999). Intelligent Design, p. 47.
11.^ William A. Dembski (2004). The Design Revolution: Answering the Toughest Questions
About Intelligent Design, p. 85.
12.^ William A. Dembski (1998) Intelligent Design as a Theory of Information.
13.^ "Searching Large Spaces: Displacement and the No Free Lunch Regress (356k PDF)", pp.
15-16, describing an argument made by Michael Shermer in How We Believe: Science, Skepticism,
and the Search for God, 2nd ed. (2003).
14.^ On Dembski's law of conservation of information Erik Tellgren. talkreason.org, 2002. (PDF
file)
15.^ William A. Dembski (2005). Specification: The Pattern that Signifies intelligence
16.^ (loc. cit. p 16)
17.^ Michael Sipser (1997). Introduction to the Theory of Computation, PWS Publishing
Company.
18.^ Seth Lloyd (2002), Computational capacity of the universe, Phys. Rev. Lett. 88(23):790 1-
4. See also arXiv:quant/ph0110141.
19.^ Elsberry and Shallit, (2003)p 14
20.^ Elsberry and Shallit, (2003)p 12
21.^ William A. Dembski, (2002). If Only Darwinists Scrutinized Their Own Work as Closely: A
Response to "Erik".
22.^ a b Jeffrey Shallit (2002) A review of Dembski's No Free Lunch
 23.^ Thomas D. Schneider. (2002) Dissecting Dembski's "Complex Specified Information"
24.^ William A. Dembski (1998) Intelligent Design as a Theory of Information.
25.^ B.G. Hall (1982). "Evolution of a regulated operon in the laboratory", Genetics, 101(3-
4):335-44. In PubMed.
26.^ Evolutionary algorithms now surpass human designers New Scientist, 28 July 2007

[edit] References
• Wesley Elsberry and Jeffrey Shallit, (2003). Information Theory, Evolutionary Computation,
and Dembski’s “Complex Specified Information”

[edit] External links

• Not a Free Lunch But a Box of Chocolates - A critique of William Dembski's book No Free
Lunch by Richard Wein, from TalkOrigins
• Information Theory and Creationism William Dembski by Rich Baldwin, from Information
Theory and Creationism, compiled by Ian Musgrave and Rich Baldwin
• Critique of No Free Lunch by H. Allen Orr from the Boston Review
• Dissecting Dembski's "Complex Specified Information" by Dr. Thomas D. Schneider.
• William Dembski's treatment of the No Free Lunch theorems is written in jello by No Free
Lunch theorems co-founder, David Wolpert
• The Evolution List - Genetic ID and the Explanatory Filter by Allen MacNeill.
• Design Inference Website - The writing of William A. Dembski
 • Committee for Skeptical Inquiry - Reality Check, The Emperor's New Designer Clothes -
Victor J. Stenger
• Darwin@Home Web site - open-source software that demonstrates evolution in artificial life,
written by Gerald de Jong

[hide]
v•d•e
Creationism topics

Types Young Earth Creationism · Old Earth creationism · Progressive

creationism · Theistic evolution · Intelligent design

Genesis Genesis creation narrative · Framework interpretation · Day-Age theory ·

interpretations Gap theory · Biblical literalism

Related concepts Creation myth · Omphalos hypothesis · Specified complexity ·

Irreducible complexity · Dating Creation · Theistic realism · Intelligent designer

Pseudoscience Creation science · Creationist cosmologies · Creation biology · Created

kinds · Flood geology · Vapor canopy · Modern geocentrism

Miscellanea Creation-evolution controversy · History of creationism · Creation and

evolution in public education · Quotes about creation and evolution · Teach the
 Controversy

Retrieved from "http://en.wikipedia.org/wiki/Specified_complexity"

Categories: Intelligent design | Creationist objections to evolution

 W000

Fine-tuned Universe
From Wikipedia, the free encyclopedia

Jump to: navigation, search

The fine-tuned Universe is the idea that the conditions that allow life in the Universe can only occur
when certain universal fundamental physical constants lie within a very narrow range, so that if any of several
fundamental constants were only slightly different the universe would be unlikely to be conducive to the
establishment and development of matter, astronomical structures, elemental diversity, or life as it is
presently understood.[1]
 The existence and extent of fine-tuning in the universe is a matter of dispute in the scientific
community. Proponents of fine-tuning include physicist Paul Davies who has stated "There is now broad
agreement among physicists and cosmologists that the universe is in several respects ‘fine-tuned' for life"[2].
Other physicists such as Victor Stenger dispute fine-tuning, saying that even though "life as we know it would
not exist if any one of several of the constants of physics were just slightly different, [we] cannot prove that
some other form of life is feasible with a different set of constants. Anyone who insists that our form of life is
the only one conceivable is making a claim based on no evidence and no theory."[3] Among scientists who
find the evidence persuasive, a variety of scientific explanations have been proposed, e.g., the anthropic
principle along with multiple universes. The idea has also attracted discussion among philosophers and
theologians, as well as creationists and proponents of the Intelligent Design movement.
Contents
[hide]
• 1 Premise
• 2 Disputes on the extent and existence of fine-tuning
• 3 Possible scientific explanations
• 3.1 Multiverse
• 3.1.1 Bubble universe theory
• 3.1.2 Top-down cosmology
• 3.2 Alien design
• 4 Religious opinions
• 4.1 Intelligent design
• 4.2 Other religious creation views
• 4.3 Counter argument to religious views
• 5 In fiction and popular culture
• 6 See also
• 7 References
• 8 Further reading
• 9 External links
[edit] Premise
The neutrality of this section is disputed. Please see the discussion on the talk
page. Please do not remove this message until the dispute is resolved. (June 2010)

Fine-tuned Universe proponents argue that deep-space structures such as the Eta Carinae Nebula
would not form in a universe with significantly different physical constants. Photo: HST / NASA / ESA.
The premise of the fine-tuned universe assertion is that a small change in several of the
dimensionless fundamental physical constants would make the universe radically different. As Stephen
Hawking has noted, "The laws of science, as we know them at present, contain many fundamental numbers,
like the size of the electric charge of the electron and the ratio of the masses of the proton and the
electron. ... The remarkable fact is that the values of these numbers seem to have been very finely adjusted
to make possible the development of life."[4]
If, for example, the strong nuclear force were 2% stronger than it is (i.e., if the coupling constant
representing its strength were 2% larger), while the other constants were left unchanged, diprotons would be
stable and hydrogen would fuse into them instead of deuterium and helium. This would drastically alter the
physics of stars, and presumably preclude the existence of life similar to what we observe on Earth.
However, many of the fundamental constants describe the properties of the unstable strange, charmed,
bottom and top quarks and mu and tau leptons which seem to play little part in the universe or the structure
of matter.
The precise formulation of the idea is made difficult by the fact that physicists do not yet know how
many independent physical constants there are. The standard model of particle physics has about 26 freely
adjustable parameters. However, because the standard model is not mathematically self-consistent under
certain conditions (e.g., at very high energies, at which both quantum mechanics and general relativity are
relevant), physicists believe that it is underlaid by some other theory, such as a grand unified theory, string
theory, or loop quantum gravity. In some candidate theories, the actual number of independent physical
constants may be as small as 1. For example, the cosmological constant may be a fundamental constant, but
attempts have also been made to calculate it from other constants, and according to the author of one such
calculation, "the small value of the cosmological constant is telling us that a remarkably precise and totally
unexpected relation exists among all the parameters of the Standard Model of particle physics, the bare
cosmological constant and unknown physics."[5]
Martin Rees[6] formulates the fine-tuning of the universe in terms of the following six dimensionless
constants:
 • N = ratio of the strength of electromagnetism to that of gravity;
• Epsilon (ε) = strength of the force binding nucleons into nuclei;
• Omega (ω) = relative importance of gravity and expansion energy in the universe;
• Lambda (λ) = cosmological constant;
• Q = ratio of the gravitational energy required to pull a large galaxy apart to the energy
equivalent of its mass;
• D = number of spatial dimensions in spacetime.

[edit] Disputes on the extent and existence of fine-tuning

Computer simulations suggest that not all of the purportedly "fine-tuned" parameters may be as fine-
tuned as has been claimed. Victor Stenger has simulated different universes in which four fundamental
parameters are varied (electromagnetic interaction, strong nuclear interaction, electron mass, and proton
mass; tuning of other parameters such as the cosmological constant was not considered). He found that
long-lived stars could exist over a wide parameter range, and concluded that "... a wide variation of constants
of physics leads to universes that are long-lived enough for life to evolve, although human life need not exist
in such universes".[7] However Stenger's work has been heavily criticised as having several fundamental
flaws by other physicists[8] [9].
Fred Adams has done a similar study to Stenger, investigating the structure of stars in universes with
different values of the gravitational constant G, the fine-structure constant α, and a nuclear reaction rate
parameter C. His study suggests that roughly 25% of this parameter space allows stars to exist, but not
necessarily to form. His analysis also does not take into account variations of other fundamental constants or
considerations of habitability.[10] Adams work has also been criticised for making incorrect assumptions[11] .
Harnik, Kribs and Perez have argued for the viability of a universe with no weak interaction at all. However,
they noted that their analysis does not extend to the supposed fine tuning of the cosmological constant, and
concluded that "the fine-tuning problems associated with the electroweak breaking scale and the
cosmological constant appear to be qualitatively different from the perspective of obtaining a habitable
universe."[12]
The validity of fine tuning examples is sometimes questioned on the grounds that such reasoning is
subjective anthropomorphism applied to natural physical constants. Critics also suggest that the fine-tuned
universe assertion and the anthropic principle are essentially tautologies.[13] The fine-tuned universe
argument has also been criticized as an argument by lack of imagination because it assumes no other forms
of life, sometimes referred to as carbon chauvinism. Conceptually, alternative biochemistry or other forms of
life are possible [14]. In addition, critics argue that humans are adapted to the universe through the process
of evolution, rather than the universe being adapted to humans (see puddle thinking). They also see it as an
example of the logical flaw of hubris or anthropocentrism in its assertion that humans are the purpose of the
universe.[15]

[edit] Possible scientific explanations

If it is accepted that the universe is fine-tuned, there are a number of scientific explanations that
attempt to account for it.
There are fine tuning arguments that are naturalistic [16]. As modern cosmology developed, various
hypotheses have been proposed. One is an oscillatory universe or a multiverse where physical constants are
postulated to resolve themselves to random values in different iterations of reality.[17] Under this hypothesis,
separate parts of reality would have wildly different characteristics. In such scenarios the issue of fine-tuning
does not arise at all, as only those "universes" with constants hospitable to life (such as what we observe)
would develop life capable of asking the question.
Based upon the Anthropic principle, physicist Robert H. Dicke proposed the "Dicke coincidence"
argument that the structure (age, physical constants, etc.) of the universe as seen by living observers is not
random, but is constrained by biological factors that require it to be roughly a "golden age".[18]

[edit] Multiverse
Main article: Multiverse
The Multiverse hypothesis assumes the existence of many universes with different physical
constants, some of which are hospitable to intelligent life (see multiverse: anthropic principle). Because we
are intelligent beings, we are by definition in a hospitable one. Mathematician Michael Ikeda and astronomer
William H. Jefferys have argued that the anthropic principle resolves the entire issue of fine-tuning,[19][20] as
does philosopher of science Elliott Sober.[21] Philosopher and theologian Richard Swinburne reaches the
opposite conclusion using Bayesian probability.[22]
This approach has led to considerable research into the anthropic principle and has been of
particular interest to particle physicists because theories of everything do apparently generate large numbers
of universes in which the physical constants vary widely. As of yet, there is no evidence for the existence of a
multiverse, but some versions of the theory do make predictions which some researchers studying M-theory
and gravity leaks hope to see some evidence of soon.[23] The existence of additional universes in a
multiverse, other than the observable universe, is not falsifiable, and thus some are reluctant to call the
multiverse idea a "scientific" idea. UNC-Chapel Hill professor Laura Mersini-Houghton claims that the WMAP
cold spot may provide testable empirical evidence for a parallel universe.
 Variants on this approach include Lee Smolin's notion of cosmological natural selection, the
Ekpyrotic universe, and the Bubble universe theory.
Critics of the multiverse-related explanations argue that there is no evidence that other universes
exist.

[edit] Bubble universe theory

Main article: Chaotic inflation theory
See also: Multiverse#Bubble theory
The bubble universe model by physicist Andrei Linde, postulates that our universe is one of many
that grew from a multiverse consisting of vacuum that had not yet decayed to its ground state.
According to this scenario, by means of a random quantum fluctuation the universe "tunneled"
from pure vacuum ("nothing") to what is called a false vacuum, a region of space that contains
no matter or radiation but is not quite "nothing." The space inside this bubble of false vacuum
was curved, or warped. A small amount of energy was contained in that curvature, somewhat
like the energy stored in a strung bow. This ostensible violation of energy conservation is
allowed by the Heisenberg uncertainty principle for sufficiently small time intervals.

The bubble then inflated exponentially and the universe grew by many orders of magnitude in a
tiny fraction of a second. (For a not-too-technical discussion, see Stenger 1990). As the bubble
expanded, its curvature energy was converted into matter and radiation, inflation stopped, and
the more linear big bang expansion we now experience commenced. The universe cooled and
 its structure spontaneously froze out, as formless water vapor freezes into snowflakes whose
unique patterns arise from a combination of symmetry and randomness.

—Victor J. Stenger, The Anthropic Coincidences[24]

In standard inflation, inflationary expansion occurred while the universe was in a false vacuum state,
halting when the universe decayed to a true vacuum state. The bubble universe model proposes that
different parts of this inflationary universe (termed a Multiverse) decayed at different times, with decaying
regions corresponding to universes not in causal contact with each other. It further supposes that each
bubble universe may have different physical constants.

[edit] Top-down cosmology

Stephen Hawking, along with Thomas Hertog of CERN, proposed that the universe's initial conditions
consisted of a superposition of many possible initial conditions, only a small fraction of which contributed to
the conditions we see today.[25] According to their theory, it is inevitable that we find our universe's "fine-
tuned" physical constants, as the current universe "selects" only those past histories that led to the present
conditions. In this way, top-down cosmology provides an anthropic explanation for why we find ourselves in a
universe that allows matter and life, without invoking the current existence of a multiverse.

[edit] Alien design

One hypothesis is that the Universe may have been designed by an alien or by aliens. This would
solve the problem of how a designer or design team capable of fine-tuning the Universe could come to exist.
Cosmologist Alan Guth believes humans will in time be able to generate new universes. By implication
previous intelligent entities may have generated our universe.[26] This idea leads to the possibility that the
extraterrestrial designer/designers are themselves the product of an evolutionary process in their own
universe, which must therefore itself be able to sustain life. For instance, Richard Dawkins maintains that an
alien designer or designers are more plausible than a supernatural designer or designers because there is a
known mechanism to produce them. He calls it the “crane” of Natural selection. Dawkins claims, though, are
criticized among philosophers[who?] to just push back the problem once (now it would be no more the case
to explain this universe, but the universe in which those aliens live), and it could be argued that the resulting
universe where the aliens live calls even more for a designer that would be eternal and uncreated, such as
God. Further, in Richard Dawkins' ultimate Boeing 747 gambit he explains that evolution is an even more
plausible "crane".
The Simulation hypothesis promoted by Nick Bostrom and others suggests that our universe may be
a computer simulation by aliens.[27]
The Biocosm hypothesis and the Meduso-anthropic principle both suggest that natural selection has
made the universe biophilic. The universe enables intelligence because intelligent entities later create new
biophilic universes. This is different from the suggestion above that aliens from a universe which is less finely
tuned than ours made our universe finely tuned.

[edit] Religious opinions

Part of a series of articles on

Intelligent design
 Concepts

Irreducible complexity
Specified complexity
Fine-tuned universe
Intelligent designer
Theistic realism
Neo-creationism

Intelligent design
movement
 Timeline
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Center for Science and Culture
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Intelligent design in politics
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As with theistic evolution, some individual scientists, theologians, and philosophers as well as certain
religious groups argue that providence or creation are responsible for fine-tuning.
 Christian philosopher Alvin Plantinga argues that random chance, applied to a single and sole
universe, only raises the question as to why this universe could be so "lucky" as to have precise conditions
that support life at least at some place (the Earth) and time (within millions of years of the present).[28]
This apparent fine-tuning of the universe is cited[29] by theologian William Lane Craig as an
evidence for the existence of God or some form of intelligence capable of manipulating (or designing) the
basic physics that governs the universe. Craig argues, however, "that the postulate of a divine Designer does
not settle for us the religious question."
Variants on this approach include:

[edit] Intelligent design

Proponents of Intelligent design argue that certain features of the universe and of living things are
best explained by an intelligent cause, not an undirected process such as natural selection. The fine-tuned
universe argument is a central premise or presented as a given in many of the published works of prominent
Intelligent Design proponents, such as William A. Dembski and Michael Behe.

[edit] Other religious creation views

Most religions have some kind of account of the creation of the universe, although they generally
differ in detail from the ones listed above. Some of these may be compatible with known scientific facts. For
example scientist-theologians such as John Polkinghorne emphasize the implications of Anthropic Fine-
Tuning within an orthodox Christian framework whilst fully accepting the scientific findings about Evolution
and the age of the Universe. This is also the position of the Roman Catholic Church and of most Anglican
theologians.[30] The Jewish physicist Gerald Schroeder argues that the apparent discrepancy between the
"days" in Genesis and the billions of years in a scientific understanding are due to the differences in frames of
reference. Many other religious creation views are either incompatible with, or indifferent to, scientific
understandings. Other scientists with similar views are physicist Freeman Dyson and astronomer Owen
Gingerich.

[edit] Counter argument to religious views

Victor Stenger argues that "... The fine-tuning argument and other recent intelligent design
arguments are modern versions of God-of-the-gaps reasoning, where a God is deemed necessary whenever
science has not fully explained some phenomenon".[7]
The "argument from imperfection" suggests that if the universe were designed to be fine-tuned for
life, it should be the best one possible and that evidence suggests that it is not.[31] In fact, most of the
universe is highly hostile to life.

[edit] In fiction and popular culture

This "In popular culture" section may contain minor or trivial references. Please
reorganize this content to explain the subject's impact on popular culture rather than simply
listing appearances, and remove trivial references. (January 2010)
• Stephen Baxter has written several novels and short stories in which the setting is an
alternative universe with different physical laws.
• Robert J. Sawyer discusses the fine-tuned universe at length in his novel Calculating God
(2000).
 • The Sphere Builders of Star Trek: Enterprise created the Delphic Expanse in an attempt to
alter the physical laws of our universe to match those of their own parallel universe.
• Author Neal Stephenson discussed the issue of fine-tuning in the conclusion to his essay In
the Beginning... was the Command Line.[32]
• L. Ron Hubbard in his novel Battlefield Earth (1982) introduced an invasion force whose
home world was in fact located in another Universe. Many of the differences between the two
universes formed the basis for plot points.
• Puddle thinking is an analogy coined by Douglas Adams to satirize the Fine-tuned Universe
argument for supernatural creationism.[33][34] As quoted in Richard Dawkins' eulogy for Douglas
Adams:[35]
. . . imagine a puddle waking up one morning and thinking, 'This is an interesting world I find
myself in, an interesting hole I find myself in, fits me rather neatly, doesn't it? In fact it fits me
staggeringly well, must have been made to have me in it!' This is such a powerful idea that as
the sun rises in the sky and the air heats up and as, gradually, the puddle gets smaller and
smaller, it's still frantically hanging on to the notion that everything's going to be all right,
because this world was meant to have him in it, was built to have him in it; so the moment he
disappears catches him rather by surprise. I think this may be something we need to be on the
watch out for.

[edit] See also

• Fine-tuning
• Origin of life
 • Puddle thinking
• Rare Earth hypothesis
• Teleology
• Ultimate fate of the universe

[edit] References
1. ^ Mark Isaak (ed.) (2005). "CI301: The Anthropic Principle". Index to Creationist Claims.
TalkOrigins Archive. http://www.talkorigins.org/indexcc/CI/CI301.html. Retrieved 2007-10-31.
2. ^ Paul Davies, "How bio-friendly is the universe?" International Journal of Astrobiology, vol.
2, no. 2 (2003): 115.
3. ^ Victor Stenger,[1] "The Uncreated Universe" Has Science Found God?: The Latest Results
in the Search for Purpose in the Universe, Prometheus Books, ISBN 1591020182.
4. ^ Stephen Hawking, 1988. A Brief History of Time, Bantam Books, ISBN 0-553-05340-X, p.
125.
5. ^ Larry Abbott, "The Mystery of the Cosmological Constant," Scientific American, vol. 3, no. 1
(1991): 78.
6. ^ Martin Rees, 1999. Just Six Numbers, HarperCollins Publishers, ISBN 0-465-03672-4.
7. ^ a b Is The Universe Fine-Tuned For Us? Victor J. Stenger, University of Colorado.
8. ^ Luke Barnes (2010). "What Chances Me? A Fine Tuned Critique of Victor Stenger (Part 1)".
Letters to Nature. Letters to Nature. http://letterstonature.wordpress.com/2010/04/11/what-chances-
me-a-fine-tuned-critique-of-victor-stenger-part-1/. Retrieved 2010-09-03.
 9. ^ Luke Barnes (2010). "No Faith In MonkeyGod: A Fine-Tuned Critique of Victor Stenger
(Part 2)". Letters to Nature. Letters to Nature. http://letterstonature.wordpress.com/2010/04/18/no-
faith-in-monkeygod-a-fine-tuned-critique-of-victor-stenger-part-2/. Retrieved 2010-09-03.
10.^ Adams, F.C. (2008). "Stars in other universes: stellar structure with different fundamental
constants". Journal of Cosmology and Astroparticle Physics 2008 (08): 010. doi:10.1088/1475-
7516/2008/08/010. http://arxiv.org/abs/0807.3697.
11.^ Luke Barnes (2010). "The Shrinking Quarter: A Fine-Tuned Critique of Fred Adams".
Letters to Nature. Letters to Nature. http://letterstonature.wordpress.com/2010/02/28/the-shrinking-
quarter-a-fine-tuned-critique-of-fred-adams/. Retrieved 2010-09-03.
12.^ Harnik, R.; Kribs, G.D. and Perez, G. (2006). "A universe without weak interactions".
Physical Review D 74: 035006. doi:10.1103/PhysRevD.74.035006. http://arxiv.org/abs/hep-
ph/0604027.
13.^ See, e.g., Our place in the Multiverse Joseph Silk. Nature, Volume 443 Number 7108,
September 14, 2006.
14.^ See, e.g. Jack Cohen and Ian Stewart: What Does a Martian Look Like: The Science of
Extraterrestrial Life, Wiley, 2002
15.^ See, e.g., Gerald Feinberg and Robert Shapiro, "A Puddlian Fable" in Huchingson,
Religion and the Natural Sciences (1993), pp. 220-221
16.^ L. Susskind, The cosmic landscape: string theory and the illusion of intelligent design .
(Little, Brown, 2005)
17.^ Wheeler, J. A. (1977) in Foundational problems in the special sciences, Reidel, Dordrecht,
pp 3–33
18.^ Dicke, R. H. (1961). "Dirac's Cosmology and Mach's Principle". Nature 192: 440–441.
doi:10.1038/192440a0.
 19.^ The Anthropic Principle Does Not Support Supernaturalism, Michael Ikeda, Bill Jefferys
20.^ Michael Ikeda and William H. Jefferys, "The Anthropic Principle Does Not Support
Supernaturalism," in The Improbability of God, Michael Martin and Ricki Monnier, Editors, pp. 150-
166. Amherst, N.Y.: Prometheus Press. ISBN 1-59102-381-5.
21.^ Elliott Sober, 2004. The Design Argument, in The Blackwell Guide to the Philosophy of
Religion, W. E. Mann, Editor. Blackwell Publishing, ISBN 0-631-22129-8.
22.^ Richard Swinburne, 1990. Argument from the fine-tuning of the universe , in Physical
cosmology and philosophy, J. Leslie, Editor. Collier Macmillan: New York. pp. 154-73.
23.^ Parallel Worlds,2005, Michio Kaku, pp. 220-221
24.^ The Anthropic Coincidences
25.^ Ball, Philip (June 21, 2006). "Hawking Rewrites History...Backwards". Nature News Online.
http://www.nature.com/news/2006/060619/full/news060619-6.html. Retrieved April 19, 2010.
26.^ BBC - Science & Nature - Horizon - Parallel Universes - Transcript
27.^ Bostrom, N. (2002). Anthropic Bias: Observation Selection Effects in Science and
Philosophy. Routledge, New York. ISBN 0-415-93858-9.
28.^ Alvin Plantinga, Books & Culture, March/April 2007 Issue
29.^ William Lane Craig, "The Teleological Argument and the Anthropic Principle," [2]
30.^ See, e.g., Alister McGrath's books Scientific Theology and The Science of God.
31.^ Avitel Pilpel, SKEPTIC, November 2007 Issue, p.18
32.^ In The Beginning Was The Command Line
33.^ Williams, Robyn (18). "The anthropic universe". The Science Show (ABC Radio National).
http://www.abc.net.au/rn/scienceshow/stories/2006/1572643.htm. Retrieved 19 November 2009.
 34.^ Redfern, Martin (24 December 1995). "Proofs of God in a photon". The Independent.
http://www.independent.co.uk/arts-entertainment/proofs-of-god-in-a-photon-1527306.html. Retrieved
19 November 2009.
35.^ Dawkins, Richard (17 September 2001). "Eulogy for Douglas Adams". Edge.
http://www.edge.org/documents/adams_index.html. Retrieved 19 November 2009.

[edit] Further reading

• [Edit this reference]
Barrow, John D.; Tipler, Frank J. (19 May 1988). The Anthropic Cosmological Principle. foreword by
John A. Wheeler. Oxford: Oxford University Press. LC 87-28148. ISBN 9780192821478.
http://books.google.com/books?id=uSykSbXklWEC&printsec=frontcover. Retrieved 31 December
2009.
• John D. Barrow, 2003. The Constants of Nature, Pantheon Books, ISBN 0-375-42221-8
• Bernard Carr, ed. (2007) Universe or Multiverse? Cambridge University Press.
• Paul Davies, 1982. The Accidental Universe, Cambridge University Press, ISBN 0-521-
24212-6
• Paul Davies, 2007. Cosmic Jackpot: Why Our Universe Is Just Right for Life, Houghton
Mifflin Harcourt, ISBN 0618592261. Reprinted as: The Goldilocks Enigma: Why Is the Universe Just
Right for Life?, 2008, Mariner Books, ISBN 0547053584.
• Alister McGrath, 2009. A Fine-Tuned Universe: The Quest for God in Science and Theology ,
Westminster John Knox Press, ISBN 0664233104.
• Simon Conway Morris, 2003. Life's Solution: Inevitable Humans in a Lonely Universe .
Cambridge Univ. Press.
 • Peter Ward and Donald Brownlee, 2000. Rare Earth: Why Complex Life is Uncommon in the
Universe. Springer Verlag.

[edit] External links

Defend fine-tuning:
• Stephen M. Barr (2001) "Overview of the Cosmological Intelligent Design argument," First
Things, the Journal of Religion, Culture, and Public Life .
• Robin Collins:
• Fine-Tuning website.
• "Cosmological fine-tuning."
• Hugh Ross:
• "Design and the Anthropic Principle."
• "Evidence For Design In The Universe."
• Gerald Schroeder: Fine-turned universe.
• The cosmos is fine-tuned to permit human life at the talk.origins index to creationist claims.
• Interview with Charles Townes discussing science and religion.
• Home page of Templeton Foundation project on fine-tuning.
Criticize fine tuning:
• Bibliography of online Links to criticisms of the Fine-Tuning Argument. Secular Web.
 • Ikeda, Michael, and Jefferys, William H., "The Anthropic Principle Does Not Support
Supernaturalism."
• Jefferys, W. H,. and J. O. Berger, "Sharpening Ockham's razor on a Bayesian strop."
• Victor Stenger:
• "Does the Cosmos Show Evidence of Purpose?"
• "Is the Universe fine-tuned for us?"
• Elliott Sober, "The Design Argument." An earlier version appeared in the Blackwell
Companion to the Philosophy of Religion (2004).
Retrieved from "http://en.wikipedia.org/wiki/Fine-tuned_Universe"

Categories: Intelligent design | Pseudophysics | Creation science | Philosophical arguments

 W000

Intelligent designer
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Intelligent design
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 Timeline
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An intelligent designer, also referred to as an intelligent agent, is the willed and self-conscious entity
that the intelligent design movement argues had some role in the origin and/or development of life and who
supposedly has left scientific evidence of this intelligent design. They also use the term "intelligent cause"
implying their teleological supposition of direction and purpose in features of the universe and of living things.
The intelligent design movement is a neo-creationist campaign that arose out of the previous
Christian fundamentalist and evangelistic creation science movement.[1][2][3] Proponents of intelligent
design argue to the public that their concept does not posit the identity of the designer as part of this effort.
But in statements to their constituency, which consists largely of Christian conservatives, they identify the
designer as God.[4][5][6][7]

Contents
[hide]
• 1 Who does the ID movement think the designer is?
• 2 What does the ID movement think the designer did?
• 3 Criticism
• 4 The Dover trial
• 5 See also
• 6 References
• 7 External links

[edit] Who does the ID movement think the designer is?

William Dembski states in his book Design Inference that the nature of the intelligent designer cannot
be inferred from intelligent design[8] and suggests that the designer, if one is even necessary for design
inference, may or may not be "the God of Scripture."[9] In December 2007 Dembski told Focus on the
Family, "I believe God created the world for a purpose. The Designer of intelligent design is, ultimately, the
Christian God."[10] Some leading intelligent design proponents have stated identifying or characterizing the
designer is beyond the scope of intelligent design as a line of inquiry. Proponents had hoped that, by
avoiding invoking creation by a specific supernatural entity, (such as that employed by creation science),
intelligent design would be considered scientific and not violate the establishment clause of the US
constitution. Proponents feared that were intelligent design identified as a restatement of previous forms of
creationism, it would be precluded from being taught in public schools after the 1987 Supreme Court of the
United States decision in Edwards vs Aguillard. This line of reasoning was not particularly persuasive to
many in the scientific community, which largely rejected intelligent design as both a line of scientific inquiry
and as a basis for a sound education in science.
On December 20, 2005 federal district court ruled in Kitzmiller v. Dover Area School District that
intelligent design was not science and was essentially religious in nature. The ruling not only rendered that
public school district's requirement endorsing intelligent design as an alternative to evolution in science
classes unconstitutional on the grounds that its inclusion violates the Establishment Clause of the First
Amendment, but validated the objections of critics who discounted proponent's claim that the identity was not
God.
Highlighting these mutually exclusive claims about the designer, Dembski, despite having said that
the intelligent designer or designers could be any god or gods, or even space aliens, has also said that
"intelligent design should be understood as the evidence that God has placed in nature to show that the
physical world is the product of intelligence and not simply the result of mindless material forces"[11] and that
"Intelligent design is just the Logos theology of John's Gospel restated in the idiom of information theory."[12]
 Michael Behe, in his book Darwin's Black Box, suggested the designer might be a time traveling cell
biologist.[13]
At various times, leading proponents in the intelligent design movement have clearly expressed that
they consider the Abrahamic God "Elohim" in his role as a creator God, to be the intelligent designer and
denied that intelligent designer is God, depending on which audience they are addressing. One example is
William Dembski, who on his blog in response to the question "Is the designer responsible for biological
complexity God?" said "not necessarily" and "To ask who or what is the designer of a particular object is to
ask for the immediate intelligent agent responsible for its design. The point is that God is able to work
through derived or surrogate intelligences, which can be anything from angels to organizing principles
embedded in nature."[14] Yet to the intelligent design movement's conservative Christian constituents
Dembski has said "intelligent design should be understood as the evidence that God has placed in nature to
show that the physical world is the product of intelligence and not simply the result of mindless material
forces. This evidence is available to all apart from the special revelation of God in salvation history as
recounted in Scripture. ... Intelligent design makes it impossible to be an intellectually fulfilled atheist. This
gives intelligent design incredible traction as a tool for apologetics, opening up the God-question to
individuals who think that science has buried God"[15] and "Thus, in its relation to Christianity, intelligent
design should be viewed as a ground-clearing operation that gets rid of the intellectual rubbish that for
generations has kept Christianity from receiving serious consideration."[16] Stephen C. Meyer, founder and
leader of the intelligent design program of the Discovery Institute admitted on national television he believes
that the designer is God.[17]
In addition, the intelligent design movement seeks as a well-documented agenda the overall goal "to
defeat materialism" and the "materialist world view" as represented by evolution, and replace it with "a
science consonant with Christian and theistic convictions."[18] Phillip E. Johnson, considered the father of
the ID movement has stated that the goal of intelligent design is to cast creationism as a scientific concept:
"Our strategy has been to change the subject a bit so that we can get the issue of intelligent
design, which really means the reality of God, before the academic world and into the schools."
-- Phillip E. Johnson, American Family Radio, January 10, 2003 [19]

"This isn't really, and never has been a debate about science. It's about religion and
philosophy." -- Phillip E. Johnson, World Magazine, November 30, 1996 [20]

The Discovery Institute's leaked Wedge document sets out the movement's governing goals,
including:
"To replace materialistic explanations with the theistic understanding that nature and human
beings are created by God." . . . "Design theory promises to reverse the stifling dominance of the
materialist worldview, and to replace it with a science consonant with Christian and theistic
convictions."[21] -- The Wedge Document, a 1999 Discovery Institute pamphlet

[edit] What does the ID movement think the designer did?

Opinion as to the amount of creation the intelligent designer has done varies within the ID movement.
Michael Behe's concept of irreducible complexity has natural selection accounting for most of evolution but
the intelligent designer contributing the design of some proteins. Others in the ID movement however contest
concepts such as common descent, particularly of humans and other apes. Though most in the ID movement
seem to be Old Earth Creationists, a few are Young Earth Creationists who believe in ex-nihilo.
 The amount of creation that the intelligent designer did has also been criticised by Young Earth
Creationists as not being specific enough, and particularly contradicting their beliefs of Biblical inerrancy and
a young earth.[22][23]
Some intelligent design proponents say the intelligent designer fine-tuned the universe's physical
constants in such a way that life is the result of the universe's physical constants being related to one another
in a fashion that permits life to exist. The fine-tuned universe argument is a central premise or presented as a
given in many of the published works of prominent intelligent design proponents, such as William A. Dembski
and Michael Behe.

[edit] Criticism
Intelligent design has been presented by its proponents as a "big tent" strategy into which several
accounts of creation can fit. Were a scientific version of intelligent design approved for inclusion in public
school science curricula, then a path would be opened for discussion of alternatives to not only natural
selection but naturalism as well, and eventually religious accounts on the origin of life. The vast majority of
scientists reject the concept of intelligent design and an intelligent designer. Instead, the most widely
accepted explanation is that physical processes such as natural selection can account for the complexity of
life and other phenomena and features of the universe. Attempts to insert theories of intelligent design into
public school science curricula fits in with the intelligent design movement's social aims, via the overturning
of Western secularism as detailed in the Wedge strategy. The concept of the intelligent designer has been
criticised as a God-of-the-gaps argument. Introducing the hypothesis of an intelligent designer introduces the
unsolved problem of accounting for the origin of such a designer (first cause).
By raising the question of the need for a designer for objects with irreducible complexity, intelligent
design also raises the question, "what designed the designer?" Richard Dawkins has argued that "If complex
organisms demand an explanation, so does a complex designer. And it's no solution to raise the theologian's
plea that God (or the intelligent designer) is simply immune to the normal demands of scientific
explanation,"[24] since such an answer would be unscientific. With religious creationism, the question "what
created God?" can be answered with theological arguments, but in intelligent design, the chain of designers
can be followed back indefinitely in an infinite regression, leaving the question of the creation of the first
designer dangling. As a result, intelligent design does not explain how the complexity happened in the first
place; it just moves it.[25]
Elliott Sober says that by intelligent design's own arguments, a designer capable of creating
irreducible complexity must also be irreducibly complex: "Any mind in nature that designs and builds an
irreducibly complex system is itself irreducibly complex"[26] Sober says that this an argument that intelligent
design proponents still need to respond to.
If intelligent design proponents invoke an uncaused causer or deity to resolve this problem,[27] they
contradict a fundamental assumption of intelligent design that design requires a designer[28][29] and reduce
intelligent design to religious creationism. Another possible counter-argument might be an infinite regression
of designers. However, admitting infinite numbers of objects also allows any arbitrarily improbable event to
occur [30], such as an object with "specific" complexity assembling itself by chance. Again, this contradicts a
fundamental assumption of intelligent design that a designer is needed for every specifically complex object,
producing a logical contradiction.
Critics contend the claim that positing a designer which explains gaps in our understanding yet does
not need to be itself explained as not a contribution to knowledge but as a thought-terminating cliché.[31][32]
[edit] The Dover trial
In 2005, intelligent design proponents arguments regarding the identity of a designer became an
issue considered by the court in Kitzmiller v. Dover Area School District, the "Dover trial," where plaintiffs
successfully argued that intelligent design is a form of creationism, and that the school board policy requiring
the presentation of intelligent design as an alternative to evolution as an "explanation of the origin of life" thus
violated the Establishment Clause of the First Amendment to the United States Constitution. In his ruling, the
judge stated
"However, as Dr. Haught testified, anyone familiar with Western religious thought would
immediately make the association that the tactically unnamed designer is God..." -- Ruling,
Kitzmiller v. Dover Area School District, page 25

Jones also commented that the appearance of design is subjective,

"It is readily apparent to the Court that the only attribute of design that biological systems appear
to share with human artifacts is their complex appearance, i.e. if it looks complex or designed, it
must have been designed. (23:73 (Behe)). This inference to design based upon the appearance
of a "purposeful arrangement of parts" is a completely subjective proposition, determined in the
eye of each beholder and his/her viewpoint concerning the complexity of a system." -- Ruling,
Kitzmiller v. Dover Area School District, page 81

and thus the analogy upon which the argument from design rests is flawed.
"For human artifacts, we know the designer's identity, human, and the mechanism of design, as
we have experience based upon empirical evidence that humans can make such things, as well
 as many other attributes including the designer's abilities, needs, and desires. With ID,
proponents assert that they refuse to propose hypotheses on the designer's identity, do not
propose a mechanism, and the designer, he/she/it/they, has never been seen. In that vein,
defense expert Professor Minnich agreed that in the case of human artifacts and objects, we
know the identity and capacities of the human designer, but we do not know any of those
attributes for the designer of biological life. In addition, Professor Behe agreed that for the
design of human artifacts, we know the designer and its attributes and we have a baseline for
human design that does not exist for design of biological systems. Professor Behe's only
response to these seemingly insurmountable points of disanalogy was that the inference still
works in science fiction movies. -- Ruling, Kitzmiller v. Dover Area School District, page 81

[edit] See also

• Anthropic principle
• Conflict thesis
• Continuity thesis
• Flying Spaghetti Monster
• Great Architect of the Universe
• Genesis creation myth

[edit] References
1. ^ Ruling, Context Kitzmiller v. Dover Area School District
 2. ^ Life in the Big Tent: Traditional Creationism and the Intelligent Design Community Paul A.
Nelson. Christian Research Journal, volume 24, number 4, 2002.
3. ^ Expert Witness Report Barbara Forrest. Kitzmiller v. Dover Area School District. (PDF file)
4. ^ Stephen C. Meyer (Discovery Institute): "I think the designer is God..." Darwin, the
marketing of Intelligent Design Nightline ABC News, with Ted Koppel, August 10, 2005.
5. ^ "Now the way that I see the logic of our movement going is like this. The first thing you
understand is that the Darwinian theory isn't true. It's falsified by all of the evidence and the logic is
terrible. When you realize that, the next question that occurs to you is, well, where might you get the
truth? When I preach from the Bible, as I often do at churches and on Sundays, I don't start with
Genesis. I start with John 1:1. In the beginning was the word. In the beginning was intelligence,
purpose, and wisdom. The Bible had that right. And the materialist scientists are deluding
themselves." How the Evolution Debate Can Be Won. Phillip Johnson. Truths that Transform.
6. ^ William Dembski. "Intelligent design is just the Logos theology of John's Gospel restated in
the idiom of information theory," Touchstone Magazine. Volume 12, Issue 4 July/August, 1999
7. ^ "The only apparent difference between the argument made by Paley and the argument for
ID, as expressed by defense expert witnesses Behe and Minnich, is that ID's "official position" does
not acknowledge that the designer is God. However, as Dr. Haught testified, anyone familiar with
Western religious thought would immediately make the association that the tactically unnamed
designer is God, as the description of the designer in Of Pandas and People (hereinafter "Pandas") is
a "master intellect," strongly suggesting a supernatural deity as opposed to any intelligent actor
known to exist in the natural world. (P-11 at 85). Moreover, it is notable that both Professors Behe
and Minnich admitted their personal view is that the designer is God and Professor Minnich testified
that he understands many leading advocates of ID to believe the designer to be God. (21:90 (Behe);
38:36-38 (Minnich)). Although proponents of the IDM occasionally suggest that the designer could be
a space alien or a time-traveling cell biologist, no serious alternative to God as the designer has been
proposed by members of the IDM, including Defendants' expert witnesses. (20:102-03 (Behe)). In
fact, an explicit concession that the intelligent designer works outside the laws of nature and science
and a direct reference to religion is Pandas’ rhetorical statement, "what kind of intelligent agent was it
[the designer]" and answer: "On its own science cannot answer this question. It must leave it to
religion and philosophy." (P-11 at 7; 9:13-14 (Haught)). A significant aspect of the IDM is that despite
Defendants' protestations to the contrary, it describes ID as a religious argument. In that vein, the
writings of leading ID proponents reveal that the designer postulated by their argument is the God of
Christianity." Dover trial ruling: Context Kitzmiller v. Dover Area School District
8. ^ William A. Dembski, The Design Inference: Eliminating Chance through Small Probabilities,
Cambridge University Press, 1998, ISBN 9780521623872, p. 9: "The effect of a design inference is
to limit our explanatory options, not to identify a cause. To identify a cause we need to investigate the
particulars of the situation in which design is inferred. Simply put, we need more details....The design
inference does not by itself deliver an intelligent agent."
9. ^ William A. Dembski, The Design Inference: Eliminating Chance through Small Probabilities,
Cambridge University Press, 1998, ISBN 9780521623872, p. 60.
10.^ "Friday Five: William A. Dembski". Focus on the Family. December 14, 2007.
http://www.citizenlink.org/content/A000006139.cfm. Retrieved 2008-05-17.
11.^ Why President Bush Got It Right about Intelligent Design William A. Dembski
DesignInference.com, August 4, 2005 (PDF file)
12.^ Signs of Intelligence A Primer on the Discernment of Intelligent Design. William A.
Dembski. Touchstone Journal, Volume 12, Issue 4, July/August 1999
13.^ Behe, Michael J. (1998). Darwin's Black Box: The Biochemical Challenge to Evolution .
Simon and Schuster. p. 249. ISBN 9780743214858.
 14.^ Who or what is the designer? William A. Dembski. Uncommon Descent, December 22,
2005.
15.^ Commending President Bush William A. Dembski. DesignInference.com
16.^ Intelligent Design's Contribution To The Debate Over Evolution: A Reply To Henry Morris
William A. Dembski. DesignInference.com.
17.^ Stephen C. Meyer: "I think the designer is God..." Darwin, the marketing of Intelligent
Design Nightline ABC News with Ted Koppel, August 10, 2005.
18.^ Evolution or design debate heats up Paul Handley. The Times of Oman, March 7, 2005
19.^ Robert T. Pennock, Ph.D. (March 31, 2005). "Kitzmiller, et al. v. Dover Area School District
- Expert Report" (pdf). http://www.msu.edu/~pennock5/research/papers/Pennock_DoverExptRpt.pdf.
Retrieved 2007-12-19. p. 4
20.^ Joel Belz (November 30, 1996). "World Magazine Article". Witnesses for the Prosecution.
World. http://www.leaderu.com/pjohnson/world2.html. Retrieved 2007-12-21.
21.^ The Wedge Document Discovery Institute pamphlet, 1999. (PDF file)
22.^ AiG’s views on the Intelligent Design Movement - by Carl Wieland, 30 August 2002 [1]
23.^ 'Design is Not Enough' - by Henry H. Morris, Back to Genesis pamphlet series, No.127a,
July 1999, Institute for Creation Research. [2]
24.^ "If complex organisms demand an explanation, so does a complex designer. And it's no
solution to raise the theologian's plea that God (or the Intelligent Designer) is simply immune to the
normal demands of scientific explanation. To do so would be to shoot yourself in the foot." Richard
Dawkins and Jerry Coyne. 1 September 2005. The Guardian [3]
25.^ Claudia Wallis. Evolution Wars. Time Magazine, 15 August 2005 edition, page 32 [4]
26.^ Intelligent Design Theory and the Supernatural - The "God or Extra-terrestrials" Reply
Elliott Sober. University of Wisconsin - Madison.
 27.^ "Christianity postulates the religious answer to this question that the designer is God who
by definition is eternally existent and has no origin. There is no logical philosophical impossibility with
this being the case (akin to Aristotle's 'unmoved mover') as a religious answer to the origin of the
designer. See also an answer to a subissue the implications of whether or not the first CSI come from
an unintelligent source." FAQ: Who designed the designer? IDEA [5]
28.^ "Intelligent design, on the other hand, involves two basic assumptions: 1) Intelligent causes
exist. 2) These causes can be empirically detected (by looking for specified complexity). " Access
Research Network. Frequently Asked Questions about Intelligent Design. [6]
29.^ "According to contemporary design theory, the presence of highly specified complexity is
an indicator of an intelligent cause." Access Research Network. Frequently Asked Questions about
Intelligent Design. [7]
30.^ "To explain the origin of the DNA/protein machine by invoking a supernatural Designer is to
explain precisely nothing, for it leaves unexplained the origin of the Designer. You have to say
something like 'God was always there', and if you allow yourself that kind of lazy way out, you might
as well just say 'DNA was always there', or "Life was always there', and be done with it." Richard
Dawkins. The Blind Watchmaker : Why the Evidence of Evolution Reveals a Universe without Design
pg 141
31.^ Who Designed the Designer? Jason Rosenhouse. Creation & Intelligent Design Watch,
Committee for the Scientific Investigation of Claims of the Paranormal.
32.^ Richard Dawkins. The Blind Watchmaker: Why the Evidence of Evolution Reveals a
Universe without Design pg 141
 [edit] External links
• Claim CI001: Intelligent design (ID) is scientific from the talk.origins Archive's Index to
Creationist Claims, and sub-claims.
• Evolution & Creation: A Theosophic Synthesis Surveys critical problems on both sides of the
ID controversy and explores ancient and modern concepts of intelligent designer(s)
• Who Designed the Designer? by Jason Rosenhouse

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Creationism topics

Types Young Earth Creationism · Old Earth creationism · Progressive

creationism · Theistic evolution · Intelligent design

Genesis Genesis creation narrative · Framework interpretation · Day-Age theory ·

interpretations Gap theory · Biblical literalism

Related concepts Creation myth · Omphalos hypothesis · Specified complexity ·

Irreducible complexity · Dating Creation · Theistic realism · Intelligent designer

Pseudoscience Creation science · Creationist cosmologies · Creation biology · Created

kinds · Flood geology · Vapor canopy · Modern geocentrism
 Creation-evolution controversy · History of creationism · Creation and
Miscellanea evolution in public education · Quotes about creation and evolution · Teach the
Controversy

Retrieved from "http://en.wikipedia.org/wiki/Intelligent_designer"

 W000

Teleonomy
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Teleonomy is the quality of apparent purposefulness and of goal-directedness of structures and
functions in living organisms that derive from their evolutionary history, adaptation for reproductive success,
or generally, due to the operation of a program.
The term was coined to stand in contrast with teleology, which applies to ends that are planned by an
agent which can internally model/imagine various alternative futures, which enables intention, purpose and
foresight. A teleonomic process, such as evolution, produces complex products without the benefit of such a
guiding foresight. Evolution largely hoards hindsight, as variations unwittingly make "predictions" about
structures and functions which could successfully cope with the future, and participate in an audition which
culls the also-rans, leaving winners for the next generation. Information accumulates about functions and
structures that are successful, exploiting feedback from the environment via the selection of fitter coalitions of
structures and functions. These features also have been described by Robert Rosen as an anticipatory
system which builds an internal model based on past and possible futures states. Teleonomy is related to
programmatic or computational aspects of purpose. Richard Dawkins has also described the properties of
"archeo-purpose" and "neo-purpose" in his talk on the "purpose of purpose."[1]

Contents
[hide]
• 1 History
• 2 Philosophy
• 3 Current status
• 4 See also
• 5 References
• 6 Further reading
• 7 External links

[edit] History
 In 1958, C.S. Pittendrigh applied the term to biology:
Biologists for a while were prepared to say a turtle came ashore and laid its eggs. These verbal
scruples were intended as a rejection of teleology but were based on the mistaken view that the
efficiency of final causes is necessarily implied by the simple description of an end-directed
mechanism. … The biologists long-standing confusion would be removed if all end-directed
systems were described by some other term, e.g., ‘teleonomic,’ in order to emphasize that
recognition and description of end-directedness does not carry a commitment to Aristotelian
teleology as an efficient causal principle.[2]

In 1962, Grace A. de Laguna's "The Role of Teleonomy in Evolution"[3] fleshed the applicability of
the term to biological history and adaptation.
In 1965 Ernst Mayr cited[4] Pittendrigh and criticized the last few words cited above for not making a
“clear distinction between the two teleologies of Aristotle”; evolution involves Aristotle's material causes and
formal causes rather than efficient causes. Mayr adopted Pittendrigh’s term, but supplied his own definition:
It would seem useful to rigidly restrict the term teleonomic to systems operating on the basis of a
program of coded information. (p.42)

In 1966 George C. Williams approved of the term in the last chapter of his Adaptation and Natural
Selection; a critique of some current evolutionary thought .[5] In 1970, Jacques Monod, in Chance and
Necessity, an Essay on the Natural Philosophy of Modern Biology ,[6] suggested teleonomy as a key feature
that defines life:
 Rather than reject this [goal-directedness] idea (as certain biologists have tried to do) it is
indispensable to recognise that it is essential to the very definition of living beings. We shall
maintain that the latter are distinct from all other structures or systems present in the universe
through this characteristic property, which we shall call teleonomy.

[..] It will be readily seen that, in this or that species situated higher or lower on the animal scale,
the achievement of the fundamental teleonomic project (i.e., invariant reproduction) calls
assorted, more or less elaborate and complex structures and performances into play. The fact
must be stressed that concerned here are not only the activities directly bound up with
reproduction itself, but all those that contribute-be it very indirectly-to the species' survival and
multiplication. For example, in higher mammals the play of the young is an important element of
psychic development and social integration. Therefore this activity has teleonomic value,
inasmuch as it furthers the cohesion of the group, a condition for its survival and for the
expansion of the species.

In 1974 Ernst Mayr illustrated[7] the difference in the statements:

"The Wood Thrush migrates in the fall in order to escape the inclemency of the weather and the food
shortages of the northern climates."
"The Wood Thrush migrates in the fall and thereby escapes the inclemency of the weather and the
food shortages of the northern climates."
If we replace the words ‘in order to escape’ by ‘and thereby escapes’, we leave the important
question unanswered as to why the Wood Thrush migrates. The teleonomic form of the
 statement implies that the goal-directed migratory activity is governed by a program. By omitting
this important message the second sentence is greatly impoverished as far as information
content is concerned, without gaining in causal strength.

Subsequently philosophers like Ernest Nagel further analysed[8] the concept of goal-directedness in
biology and by 1982, philosopher and historian of science David Hull joked[9] about the use of teleology and
teleonomy by biologists:
Haldane [in the 1930s] can be found remarking, ‘Teleology is like a mistress to a biologist: he
cannot live without her but he’s unwilling to be seen with her in public.’ Today the mistress has
become a lawfully wedded wife. Biologists no longer feel obligated to apologize for their use of
teleological language; they flaunt it. The only concession which they make to its disreputable
past is to rename it ‘teleonomy’.

Although Aristotle believed the world as a whole has a purpose and is teleologically guided, his
ancient closing argument could be taken as a statement about teleonomy given modern understanding of
adaptation and feedback as "nature's craft":
It is absurd to suppose that purpose is not present because we do not observe an agent
deliberating. Craft does not deliberate. If the ship-building craft were in the wood, it would
produce the same results by nature. If, therefore, purpose is present in craft, it is present also in
nature. The best illustration is a doctor doctoring himself: nature is like that. It is plain that nature
is a cause, a cause that operates for a purpose.
—Physics, Book II, ch.8, 199b
[edit] Philosophy
In teleology, Kant's positions were neglected for many years because in the minds of many scientists
they were associated with vitalist views of evolution. Their gradual rehabilitation recently is evident in
teleonomy which bears a number of features, such as the description of organisms, that are reminiscent of
the Kantian conception of final causes as essentially recursive in nature. The gist of Kant's position is that
even though we cannot know whether there are final causes in nature, we are constrained by the peculiar
nature of the human understanding to view organisms teleologically. Thus the Kantian view sees teleology as
a necessary principle for the study of organisms, but only as a regulative principle, and with no ontological
implications.

[edit] Current status

Teleonomy is closely related to concepts of emergence, complexity theory[10] and self-organizing
systems.[11] It has extended beneath biology to be applied in the context of chemistry.[12][13] Some
philosophers of biology resist the term and still employ "teleology" when analyzing biological function[14] and
the language used to describe it.[15] while others endorse it.[16] The debate also intersects with theoretical
physics and the foundations of mathematical modeling, if space-time evolves concurrently with information-
cognition, then reflexive self-processing (Wheeler's It from Bit) may allow life the ability to promote the
environmental conditions for its survival. Whether through coincidence via multiversal self-selection (as
discussed in Susskind's Landscape) or through cybernetic systems, both "teleology" and "teleonomy" lack
the precision to describe the physical mechanisms involved and do not address the potential role of
hyperincursion in anticipatory computing.
[edit] See also
Look up teleonomy in Wiktionary, the free dictionary.

• Autopoiesis
• Naturalism (philosophy)
• Orthogenesis
• Polytely
• Purpose
• Telic Recursion
• T-symmetry

[edit] References
1. ^ http://richarddawkins.net/article,3956,The-Purpose-of-Purpose,Richard-Dawkins
2. ^ Pittendrigh, C. S. "Adaptation, natural selection, and behavior," in Behavior and Evolution,
ed. A. Roe and George Gaylord Simpson, New Haven: Yale University Press, 1958, 390–416; p. 394.
3. ^ de Laguna, Grace A. (April 1962). "The Role of Teleonomy in Evolution". Philosophy of
Science 29 (2): 117–131. doi:10.1086/287855. http://links.jstor.org/sici?sici=0031-
8248(196204)29%3A2%3C117%3ATROTIE%3E2.0.CO%3B2-N#abstract.
4. ^ Mayr, E. (1965). "Cause and effect in biology". In Lerner, D.. Cause and effect. New York:
Free Press. pp. 33–50.
 5. ^ Williams, G.C. (1966). Adaptation and natural selection; a critique of some current
evolutionary thought. Princeton, NJ: Princeton University Press. ISBN 0691026157.
6. ^ Monod, Jacques (1971). Chance and Necessity: An Essay on the Natural Philosophy of
Modern Biology. New York: Alfred A. Knopf. ISBN 0-394-46615-2.
7. ^ Mayr, E. (1974). "Teleological and Teleonomic, a New Analysis". Boston Studies in the
Philosophy of Science 14.
8. ^ Nagel, E. (1977). "Teleology Revisited: Goal-Directed Processes in Biology". Journal of
Philosophy 74 (5): 261–301. doi:10.2307/2025745. http://jstor.org/stable/2025745. Reprinted in Allen,
Bekoff & Lauder, 1998.
9. ^ Hull, D.L. (1982). "Philosophy and Biology". In Fløistad, Guttorm. Philosophy of Science.
Contemporary Philosophy: A New Survey. 2. Hague: Nijhoff. pp. 280–316. ISBN 90-247-2518-6.
10.^ Christensen, W.D. (1996). "A complex systems theory of teleology" (PDF). Biology and
Philosophy 11: 301–320. doi:10.1007/BF00128784.
http://www.springerlink.com/index/N065P12442750447.pdf.
11.^ Lifson S (May 1987). "Chemical selection, diversity, teleonomy and the second law of
thermodynamics. Reflections on Eigen's theory of self-organization of matter". Biophys. Chem. 26 (2-
3): 303–11. doi:10.1016/0301-4622(87)80031-5. PMID 3607231.
http://linkinghub.elsevier.com/retrieve/pii/0301-4622(87)80031-5.
12.^ Pross, Addy (August 2005). "On the Chemical Nature and Origin of Teleonomy". Origins of
Life and Evolution of Biospheres 35 (4). http://www.springerlink.com/content/7355r78446n0ppr6/.
13.^ ibid. On the Chemical Nature of Purpose (Teleonomy), Journal of 18th Conference on
Physical Organic Chemistry
14.^ Neander, K. (1991). "The Teleological Notion of ‘Function’". Australasian Journal of
Philosophy 69 (4): 454–468. doi:10.1080/00048409112344881.
 15.^ Nissen, Lowell (1997). Teleological Language in the Life Sciences. Rowman & Littlefield.
ISBN 0-8476-8694-9.
16.^ Kober, G. "Teleology's New Clothes: Teleonomy and the Notion of Program", International
Society for the History, Philosophy, and Social Studies of Biology (Feb 14, 2005)

[edit] Further reading

• Allen, C., M. Bekoff, G. Lauder, eds., Nature’s Purposes: Analyses Of Function and Design
in Biology. MIT Press, 1998. (ISBN 0-2625-1097-9)
• Mayr, E., The Growth of Biological Thought. Diversity, Evolution, and Inheritance. Cambridge
(MA): Belknap Press of Harvard University Press, 1982 : pp. 47-51 (differentiating four kinds of
teleology).
• Mayr, E., What Makes Biology Unique?: Considerations on the Autonomy of a Scientific
Discipline, Cambridge University Press, 2004. (ISBN 0-5218-4114-3).
• Ruse, M. Darwin and Design, Harvard University Press; 2004. (ISBN 0-6740-1631-9)

[edit] External links

• Merriam Webster definition
• Nonlinearity and Teleology
• Biological Information
Retrieved from "http://en.wikipedia.org/wiki/Teleonomy"

Categories: Philosophical terminology

 W000

Téléonomie
 Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

La téléonomie (du grec telos, but et nomos, loi) est un terme inventé par Colin S. Pittendrigh en
1958[1], puis repris par de nombreux autres biologistes de renom, dont Jacques Monod dans les années
1970[2],[3],[4]. Il désigne le concept scientifique de finalité. Il s'agit ne pas confondre avec la téléologie (de
logos = raison, parole) car Monod refuse, par principe, tout recours à une Raison créatrice dans le
développement des processus biologiques.
Le mot de téléonomie résume entre autres la suite de raisons pour lesquelles trois animaux aussi
différents que l'ichtyosaure, le requin et le dauphin ont des formes hydrodynamiques similaires bien
qu'appartenant à des branches différentes du vivant.
Téléonomie ou qualité de la matière vivante en tant qu'elle matérialise un projet.
Notes et références [modifier]
1. ↑ (en) « Biologists for a while were prepared to say a turtle came ashore and laid its eggs.
These verbal scruples were intended as a rejection of teleology but were based on the mistaken view
that the efficiency of final causes is necessarily implied by the simple description of an end-directed
mechanism. … The biologists long-standing confusion would be removed if all end-directed systems
were described by some other term, e.g., ‘teleonomic,’ in order to emphasize that recognition and
description of end-directedness does not carry a commitment to Aristotelian teleology as an efficient
causal principle. » Colin S. Pittendrigh, Adaptation, natural selection, and behavior in Behavior and
Evolution, ed. A. Roe and George Gaylord Simpson, New Haven: Yale University Press, 1958, 390-
416; p. 394
2. ↑ « La pierre angulaire de la méthode scientifique est le postulat de l'objectivité de la Nature.
C'est-à-dire le refus systématique de considérer comme pouvant conduire à une connaissance
"vraie" toute interprétation des phénomènes donnée en termes de causes finales, c'est-à-dire de
"projet". [...] Postulat pur, à jamais indémontrable, car il est évidemment impossible d'imaginer une
expérience qui pourrait prouver la non-existence d'un projet, d'un but poursuivi, où que ce soit dans
la nature. Mais le postulat d'objectivité est consubstantiel à la science, il a guidé tout son prodigieux
développement depuis trois siècles. Il est impossible de s'en défaire, fût-ce provisoirement, ou dans
un domaine limité, sans sortir de celui de la science elle-même. L'objectivité cependant nous oblige à
reconnaître le caractère téléonomique des êtres vivants, à admettre que dans leurs structures et
performances, ils réalisent et poursuivent un projet. Il y a donc là, au moins en apparence, une
contradiction épistémologique profonde. Le problème central de la biologie, c'est cette contradiction
elle-même, qu'il s'agit de résoudre si elle n'est qu'apparente, ou de prouver radicalement insoluble si
en vérité il en est bien ainsi. » (Jacques Monod, Le Hasard et la Nécessité, éd. du Seuil, coll.
« Points », p. 37-38).
3. ↑ « L'invariance précède nécessairement la téléonomie. Ou, pour être plus explicite, l'idée
darwinienne que l'apparition, l'évolution, le raffinement progressif de structures de plus en plus
intensément téléonomiques sont dus à des perturbations survenant dans une structure possédant
déjà la propriété d'invariance, capable par conséquent de "conserver le hasard" et par là d'en
soumettre les effets au jeu de la sélection naturelle. » (Monod, op. cit., p. 37).
4. ↑ « Toutes les autres conceptions qui ont été explicitement proposées pour rendre compte
de l'étrangeté des êtres vivants, ou qui sont implicitement enveloppées par des idéologies
religieuses comme par la plupart des grands systèmes philosophiques, supposent l'hypothèse
inverse : à savoir que l'invariance est protégée, l'ontogénie guidée, l'évolution orientée par un
principe téléonomique initial, dont tous les phénomènes seraient des manifestations. » (Monod,
op. cit., p. 38).

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W000
INTERNATIONAL THEOLOGICAL
COMMISSION
COMMUNION AND STEWARDSHIP:

Human Persons Created in the Image of God*

INTRODUCTION
1. The explosion of scientific understanding and technological capability in modern times
has brought many advantages to the human race, but it also poses serious challenges. Our
knowledge of the immensity and age of the universe has made human beings seem smaller and less
secure in their position and significance within it. Technological advances have greatly increased
our ability to control and direct the forces of nature, but they have also turned out to have an
unexpected and possibly uncontrollable impact on our environment and even on ourselves.
 2. The International Theological Commission offers the following theological meditation on
the doctrine of the imago Dei to orient our reflection on the meaning of human existence in the face
of these challenges. At the same time, we want to present the positive vision of the human person
within the universe which is afforded by this newly retrieved doctrinal theme.
3. Especially since Vatican Council II, the doctrine of the imago Dei has begun to enjoy a
greater prominence in magisterial teaching and theological research. Previously, various factors had
led to the neglect of the theology of the imago Dei among some modern western philosophers and
theologians. In philosophy, the very notion of the "image" was subjected to a powerful critique by
theories of knowledge which either privileged the role of the "idea" at the expense of the image
(rationalism) or made experience the ultimate criterion of truth without reference to the role of the
image (empiricism). In addition, cultural factors, such as the influence of secular humanism and,
more recently, the very profusion of images by the mass media, have made it difficult to affirm the
human orientation to the divine, on the one hand, and, on the other, the ontological reference of the
image which are essential to any theology of the imago Dei. Contributing to the neglect of the
theme within western theology itself were biblical interpretations that stressed the permanent
validity of the injunction against images (cf. Exodus 20:3-4) or posited a Hellenistic influence on
the emergence of the theme in the Bible.
4. It was not until the eve of Vatican Council II that theologians began to rediscover the
fertility of this theme for understanding and articulating the mysteries of the Christian faith. Indeed,
the documents of this council both express and confirm this significant development in twentieth
century theology. In continuity with the deepening recovery of the theme of the imago Dei since
Vatican Council II, the International Theological Commission seeks in the following pages to
reaffirm the truth that human persons are created in the image of God in order to enjoy personal
communion with the Father, Son and Holy Spirit and with one another in them, and in order to
exercise, in God's name, responsible stewardship of the created world. In the light of this truth, the
world appears not as something merely vast and possibly meaningless, but as a place created for the
sake of personal communion.
5. As we seek to demonstrate in the following chapters, these profound truths have lost
neither their relevance nor their power. After a summary review of the scriptural and traditional
basis of the imago Dei in Chapter I, we move on to an exploration of the two great themes of the
theology of the imago Dei: in Chapter II, the imago Dei as the basis of communion with the triune
God and among human persons and then, in Chapter III, the imago Dei as the basis of a share in
God's governance of visible creation. These reflections gather together the main elements of
Christian anthropology and certain elements of moral theology and ethics as they are illumined by
the theology of the imago Dei. We are well aware of the breadth of the issues we have sought to
address here, but we offer these reflections to recall for ourselves and for our readers the immense
explanatory power of the theology of the imago Dei precisely in order to reaffirm the divine truth
about the universe and about the meaning of human life.
 CHAPTER ONE
HUMAN PERSONS CREATED IN THE IMAGE OF GOD
6. As the witness of Scripture, Tradition and the Magisterium makes clear, the truth that
human beings are created in the image of God is at the heart of Christian revelation. This truth was
recognized and its broad implications expounded by the Fathers of the Church and by the great
scholastic theologians. Although, as we shall note below, this truth was challenged by some
influential modern thinkers, today biblical scholars and theologians join with the Magisterium in
reclaiming and reaffirming the doctrine of the imago Dei.
1. The imago Dei in Scripture and Tradition
7. With few exceptions, most exegetes today acknowledge that the theme of the imago Dei is
central to biblical revelation (cf. Gen. 1:26f; 5:1-3; 9:6). The theme is seen as the key to the biblical
understanding of human nature and to all the affirmations of biblical anthropology in both the Old
and New Testaments. For the Bible, the imago Dei constitutes almost a definition of man: the
mystery of man cannot be grasped apart from the mystery of God.
8. The Old Testament understanding of man as created in the imago Dei in part reflects the
ancient Near Eastern idea that the king is the image of God on earth. The biblical understanding,
however, is distinctive in extending the notion of the image of God to include all men. An additional
contrast with ancient Near Eastern thought is that the Bible sees man as directed, not first of all to
the worship of the gods, but rather to the cultivation of the earth (cf. Gen 2:15). Connecting cult
more directly with cultivation, as it were, the Bible understands that human activity in the six days
of the week is ordered to the Sabbath, a day of blessing and sanctification.
9. Two themes converge to shape the biblical perspective. In the first place, the whole of
man is seen as created in the image of God. This perspective excludes interpretations which locate
the imago Dei in one or another aspect of human nature (for example, his upright stature or his
intellect) or in one of his qualities or functions (for example, his sexual nature or his domination of
the earth). Avoiding both monism and dualism, the Bible presents a vision of the human being in
which the spiritual is understood to be a dimension together with the physical, social and historical
dimensions of man.
10. Secondly, the creation accounts in Genesis make it clear that man is not created as an
isolated individual: “God created mankind in his image, in the image of God he created them, male
and female he created them” (Gen. 1:27). God placed the first human beings in relation to one
another, each with a partner of the other sex. The Bible affirms that man exists in relation with other
persons, with God, with the world, and with himself. According to this conception, man is not an
isolated individual but a person -- an essentially relational being. Far from entailing a pure
actualism that would deny its permanent ontological status, the fundamentally relational character
of the imago Dei itself constitutes its ontological structure and the basis for its exercise of freedom
and responsibility.
11. The created image affirmed by the Old Testament is, according to the New Testament, to
be completed in the imago Christi. In the New Testament development of this theme, two
distinctive elements emerge: the christological and Trinitarian character of the imago Dei, and the
role of sacramental mediation in the formation of the imago Christi.
12. Since it is Christ himself who is the perfect image of God (2 Cor 4:4; Col 1:15; Heb 1:3),
man must be conformed to him (Rom 8:29) in order to become the son of the Father through the
power of the Holy Spirit (Rom 8:23). Indeed, to "become" the image of God requires an active
participation on man’s part in his transformation according to the pattern of the image of the Son
(Col 3:10) who manifests his identity by the historical movement from his incarnation to his glory.
According to the pattern first traced out by the Son, the image of God in each man is constituted by
his own historical passage from creation, through conversion from sin, to salvation and
consummation. Just as Christ manifested his lordship over sin and death through his passion and
resurrection, so each man attains his lordship through Christ in the Holy Spirit -- not only over the
earth and the animal kingdom (as the Old Testament affirms) – but principally over sin and death.
13. According to the New Testament, this transformation into the image of Christ is
accomplished through the sacraments, in the first place as an effect of the illumination of the
message of Christ (2 Cor 3:18-4:6) and of Baptism (1 Cor 12:13). Communion with Christ is a
result of faith in him, and Baptism through which one dies to the old man through Christ (Gal 3:26-
28) and puts on the new man (Gal 3:27; Rom 13:14). Penance, the Eucharist, and the other
sacraments confirm and strengthen us in this radical transformation according to the pattern of
Christ’s passion, death and resurrection. Created in the image of God and perfected in the image of
Christ by the power of the Holy Spirit in the sacraments, we are embraced in love by the Father. 14.
The biblical vision of the image of God continued to occupy a prominent place in Christian
anthropology in the Fathers of the Church and in later theology, right up to the beginning of modern
times. An indication of the centrality of this theme can be found in the endeavor of early Christians
to interpret the biblical prohibition against artistic representations of God (cf. Ex 20:2f; Dt 27:15) in
the light of the incarnation. For the mystery of the incarnation demonstrated the possibility of
representing the God-made-man in his human and historical reality. Defense of artistic
representation of the Incarnate Word and of the events of salvation during the iconoclastic
controversies of the seventh and eighth centuries rested on a profound understanding of the
hypostatic union which refused to separate the divine and the human in the “image.”
15. Patristic and medieval theology diverged at certain points from biblical anthropology,
and developed it at other points. The majority of the representatives of the tradition, for example,
did not fully embrace the biblical vision which identified the image with the totality of man. A
significant development of the biblical account was the distinction between image and likeness,
introduced by St. Irenaeus, according to which “image” denotes an ontological participation
(methexis) and “likeness” (mimêsis) a moral transformation (Adv. Haer. V,6,1; V,8,1; V,16,2).
According to Tertullian, God created man in his image and gave him the breath of life as his
likeness. While the image can never be destroyed, the likeness can be lost by sin (Bapt. 5, 6.7). St.
Augustine did not take up this distinction, but presented a more personalistic, psychological and
existential account of the imago Dei. For him, the image of God in man has a Trinitarian structure,
reflecting either the tripartite structure of the human soul (spirit, self-consciousness, and love) or the
threefold aspects of the psyche (memory, intelligence, and will). According to Augustine, the image
of God in man orients him to God in invocation, knowledge and love (Confessions I, 1,1).
16. In Thomas Aquinas, the imago Dei possesses an historical character, since it passes
through three stages: the imago creationis (naturae), the imago recreationis (gratiae), and the
similitudinis (gloriae) (S.Th. I q.93 a.4). For Aquinas, the imago Dei is the basis for participation in
the divine life. The image of God is realized principally in an act of contemplation in the intellect
(S.Th. I q.93 a.4 and 7). This conception can be distinguished from that of Bonaventure, for whom
the image is realized chiefly through the will in the religious act of man (Sent. II d.16 a.2 q.3).
Within a similar mystical vision, but with a greater boldness, Meister Eckhart tends to spiritualize
the imago Dei by placing it at the summit of the soul and detaching it from the body (Quint. I,5,5-
7;V, 6.9s).
17. Reformation controversies demonstrated that the theology of the imago Dei remained
important for both Protestant and Catholic theologians. The Reformers accused the Catholics of
reducing the image of God to an “imago naturae” which presented a static conception of human
nature and encouraged the sinner to constitute himself before God. On the other side, the Catholics
accused the Reformers of denying the ontological reality of the image of God and reducing it to a
pure relation. In addition, the Reformers insisted that the image of God was corrupted by sin,
whereas Catholic theologians viewed sin as a wounding of the image of God in man.
2. The modern critique of the theology of the imago Dei
18. Until the dawn of the modern period, the theology of the imago Dei retained its central
position in theological anthropology. Throughout the history of Christian thought, such was the
power and fascination of this theme that it could withstand those isolated critiques (as, for example,
in iconoclasm) which charged that its anthropomorphism fostered idolatry. But, in the modern
period, the theology of the imago Dei came under a more sustained and systematic critique.
19. The view of the universe advanced by modern science displaced the classical notion of a
cosmos made in the divine image and thus dislodged an important part of the conceptual framework
supporting the theology of the imago Dei. The theme was regarded as ill-adapted to experience by
empiricists, and as ambiguous by rationalists. But more significant among the factors undermining
the theology of the imago Dei was the conception of man as a self-constituting autonomous subject,
apart from any relationship to God. With this development, the notion of the imago Dei could not be
sustained. It was but a short step from these ideas to the reversal of biblical anthropology which
took various forms in the thought of Ludwig Feuerbach, Karl Marx and Sigmund Freud: it is not
man who is made in the image of God, but God who is nothing else than an image projected by
man. In the end, atheism appeared to be required if man was to be self-constituting.
20. At first, the climate of twentieth century western theology was unfavorable to the theme
of the imago Dei. Given the nineteenth century developments just mentioned, it was perhaps
inevitable that some forms of dialectical theology regarded the theme as an expression of human
arrogance by which man compares or equates himself to God. Existential theology, with its stress
on the event of the encounter with God, undermined the notion of a stable or permanent relationship
with God which is entailed by the doctrine of the imago Dei. Secularization theology rejected the
notion of an objective reference in the world locating man with respect to God. The "God without
properties," - in effect, an impersonal God - espoused by some versions of negative theology could
not serve as the model for man made in his image. In political theology, with its overriding concern
for orthopraxis, the theme of the imago Dei receded from view. Finally, secular and theological
critics alike blamed the theology of the imago Dei for promoting a disregard of the natural
environment and animal welfare.
3. The imago Dei at Vatican Council II and in current theology
21. Despite these unfavorable trends, interest in the recovery of the theology of the imago
Dei rose steadily throughout the mid- twentieth century. Intense study of the Scriptures, of the
Fathers of the Church, and of the great scholastic theologians produced a renewed awareness of the
ubiquity and importance of the theme of the imago Dei. This recovery was well underway among
Catholic theologians before the Second Vatican Council. The council gave new impetus to the
theology of the imago Dei, most especially in the Constitution on the Church in the Modern World
Gaudium et Spes.
22. Invoking the theme of the image of God, the Council affirmed in Gaudium et Spes the
dignity of man as it is taught in Genesis 1;26 and Psalm 8:6 (GS 12). Within the conciliar vision, the
imago Dei consists in man's fundamental orientation to God, which is the basis of human dignity
and of the inalienable rights of the human person. Because every human being is an image of God,
he cannot be made subservient to any this-worldly system or finality. His sovereignty within the
cosmos, his capacity for social existence, and his knowledge and love of the Creator - all are rooted
in man's being made in the image of God. Basic to the conciliar teaching is the christological
determination of the image: it is Christ who is the image of the invisible God (Col 1:15) (GS 10).
The Son is the perfect Man who restores the divine likeness to the sons and daughters of Adam
which was wounded by the sin of the first parents (GS 22). Revealed by God who created man in
his image, it is the Son who gives to man the answers to his questions about the meaning of life and
death (GS 41). The Council also underscores the trinitarian structure of the image: by conformity to
Christ (Rm 8:29) and through the gifts of the Holy Spirit (Rm 8:23), a new man is created, capable
of fulfilling the new commandment (GS 22). It is the saints who are fully transformed in the image
of Christ (cf. 2 Cor 3:18); in them, God manifests his presence and grace as a sign of his kingdom
(GS 24). On the basis of the doctrine of the image of God, the Council teaches that human activity
reflects the divine creativity which is its model (GS 34) and must be directed to justice and human
fellowship in order to foster the establishment of one family in which all are brothers and sisters
(GS 24).
24. The renewed interest in the theology of the imago dei which emerged at the Second
Vatican Council is reflected in contemporary theology, where it is possible to note developments in
several areas. In the first place, theologians are working to show how the theology of the imago Dei
illumines the connections between anthropology and Christology. Without denying the unique grace
which comes to the human race through the incarnation, theologians want to recognize the intrinsic
value of the creation of man in God’s image. The possibilities that Christ opens up for man do not
involve the suppression of the human reality in its creatureliness but its transformation and
realization according to the perfect image of the Son. In addition, with this renewed understanding
of the link between Christology and anthropology comes a deeper understanding of the dynamic
character of the imago Dei. Without denying the gift of man’s original creation in the image of God,
theologians want to acknowledge the truth that, in the light of human history and the evolution of
human culture, the imago Dei can in a real sense be said to be still in the process of becoming. What
is more, the theology of the imago Dei also links anthropology with moral theology by showing
that, in his very being, man possesses a participation in the divine law. This natural law orients
human persons to the pursuit of the good in their actions. It follows, finally, that the imago Dei has a
teleological and eschatological dimension which defines man as homo viator, oriented to the
parousia and to the consummation of the divine plan for the universe as it is realized in the history
of grace in the life of each individual human being and in the history of the whole human race.
CHAPTER TWO
IN THE IMAGE OF GOD: PERSONS IN COMMUNION
25. Communion and stewardship are the two great strands out of which the fabric of the
doctrine of the imago Dei is woven. The first strand, which we take up in this chapter, can be
summarized in the following way: The triune God has revealed his plan to share the communion of
Trinitarian life with persons created in his image. Indeed, it is for the sake of this Trinitarian
communion that human persons are created in the divine image. It is precisely this radical likeness
to the triune God that is the basis for the possibility of the communion of creaturely beings with the
uncreated persons of the Blessed Trinity. Created in the image of God, human beings are by nature
bodily and spiritual, men and women made for one another, persons oriented towards communion
with God and with one another, wounded by sin and in need of salvation, and destined to be
conformed to Christ, the perfect image of the Father, in the power of the Holy Spirit.
1. Body and soul
26. Human beings, created in the image of God, are persons called to enjoy communion and
to exercise stewardship in a physical universe. The activities entailed by interpersonal communion
and responsible stewardship engage the spiritual - intellectual and affective - capacities of human
persons, but they do not leave the body behind. Human beings are physical beings sharing a world
with other physical beings. Implicit in the Catholic theology of the imago Dei is the profound truth
that the material world creates the conditions for the engagement of human persons with one
another.
27. This truth has not always received the attention it deserves. Present-day theology is
striving to overcome the influence of dualistic anthropologies that locate the imago Dei exclusively
with reference to the spiritual aspect of human nature. Partly under the influence first of Platonic
and later of Cartesian dualistic anthropologies, Christian theology itself tended to identify the
imago Dei in human beings with what is the most specific characteristic of human nature, viz., mind
or spirit. The recovery both of elements of biblical anthropology and of aspects of the Thomistic
synthesis has contributed to the effort in important ways.
28. The view that bodiliness is essential to personal identity is fundamental, even if not
explicitly thematized, in the witness of Christian revelation. Biblical anthropology excludes mind-
body dualism. It speaks of man as a whole. Among the basic Hebrew terms for man used in the Old
Testament, nèfèš means the life of a concrete person who is alive (Gen 9:4; Lev. 24:17-18, Proverbs
8:35). But man does not have a nèfèš; he is a nèfèš (Gen 2:7; Lev 17:10). Basar refers to the flesh of
animals and of men, and sometimes the body as a whole (Lev 4:11; 26:29). Again, one does not
have a basar, but is a basar. The New Testament term sarx (flesh) can denote the material
corporality of man (2 Cor 12:7), but on the other hand also the whole person (Rom. 8:6). Another
Greek term, soma (body) refers to the whole man with emphasis on his outward manifestation. Here
too man does not have his body, but is his body. Biblical anthropology clearly presupposes the unity
of man, and understands bodiliness to be essential to personal identity.
29. The central dogmas of the Christian faith imply that the body is an intrinsic part of the
human person and thus participates in his being created in the image of God. The Christian doctrine
of creation utterly excludes a metaphysical or cosmic dualism since it teaches that everything in the
universe, spiritual and material, was created by God and thus stems from the perfect Good. Within
the framework of the doctrine of the incarnation, the body also appears as an intrinsic part of the
person. The Gospel of John affirms that "the Word became flesh (sarx)," in order to stress, against
Docetism, that Jesus had a real physical body and not a phantom-body. Furthermore, Jesus redeems
us through every act he performs in his body. His Body which is given up for us and His Blood
which is poured out for us mean the gift of his Person for our salvation. Christ's work of redemption
is carried on in the Church, his mystical body, and is made visible and tangible through the
sacraments. The effects of the sacraments, though in themselves primarily spiritual, are
accomplished by means of perceptible material signs, which can only be received in and through the
body. This shows that not only man's mind but also his body is redeemed. The body becomes a
temple of the Holy Spirit. Finally, that the body belongs essentially to the human person is inherent
to the doctrine of the resurrection of the body at the end of time, which implies that man exists in
eternity as a complete physical and spiritual person.
 30. In order to maintain the unity of body and soul clearly taught in revelation, the
Magisterium adopted the definition of the human soul as forma substantialis (cf. Council of Vienne
and the Fifth Lateran Council). Here the Magisterium relied on Thomistic anthropology which,
drawing upon the philosophy of Aristotle, understands body and soul as the material and spiritual
principles of a single human being. It may be noted that this account is not incompatible with
present-day scientific insights. Modern physics has demonstrated that matter in its most elementary
particles is purely potential and possesses no tendency toward organization. But the level of
organization in the universe, which contains highly organized forms of living and non-living
entities, implies the presence of some "information." This line of reasoning suggests a partial
analogy between the Aristotelian concept of substantial form and the modern scientific notion of
"information." Thus, for example, the DNA of the chromosomes contains the information necessary
for matter to be organized according to what is typical of a certain species or individual.
Analogically, the substantial form provides to prime matter the information it needs to be organized
in a particular way. This analogy should be taken with due caution because metaphysical and
spiritual concepts cannot be simply compared with material, biological data.
31. These biblical, doctrinal and philosophical indications converge in the affirmation that
human bodiliness participates in the imago Dei. If the soul, created in God's image, forms matter to
constitute the human body, then the human person as a whole is the bearer of the divine image in a
spiritual as well as a bodily dimension. This conclusion is strengthened when the christological
implications of the image of God are taken fully into account. "In reality it is only in the mystery of
the Word made flesh that the mystery of man truly becomes clear….Christ fully reveals man to
himself and brings to light his most high calling" (Gaudium et Spes 22). Spiritually and physically
united to the incarnate and glorified Word, especially in the sacrament of the Eucharist, man arrives
at his destination: the resurrection of his own body and the eternal glory in which he participates as
a complete human person, body and soul, in the Trinitarian communion shared by all the blessed in
the company of heaven.
2. Man and woman
32. In Familiaris Consortio, Pope John Paul II affirmed: “As an incarnate spirit, that is a
soul which expresses itself in a body and a body informed by an immortal spirit, man is called to
love in his unified totality. Love includes the human body, and the body is made a sharer in spiritual
love” (11). Created in the image of God, human beings are called to love and communion. Because
this vocation is realized in a distinctive way in the procreative union of husband and wife, the
difference between man and woman is an essential element in the constitution of human beings
made in the image of God.
33. "God created man in his image; in the image of God he created him; male and female, he
created them" (Gen. 1:27; cf. Gen. 5:1-2). According to the Scripture, therefore, the imago Dei
manifests itself, at the outset, in the difference between the sexes. It could be said that human being
exist only as masculine or feminine, since the reality of the human condition appears in the
difference and plurality of the sexes. Hence, far from being an accidental or secondary aspect of
personality, it is constitutive of person identity. Each of us possesses a way of being in the world, to
see, to think, to feel, to engage in mutual exchange with other persons who are also defined by their
sexual identity. According to the Catechism of the Catholic Church: "Sexuality affects all aspects of
the human person in the unity of his body and soul. It especially concerns affectivity, the capacity to
love and to procreate, and in a more general way the aptitude for forming bonds of communion with
others" (2332). The roles attributed to one or the other sex may vary across time and space, but the
sexual identity of the person is not a cultural or social construction. It belongs to the specific
manner in which the imago Dei exists.
34. The incarnation of the Word reinforces this specificity. He assumed the human condition
in its totality, taking up one sex, but he became man in both senses of the term: as a member of the
human community, and as a male. The relation of each one to Christ is determined in two ways: it
depends on one’s own proper sexual identity and that of Christ.
35. In addition, the incarnation and resurrection extend the original sexual identity of the
imago Dei into eternity. The risen Lord remains a man when he sits now at the right hand of the
Father. We may also note that the sanctified and glorified person of the Mother of God, now
assumed bodily into heaven, continues to be a woman. When in Galatians 3:28, St. Paul announces
that in Christ all differences – including that between man and woman – would be erased, he is
affirming that no human differences can impede our participation in the mystery of Christ. The
Church has not followed St. Gregory of Nyssa and some other Fathers of the Church who held that
sexual differences as such would be annulled by the resurrection. The sexual differences between
man and woman, while certainly manifesting physical attributes, in fact transcend the purely
physical and touch the very mystery of the person.
36. The Bible lends no support to the notion of a natural superiority of the masculine over
the feminine sex. Their differences notwithstanding, the two sexes enjoy an inherent equality. As
Pope John Paul II wrote in Familiaris Consortio: “Above all it is important to underline the equal
dignity and responsibility of women with men. This equality is realized in a unique manner in that
reciprocal self-giving by each one to the other and by both to the children which is proper to
marriage and the family….In creating the human race ‘male and female,’ God gives man and
woman an equal personal dignity, endowing them with the inalienable rights and responsibilities
proper to the human person” (22). Man and woman are equally created in God’s image. Both are
persons, endowed with intelligence and will, capable of orienting their lives through the exercise of
freedom. But each does so in a manner proper and distinctive to their sexual identity, in such wise
that the Christian tradition can speak of a reciprocity and complementarity. These terms, which have
lately become somewhat controversial, are nonetheless useful in affirming that man and woman
each needs the other in order to achieve fullness of life.
 37. To be sure, the original friendship between man and woman was deeply impaired by sin.
Through his miracle at the wedding feast of Cana (John 2:1 ff), our Lord shows that he has come to
restore the harmony that God intended in the creation of man and woman.
38. The image of God, which is to be found in the nature of the human person as such, can
be realized in a special way in the union between human beings. Since this union is directed to the
perfection of divine love, Christian tradition has always affirmed the value of virginity and celibacy
which foster chaste friendship among human persons at the same time that they point to the
eschatological fulfillment of all created love in the uncreated love of the Blessed Trinity. In this
very connection, the Second Vatican Council drew an analogy between the communion of the
divine persons among themselves, and that which human beings are invited to establish on earth (cf.
Gaudium et Spes, 24). While it is certainly true that union between human beings can be realized in
a variety of ways, Catholic theology today affirms that marriage constitutes an elevated form of the
communion between human persons and one of the best analogies of the Trinitarian life. When a
man and a woman unite their bodies and spirits in an attitude of total openness and self-giving, they
form a new image of God. Their union as one flesh does not correspond simply to a biological
necessity, but to the intention of the Creator in leading them to share the happiness of being made in
his image. The Christian tradition speaks of marriage as an eminent way of sanctity. “God is love,
and in himself he lives a mystery of personal loving communion. Creating man and woman in his
image…, God inscribed in the humanity of man and woman the vocation, and thus the capacity and
responsibility of love and communion” (Catechism of the Catholic Church 2331). The Second
Vatican Council also underlined the profound significance of marriage: “Christian spouses, in virtue
of the sacrament of matrimony, signify and partake of the mystery of that unity and fruitful love
which exists between Christ and His Church (cf. Eph. 5:32). The spouses thereby help each other to
attain to holiness in their married life and by the rearing of their children” (Lumen Gentium 11; cf.
Gaudium et Spes 48).
3. Person and community
40. Persons created in the image of God are bodily beings whose identity as male or female
orders them to a special kind of communion with one another. As Pope John Paul II has taught, the
nuptial meaning of the body finds its realization in the human intimacy and love that mirror the
communion of the Blessed Trinity whose mutual love is poured out in creation and redemption.
This truth is at the center of Christian anthropology. Human beings are created in the imago Dei
precisely as persons capable of a knowledge and love that are personal and interpersonal. It is of the
essence of the imago Dei in them that these personal beings are relational and social beings,
embraced in a human family whose unity is at once realized and prefigured in the Church.
41. When one speaks of the person, one refers both to the irreducible identity and interiority
that constitutes the particular individual being, and to the fundamental relationship to other persons
that is the basis for human community. In the Christian perspective, this personal identity that is at
once an orientation to the other is founded essentially on the Trinity of divine Persons. God is not a
solitary being, but a communion of three Persons. Constituted by the one divine nature, the identity
of the Father is his paternity, his relation to the Son and the Spirit; the identity of the Son is his
relation to the Father and the Spirit; the identity of the Spirit is his relation to the Father and the
Son. Christian revelation led to the articulation of the concept of person, and gave it a divine,
christological, and Trinitarian meaning. In effect, no person is as such alone in the universe, but is
always constituted with others and is summoned to form a community with them.
42. It follows that personal beings are social beings as well. The human being is truly human
to the extent that he actualizes the essentially social element in his constitution as a person within
familial, religious, civil, professional, and other groups that together form the surrounding society to
which he belongs. While affirming the fundamentally social character of human existence, Christian
civilization has nonetheless recognized the absolute value of the human person as well as the
importance of individual rights and cultural diversity. In the created order, there will always be a
certain tension between the individual person and the demands of social existence. In the Blessed
Trinity there is a perfect harmony between the Persons who share the communion of a single divine
life.
43. Every individual human being as well as the whole human community are created in the
image of God. In its original unity – of which Adam is the symbol – the human race is made in the
image of the divine Trinity. Willed by God, it makes its way through the vicissitudes of human
history towards a perfect communion, also willed by God, but yet to be fully realized. In this sense,
human beings share the solidarity of a unity that both already exists and is still to be attained.
Sharing in a created human nature and confessing the triune God who dwells among us, we are
nonetheless divided by sin and await the victorious coming of Christ who will restore and recreate
the unity God wills in a final redemption of creation (cf. Rom 8:18-19). This unity of the human
family is yet to be realized eschatologically. The Church is the sacrament of salvation and of the
kingdom of God: catholic, in bringing together man of every race and culture; one, in being the
vanguard of the unity of the human community willed by God; holy, sanctified herself by the power
of the Holy Spirit, and sanctifying all men through the Sacraments; and, apostolic, in continuing the
mission of the men chosen by Christ to accomplish progressively the divinely willed unity of the
human race and the consummation of creation and redemption.
4. Sin and salvation
44. Created in the image of God to share in the communion of Trinitarian life, human beings
are persons who are so constituted as to be able freely to embrace this communion. Freedom is the
divine gift that enables human persons to choose the communion which the triune God offers to
them as their ultimate good. But with freedom comes the possibility of the failure of freedom.
Instead of embracing the ultimate good of participation in the divine life, human persons can and do
turn away from it in order to enjoy transitory or even only imaginary goods. Sin is precisely this
failure of freedom, this turning away from the divine invitation to communion.
 45. Within the perspective of the imago Dei, which is essentially dialogical or relational in
its ontological structure, sin, as a rupture of the relationship with God, causes a disfigurement of the
imago Dei. The dimensions of sin can be grasped in the light of those dimensions of the imago Dei
which are affected by sin. This fundamental alienation from God also upsets man’s relationship with
others (cf. 1 John 3:17) and, in a real sense, produces a division within himself between body and
spirit, knowing and willing, reason and emotions (Rom. 7:14 f). It also affects his physical
existence, bringing suffering, illness and death. In addition, just as the imago Dei has an historical
dimension, so too does sin. The witness of Scripture (cf. Rom. 5:12ff) presents us with a vision of
the history of sin, caused by a rejection of the divine invitation to communion which occurred at the
beginning of the history of the human race. Finally, sin affects the social dimension of the imago
Dei; it is possible to discern ideologies and structures which are the objective manifestation of sin
and which obstruct the realization of the image of God on the part of human beings.
46. Catholic and Protestant exegetes today agree that the imago Dei cannot be totally
destroyed by sin since it defines the whole structure of human nature. For its part, Catholic tradition
has always insisted that, while the imago Dei is impaired or disfigured, it cannot be destroyed by
sin. The dialogical or relational structure of the image of God cannot be lost but, under the reign of
sin, it is disrupted in its orientation towards its christological realization. Furthermore, the
ontological structure of the image, while affected in its historicity by sin, remains despite the reality
of sinful actions. In this connection – as many Fathers of the Church argued in their response to
Gnosticism and Manicheanism and -- the freedom which as such defines what it is to be human and
is fundamental to the ontological structure of the imago Dei, cannot be suppressed, even if the
situation in which freedom is exercised is in part determined by the consequences of sinfulness.
Finally, against the notion of the total corruption of the imago Dei by sin, the Catholic tradition has
insisted that grace and salvation would be illusory if they did not in fact transform the existing,
albeit sinful, reality of human nature.
47. Understood in the perspective of the theology of the imago Dei, salvation entails the
restoration of the image of God by Christ who is the perfect image of the Father. Winning our
salvation through his passion, death and resurrection, Christ conforms us to himself through our
participation in the paschal mystery and thus reconfigures the imago Dei in its proper orientation to
the blessed communion of Trinitarian life. In this perspective, salvation is nothing less than a
transformation and fulfillment of the personal life of the human being, created in the image of God
and now newly directed to a real participation in the life of the divine persons, through the grace of
the incarnation and the indwelling of the Holy Spirit. The Catholic tradition rightly speaks here of a
realization of the person. Suffering from a deficiency of charity because of sin, the person cannot
achieve self-realization apart from the absolute and gracious love of God in Christ Jesus. Through
this saving transformation of the person through Christ and the Holy Spirit, everything in the
universe is also transformed and comes to share in the glory of God (Rom. 8:21).
 48. For the theological tradition, man affected by sin is always in need of salvation, yet
having a natural desire to see God - a capax Dei - which, as an image of the divine, constitutes a
dynamic orientation to the divine. While this orientation is not destroyed by sin, neither can it be
realized apart from God's saving grace. God the savior addresses an image of himself, disturbed in
its orientation to him, but nonetheless capable of receiving the saving divine activity. These
traditional formulations affirm both the indestructibility of man's orientation to God and the
necessity of salvation. The human person, created in the image of God, is ordered by nature to the
enjoyment of divine love, but only divine grace makes the free embrace of this love possible and
effective. In this perspective, grace is not merely a remedy for sin, but a qualitative transformation
of human liberty, made possible by Christ, as a freedom freed for the Good.
49. The reality of personal sin shows that the image of God is not unambiguously open to
God but can close in upon itself. Salvation entails a liberation from this self-glorification through
the cross. The paschal mystery, which is originally constituted by the passion, death and
resurrection of Christ, makes it possible for each person to participate in the death to sin that leads
to life in Christ. The cross entails, not the destruction of the human, but the passage that leads to
new life.
50. The effects of salvation for man created in the image of God are obtained through the
grace of Christ who, as the second Adam, is the head of a new humanity and who creates for man a
new salvific situation through his death for sinners and through his resurrection (cf. 1 Cor 15:47-49;
2 Cor 5:2; Rom 5:6ff). In this way, man becomes a new creature (2 Cor 5:17) who is capable of a
new life of freedom, a life "freed from" and "freed for."
51. Man is freed from sin, from the law, and from suffering and death. In the first place,
salvation is a liberation from sin which reconciles man with God, even in the midst of a continuing
struggle against sin conducted in the power of the Holy Spirit (cf. Eph 6:10-20). In addition,
salvation is not a liberation from the law as such but from any legalism that is opposed to the Holy
Spirit (2 Cor 3:6) and to the realization of love (Rom 13:10). Salvation brings a liberation from
suffering and death which acquire new meaning as a saving participation through the suffering,
death and resurrection of the Son. In addition, according to the Christian faith, "freed from" means
"freed for": freedom from sin signifies a freedom for God in Christ and the Holy Spirit; freedom
from the law means a freedom for authentic love; freedom from death means a freedom for new life
in God. This "freedom for" is made possible by Jesus Christ, the perfect icon of the Father, who
restores the image of God in man.
5. Imago Dei and imago Christi
52. "In reality it is only in the mystery of the Word made flesh that the mystery of man truly
becomes clear. For Adam, the first man, was a type of him who was to come, Christ the Lord. Christ
the new Adam, in the very revelation of the mystery of the Father and of his love, fully reveals man
to himself and brings to light his most high calling. No wonder, then, that all the truths mentioned
so far should find in him their source and their most perfect embodiment" (Gaudium et Spes, 22).
This famous passage from the Second Vatican Council's Constitution on the Church in the Modern
World serves well to conclude this summary of the main elements of the theology of the imago Dei.
For it is Jesus Christ who reveals to man the fullness of his being, in its original nature, in its final
consummation, and in its present reality.
53. The origins of man are to be found in Christ: for he is created "through him and in him"
(Col 1:16), "the Word [who is] the life…and the light of every man who is coming into the world"
(John 1:3-4, 9). While it is true that man is created ex nihilo, it can also be said that he is created
from the fullness (ex plenitudine) of Christ himself who is at once the creator, the mediator and the
end of man. The Father destined us to be his sons and daughters, and "to be conformed to the image
of his Son, who is the firstborn of many brothers" (Rom. 8:29). Thus, what it means to be created in
the imago Dei is only fully revealed to us in the imago Christi. In him, we find the total receptivity
to the Father which should characterize our own existence, the openness to the other in an attitude
of service which should characterize our relations with our brothers and sisters in Christ, and the
mercy and love for others which Christ, as the image of the Father, displays for us.
54. Just as man's beginnings are to be found in Christ, so is his finality. Human beings are
oriented to the kingdom of Christ as to an absolute future, the consummation of human existence.
Since "all things have been created through him and for him" (Col 1:16), they find their direction
and destiny in him. The will of God that Christ should be the fullness of man is to find an
eschatological realization. While the Holy Spirit will accomplish the ultimate configuration of
human persons to Christ in the resurrection of the dead, human beings already participate in this
eschatological likeness to Christ here below, in the midst of time and history. Through the
Incarnation, Resurrection and Pentecost, the eschaton is already here; they inaugurate it and
introduce it into the world of men, and anticipate its final realization. The Holy Spirit works
mysteriously in all human beings of good will, in societies and in the cosmos to transfigure and
divinize human beings. Moreover, the Holy Spirit works through all the sacraments, particularly the
Eucharist which is the anticipation of the heavenly banquet, the fullness of communion in the
Father, Son and Holy Spirit.
55. Between the origins of man and his absolute future lies the present existential situation
of the human race whose full meaning is likewise to be found only in Christ. We have seen that it is
Christ - in his incarnation, death and resurrection - who restores the image of God in man to its
proper form. "Through him, God was pleased to reconcile to himself all things, whether on earth or
in heaven, by making peace through the blood of his cross"(Col 1:20). At the core of his sinful
existence, man is pardoned and, through the grace of the Holy Spirit, he knows that he is saved and
justified through Christ. Human beings grow in their resemblance to Christ and collaborate with the
Holy Spirit who, especially through the sacraments, fashions them in the image of Christ. In this
way, man's everyday existence is defined as an endeavor to be conformed ever more fully to the
image of Christ and to dedicate his life to the struggle to bring about the final victory of Christ in
the world.
 CHAPTER THREE
IN THE IMAGE OF GOD: STEWARDS OF VISIBLE CREATION
56. The first great theme within the theology of the imago Dei concerns participation in the
life of divine communion. Created in the image of God, as we have seen, human beings are beings
who share the world with other bodily beings but who are distinguished by their intellect, love and
freedom and are thus ordered by their very nature to interpersonal communion. The prime instance
of this communion is the procreative union of man and woman which mirrors the creative
communion of Trinitarian love. The disfigurement of the imago Dei by sin, with its inevitably
disruptive consequences for personal and interpersonal life, is overcome by the passion, death and
resurrection of Christ. The saving grace of participation in the paschal mystery reconfigures the
imago Dei according to the pattern of the imago Christi.
57. In the present chapter, we consider the second of the main themes of the theology of the
imago Dei. Created in the image of God to share in the communion of Trinitarian love, human
beings occupy a unique place in the universe according to the divine plan: they enjoy the privilege
of sharing in the divine governance of visible creation. This privilege is granted to them by the
Creator who allows the creature made in his image to participate in his work, in his project of love
and salvation, indeed in his own lordship over the universe. Since man's place as ruler is in fact a
participation in the divine governance of creation, we speak of it here as a form of stewardship.
 58. According to Gaudium et Spes: "Man was created in God's image and was commanded
to conquer the earth and to rule the world in justice and holiness: he was to acknowledge God as
maker of all things and relate himself and the totality of creation to him, so that through the
dominion of all things by man the name of God would be majestic in all the earth" (34). This
concept of man's rule or sovereignty plays an important role in Christian theology. God appoints
man as his steward in the manner of the master in the Gospel parables (cf. Luke 19:12). The only
creature willed expressly by God for his own sake occupies a unique place at the summit of visible
creation (Gen. 1:26; 2:20; Ps 8:6-7, Wisdom 9:2-3).
59. Christian theology uses both domestic and royal imagery to describe this special role.
Employing royal imagery, it is said that human beings are called to rule in the sense of holding an
ascendancy over the whole of visible creation, in the manner of a king. But the inner meaning of
this kingship is, as Jesus reminds his disciples, one of service: only by willingly suffering as a
sacrificial victim does Christ become the king of the universe, with the Cross as his throne.
Employing domestic imagery, Christian theology speaks of man as the master of a household to
whom God has confided care of all his goods (cf. Mt 24:45). Man can deploy all the resources of
visible creation according to his ingenuity, and exercises this participated sovereignty over visible
creation in through science, technology and art.
60. Above himself and yet in the intimacy of his own conscience, man discovers the
existence of a law which the tradition calls the "natural law." This law is of divine origin, and man's
awareness of it is itself a participation in the divine law. It refers man to the true origins of the
universe as well as to his own (Veritatis Splendor, 20). This natural law drives the rational creature
to search for the truth and the good in his sovereignty of the universe. Created in the image of God,
man exercises this sovereignty over visible creation only in virtue of the privilege conferred upon
him by God. He imitates the divine rule, but he cannot displace it. The Bible warns against the sin
of this usurpation of the divine role. It is a grave moral failure for human beings to act as rulers of
visible creation who separate themselves from the higher, divine law. They act in place of the
master as stewards (cf. Mt 25:14 ff) who have the freedom they need to develop the gifts which
have been confided to them and to do so with a certain bold inventiveness.
61. The steward must render an account of his stewardship, and the divine Master will judge
his actions. The moral legitimacy and efficacy of the means employed by the steward provide the
criteria for this judgment. Neither science nor technology are ends in themselves; what is
technically possible is not necessarily also reasonable or ethical. Science and technology must be
put in the service of the divine design for the whole of creation and for all creatures. This design
gives meaning to the universe and to human enterprise as well. Human stewardship of the created
world is precisely a stewardship exercised by way of participation in the divine rule and is always
subject to it. Human beings exercise this stewardship by gaining scientific understanding of the
universe, by caring responsibly for the natural world (including animals and the environment), and
by guarding their own biological integrity.
 1. Science and the stewardship of knowledge
62. The endeavor to understand the universe has marked human culture in every period and
in nearly every society. In the perspective of the Christian faith, this endeavor is precisely an
instance of the stewardship which human beings exercise in accordance with God's plan. Without
embracing a discredited concordism, Christians have the responsibility to locate the modern
scientific understanding of the universe within the context of the theology of creation. The place of
human beings in the history of this evolving universe, as it has been charted by modern sciences,
can only be seen in its complete reality in the light of faith, as a personal history of the engagement
of the triune God with creaturely persons.
63. According to the widely accepted scientific account, the universe erupted 15 billion
years ago in an explosion called the “Big Bang” and has been expanding and cooling ever since.
Later there gradually emerged the conditions necessary for the formation of atoms, still later the
condensation of galaxies and stars, and about 10 billion years later the formation of planets. In our
own solar system and on earth (formed about 4.5 billion years ago), the conditions have been
favorable to the emergence of life. While there is little consensus among scientists about how the
origin of this first microscopic life is to be explained, there is general agreement among them that
the first organism dwelt on this planet about 3.5-4 billion years ago. Since it has been demonstrated
that all living organisms on earth are genetically related, it is virtually certain that all living
organisms have descended from this first organism. Converging evidence from many studies in the
physical and biological sciences furnishes mounting support for some theory of evolution to
account for the development and diversification of life on earth, while controversy continues over
the pace and mechanisms of evolution. While the story of human origins is complex and subject to
revision, physical anthropology and molecular biology combine to make a convincing case for the
origin of the human species in Africa about 150,000 years ago in a humanoid population of
common genetic lineage. However it is to be explained, the decisive factor in human origins was a
continually increasing brain size, culminating in that of homo sapiens. With the development of the
human brain, the nature and rate of evolution were permanently altered: with the introduction of the
uniquely human factors of consciousness, intentionality, freedom and creativity, biological
evolution was recast as social and cultural evolution.
64. Pope John Paul II stated some years ago that “new knowledge leads to the recognition of
the theory of evolution as more than a hypothesis. It is indeed remarkable that this theory has been
progressively accepted by researchers following a series of discoveries in various fields of
knowledge”(“Message to the Pontifical Academy of Sciences on Evolution”1996). In continuity
with previous twentieth century papal teaching on evolution (especially Pope Pius XII’s encyclical
Humani Generis ), the Holy Father’s message acknowledges that there are “several theories of
evolution” that are “materialist, reductionist and spiritualist” and thus incompatible with the
Catholic faith. It follows that the message of Pope John Paul II cannot be read as a blanket
approbation of all theories of evolution, including those of a neo-Darwinian provenance which
explicitly deny to divine providence any truly causal role in the development of life in the universe.
Mainly concerned with evolution as it “involves the question of man,” however, Pope John Paul’s
message is specifically critical of materialistic theories of human origins and insists on the
relevance of philosophy and theology for an adequate understanding of the “ontological leap” to the
human which cannot be explained in purely scientific terms. The Church’s interest in evolution thus
focuses particularly on “the conception of man” who, as created in the image of God, “cannot be
subordinated as a pure means or instrument either to the species or to society.” As a person created
in the image of God, he is capable of forming relationships of communion with other persons and
with the triune God, as well as of exercising sovereignty and stewardship in the created universe.
The implication of these remarks is that theories of evolution and of the origin of the universe
possess particular theological interest when they touch on the doctrines of the creation ex nihilo and
the creation of man in the image of God.
65. We have seen human persons are created in the image of God in order to become
partakers of the divine nature (cf. 2 Pet 1:3-4) and thus to share in the communion of trinitarian life
and in the divine dominion over visible creation. At the heart of the divine act of creation is the
divine desire to make room for created persons in the communion of the uncreated Persons of the
Blessed Trinity through adoptive participation in Christ. What is more, the common ancestry and
natural unity of the human race are the basis for a unity in grace of redeemed human persons under
the headship of the New Adam in the ecclesial communion of human persons united with one
another and with the uncreated Father, Son, and Holy Spirit. The gift of natural life is the basis for
the gift of the life of grace. It follows that, where the central truth concerns a person acting freely, it
is impossible to speak of a necessity or an imperative to create, and it is, in the end, inappropriate to
speak of the Creator as a force, or energy, or ground. Creation ex nihilo is the action of a
transcendent personal agent, acting freely and intentionally, with a view toward the all-
encompassing purposes of personal engagement. In Catholic tradition, the doctrine of the origin of
human beings articulates the revealed truth of this fundamentally relational or personalist
understanding of God and of human nature. The exclusion of pantheism and emanationism in the
doctrine of creation can be interpreted at root as a way of protecting this revealed truth. The
doctrine of the immediate or special creation of each human soul not only addresses the ontological
discontinuity between matter and spirit, but also establishes the basis for a divine intimacy which
embraces every single human person from the first moment of his or her existence.
66. The doctrine of creatio ex nihilo is thus a singular affirmation of the truly personal
character of creation and its order toward a personal creature who is fashioned as the imago Dei and
who responds not to a ground, force or energy, but to a personal creator. The doctrines of the imago
Dei and the creatio ex nihilo teach us that the existing universe is the setting for a radically
personal drama, in which the triune Creator calls out of nothingness those to whom He then calls
out in love. Here lies the profound meaning of the words of Gaudium et Spes: “Man is the only
creature on earth that God willed for his own sake” (24). Created in God’s image, human beings
assume a place of responsible stewardship in the physical universe. Under the guidance of divine
providence and acknowledging the sacred character of visible creation, the human race reshapes the
natural order, and becomes an agent in the evolution of the universe itself. In exercising their
stewardship of knowledge, theologians have the responsibility to locate modern scientific
understandings within a Christian vision of the created universe.
67. With respect to the creatio ex nihilo, theologians can note that the Big Bang theory does
not contradict this doctrine insofar as it can be said that the supposition of an absolute beginning is
not scientifically inadmissible. Since the Big Bang theory does not in fact exclude the possibility of
an antecedent stage of matter, it can be noted that the theory appears to provide merely indirect
support for the doctrine of creatio ex nihilo which as such can only be known by faith.
68. With respect to the evolution of conditions favorable to the emergence of life, Catholic
tradition affirms that, as universal transcendent cause, God is the cause not only of existence but
also the cause of causes. God’s action does not displace or supplant the activity of creaturely causes,
but enables them to act according to their natures and, nonetheless, to bring about the ends he
intends. In freely willing to create and conserve the universe, God wills to activate and to sustain in
act all those secondary causes whose activity contributes to the unfolding of the natural order which
he intends to produce. Through the activity of natural causes, God causes to arise those conditions
required for the emergence and support of living organisms, and, furthermore, for their reproduction
and differentiation. Although there is scientific debate about the degree of purposiveness or design
operative and empirically observable in these developments, they have de facto favored the
emergence and flourishing of life. Catholic theologians can see in such reasoning support for the
affirmation entailed by faith in divine creation and divine providence. In the providential design of
creation, the triune God intended not only to make a place for human beings in the universe but
also, and ultimately, to make room for them in his own trinitarian life. Furthermore, operating as
real, though secondary causes, human beings contribute to the reshaping and transformation of the
universe.
69. The current scientific debate about the mechanisms at work in evolution requires
theological comment insofar as it sometimes implies a misunderstanding of the nature of divine
causality. Many neo-Darwinian scientists, as well as some of their critics, have concluded that, if
evolution is a radically contingent materialistic process driven by natural selection and random
genetic variation, then there can be no place in it for divine providential causality. A growing body
of scientific critics of neo-Darwinism point to evidence of design (e.g., biological structures that
exhibit specified complexity) that, in their view, cannot be explained in terms of a purely contingent
process and that neo-Darwinians have ignored or misinterpreted. The nub of this currently lively
disagreement involves scientific observation and generalization concerning whether the available
data support inferences of design or chance, and cannot be settled by theology. But it is important to
note that, according to the Catholic understanding of divine causality, true contingency in the
created order is not incompatible with a purposeful divine providence. Divine causality and created
causality radically differ in kind and not only in degree. Thus, even the outcome of a truly
contingent natural process can nonetheless fall within God’s providential plan for creation.
According to St. Thomas Aquinas: “The effect of divine providence is not only that things should
happen somehow, but that they should happen either by necessity or by contingency. Therefore,
whatsoever divine providence ordains to happen infallibly and of necessity happens infallibly and
of necessity; and that happens from contingency, which the divine providence conceives to happen
from contingency” (Summa theologiae, I, 22,4 ad 1). In the Catholic perspective, neo-Darwinians
who adduce random genetic variation and natural selection as evidence that the process of evolution
is absolutely unguided are straying beyond what can be demonstrated by science. Divine causality
can be active in a process that is both contingent and guided. Any evolutionary mechanism that is
contingent can only be contingent because God made it so. An unguided evolutionary process – one
that falls outside the bounds of divine providence – simply cannot exist because “the causality of
God, Who is the first agent, extends to all being, not only as to constituent principles of species, but
also as to the individualizing principles....It necessarily follows that all things, inasmuch as they
participate in existence, must likewise be subject to divine providence” (Summa theologiae I, 22, 2).
70. With respect to the immediate creation of the human soul, Catholic theology affirms that
particular actions of God bring about effects that transcend the capacity of created causes acting
according to their natures. The appeal to divine causality to account for genuinely causal as distinct
from merely explanatory gaps does not insert divine agency to fill in the “gaps” in human scientific
understanding (thus giving rise to the so-called "God of the gaps”). The structures of the world can
be seen as open to non-disruptive divine action in directly causing events in the world. Catholic
theology affirms that that the emergence of the first members of the human species (whether as
individuals or in populations) represents an event that is not susceptible of a purely natural
explanation and which can appropriately be attributed to divine intervention. Acting indirectly
through causal chains operating from the beginning of cosmic history, God prepared the way for
what Pope John Paul II has called “an ontological leap...the moment of transition to the spiritual.”
While science can study these causal chains, it falls to theology to locate this account of the special
creation of the human soul within the overarching plan of the triune God to share the communion of
trinitarian life with human persons who are created out of nothing in the image and likeness of God,
and who, in his name and according to his plan, exercise a creative stewardship and sovereignty
over the physical universe.
2. Responsibility for the created world
71. Accelerated scientific and technological advances over the past one hundred and fifty
years have produced a radically new situation for all living things on our planet. Along with the
material abundance, higher living standards, better health and longer life spans have come air and
water pollution, toxic industrial wastage, exploitation and sometimes destruction of delicate
habitats. In this situation, human beings have developed a heightened awareness that they are
organically linked with other living beings. Nature has come to be seen as a biosphere in which all
living things form a complex yet carefully organized network of life. Moreover, it has now been
recognized that there are limits both to nature's resourcefulness and to its capacity to recover from
the harms produced by relentless exploitation of its resources.
72. An unfortunate aspect of this new ecological awareness is that Christianity has been
accused by some as in part responsible for the environmental crisis, for the very reason that it has
maximized the place of human beings created in the image of God to rule of visible creation. Some
critics go so far as to claim that the Christian tradition lacks the resources to field a sound ecological
ethics because it regards man as essentially superior to the rest of the natural world, and that it will
be necessary to turn to Asian and traditional religions to develop the needed ecological ethics.
73. But this criticism arises from a profound misunderstanding of the Christian theology of
creation and of the imago Dei. Speaking of the need for an “ecological conversion,” Pope John Paul
II remarked: “Man’s lordship is not absolute, but ministerial,…not the mission of an absolute and
unquestionable master, but of a steward of God’s kingdom” (Discourse, January 17, 2001). A
misunderstanding of this teaching may have led some to act in reckless disregard of the natural
environment, but it is no part of the Christian teaching about creation and the imago Dei to
encourage unrestrained development and possible depletion of the earth’s resources. Pope John Paul
II’s remarks reflect a growing concern with the ecological crisis on the part of the Magisterium
which is rooted in a long history of teaching found in the social encyclicals of the modern papacy.
In the perspective of this teaching, the ecological crisis is a human and a social problem, connected
with the infringement of human rights and unequal access to the earth’s resources. Pope John Paul
II summarized this tradition of social teaching when he wrote in Centesimus Annus: “Equally
worrying is the ecological question which accompanies the problem of consumerism and which is
closely connected to it. In their desire to have and to enjoy rather than to be and grow, people
consume the resources of the earth and their own lives in an excessive and disordered way. At the
root of senseless destruction of the natural environment lies an anthropological error, which
unfortunately is widespread in our day. Humankind, which discovers its capacity to transform and
in a certain sense create the world through its own work, forgets that this is always based on God’
prior and original gift of the things that are” (37).
74. The Christian theology of creation contributes directly to the resolution of the ecological
crisis by affirming the fundamental truth that visible creation is itself a divine gift, the “original
gift,” that establishes a “space” of personal communion. Indeed, we could say that a properly
Christian theology of ecology is an application of the theology of creation. Noting that the term
“ecology” combines the two Greek words oikos (house) and logos (word), the physical environment
of human existence can be conceived us a kind of “house” for human life. Given that the inner life
of the Blessed Trinity is one of communion, the divine act of creation is the gratuitous production of
partners to share in this communion. In this sense, one can say that the divine communion now
finds itself “housed” in the created cosmos. For this reason, we can speak of the cosmos as a place
of personal communion.
 75. Christology and eschatology together serve to make this truth even more profoundly
clear. In the hypostatic union of the Person of the Son with a human nature, God comes into the
world and assumes the bodiliness which he himself created. In the incarnation, through the only
begotten Son who was born of a Virgin by the power of the Holy Spirit, the triune God establishes
the possibility of an intimate personal communion with human beings. Since God graciously
intends to elevate creaturely persons to dialogical participation in his life, he has, so to speak, come
down to the creaturely level. Some theologians speak of this divine condescension as a kind of
“hominization” by which God freely makes possible our divinization. God not only manifests his
glory in the cosmos through theophanic acts, but also by assuming its bodiliness. In this
christological perspective, God’s “hominization” is his act of solidarity, not only with creaturely
persons, but with the entire created universe and its historical destiny. What is more, in the
perspective of eschatology, the second coming of Christ may be seen as the event of God’s physical
indwelling in the perfected universe which consummates the original plan of creation.
76. Far from encouraging a recklessly homocentric disregard of the natural environment, the
theology of the imago Dei affirms man’s crucial role in sharing in the realization of this eternal
divine indwelling in the perfect universe. Human beings, by God’s design, are the stewards of this
transformation for which all creation longs. Not only human beings, but the whole of visible
creation, are called to participate in the divine life. “We know that all creation is groaning in labor
pains even until now; and not only that, but we ourselves, who have the first fruits of the Spirit, we
also groan with ourselves as we wait for adoption, the redemption of our bodies” (Rm 8:23). In the
Christian perspective, our ethical responsibility for the natural environment – our “housed
existence” – is thus rooted in a profound theological understanding of visible creation and our place
within it.
77. Referring to this responsibility in an important passage in Evangelium Vitae, Pope John
Paul II wrote: “As one called to till and look after the garden of the world (cf. Gen 2:15), man has a
specific responsibility towards the environment in which he lives, towards the creation which God
has put at the service of his personal dignity. It is the ecological question – ranging from the
preservation of the natural habitats of the different species of animals and other forms to “human
ecology” properly speaking – which one finds in the Bible a clear and strong ethical direction
leading to a solution which respects the great good of life, of every life….When it comes to the
natural world, we are subject not only to biological laws but also to moral ones, which cannot be
violated with impunity” (42).
78. In the end, we must note that theology will not be able to provide us with a technical
recipe for the resolution of the ecological crisis, but, as we have seen, it can help us to see our
natural environment as God sees it, as the space of personal communion in which human beings,
created in the image of God, must seek communion with one another and the final perfection of the
visible universe.
 79. This responsibility extends to the animal world. Animals are the creatures of God, and,
according to the Scriptures, he surrounds them with his providential care (Mt 6:26). Human beings
should accept them with gratitude and, even adopting a eucharistic attitude with regard to every
element of creation, to give thanks to God for them. By their very existence the animals bless God
and give him glory: “Bless the Lord, all you birds of the air. All you beasts, wild and tame, bless the
Lord” (Dn 3:80-81). In addition, the harmony which man must establish, or restore, in the whole of
creation includes his relationship to the animals. When Christ comes in his glory, he will
“recapitulate” the whole of creation in an eschatological and definitive moment of harmony.
80. Nonetheless, there is an ontological difference between human beings and animals
because only man is created in the image of God and God has given him sovereignty over the
animal world (Gen. 1:26,28; Gen. 2: 19-20). Reflecting the Christian tradition about a just use of the
animals, the Catechism of the Catholic Church affirms: “God entrusted animals to the stewardship
of those whom he created in his own image. Hence it is legitimate to use animals for food and
clothing. They may be domesticated to help man in his work and leisure” (2417). This passage also
recalls the legitimate use of animals for medical and scientific experimentation, but always
recognizing that it is "contrary to human dignity to cause animals to suffer needlessly" (2418).
Thus, any use of animals must always be guided by the principles already articulated: human
sovereignty over the animal world is essentially a stewardship for which human beings must give an
account to God who is the lord of creation in the truest sense.
 3. Responsibility for the biological integrity of human beings
81. Modern technology, along with the latest developments in biochemistry and molecular
biology, continues to provide contemporary medicine with new diagnostic and therapeutic
possibilities. These techniques not only offer new and more effective treatments for disease,
however, but also the potential to alter man himself. The availability and feasibility of these
technologies lend new urgency to the question, how far is man allowed to remake himself? The
exercise of a responsible stewardship in the area of bioethics requires profound moral reflection on
a range of technologies that can affect the biological integrity of human beings. Here, we can offer
only some brief indications of the specific moral challenges posed by the new technologies and
some of the principles which must be applied if we are to exercise a responsible stewardship over
the biological integrity of human beings created in the image of God.
82. The right fully to dispose of the body would imply that the person may use the body as a
means to an end he himself has chosen: i.e., that he may replace its parts, modify or terminate it. In
other words, a person could determine the finality or teleological value of the body. A right to
dispose of something extends only to objects with a merely instrumental value, but not to objects
which are good in themselves, i.e., ends in themselves. The human person, being created in the
image of God, is himself such a good. The question, especially as it arises in bioethics, is whether
this also applies to the various levels that can be distinguished in the human person: the biological-
somatic, the emotional and the spiritual levels.
 83. Everyday clinical practice generally accepts a limited form of disposing of the body and
certain mental functions in order to preserve life, as for example in the case of the amputation of
limbs or the removal of organs. Such practice is permitted by the principle of totality and integrity
(also known as the therapeutic principle). The meaning of this principle is that the human person
develops, cares for, and preserves all his physical and mental functions in such a way that (1) lower
functions are never sacrificed except for the better functioning of the total person, and even then
with an effort to compensate for what is being sacrificed; and (2) the fundamental faculties which
essentially belong to being human are never sacrificed, except when necessary to save life.
84. The various organs and limbs together constituting a physical unity are, as integral parts,
completely absorbed in the body and subordinate to it. But lower values cannot simply be sacrificed
for the sake of higher ones: these values together constitute an organic unity and are mutually
dependent. Because the body, as an intrinsic part of the human person, is good in itself, fundamental
human faculties can only be sacrificed to preserve life. After all, life is a fundamental good that
involves the whole of the human person. Without the fundamental good of life, the values – like
freedom—that are in themselves higher than life itself also expire. Given that man was also created
in God’s image in his bodiliness, he has no right of full disposal of his own biological nature. God
himself and the being created in his image cannot be the object of arbitrary human action.
85. For the application of the principle of totality and integrity, the following conditions
must be met: (1) there must be a question of an intervention in the part of the body that is either
affected or is the direct cause of the life-threatening situation; (2) there can be no other alternatives
for preserving life; (3) there is a proportionate chance of success in comparison with drawbacks;
and (4) the patient must give assent to the intervention. The unintended drawbacks and side-effects
of the intervention can be justified on the basis of the principle of double effect.
86. Some have attempted to interpret this hierarchy of values to permit the sacrifice of lower
functions, like the procreative capacity, for the sake of higher values, like preserving mental health
and improving relationships with others. However, the reproductive faculty is here sacrificed in
order to preserve elements that may be essential to the person as a functioning totality but are not
essential to the person as a living totality. In fact, the person as a functioning totality is actually
violated by the loss of the reproductive faculty, and at a moment when the threat to his mental
health is not imminent and could be averted in another way. Furthermore, this interpretation of the
principle of totality suggests the possibility of sacrificing a part of the body for the sake of social
interests. On the basis of the same reasoning, sterilization for eugenic reasons could be justified on
the basis of the interest of the state.
87. Human life is the fruit of conjugal love – the mutual, total, definitive, and exclusive gift
of man and woman to one another – reflecting the mutual gift in love between the three Divine
Persons which becomes fruitful in creation, and the gift of Christ to his Church which becomes
fruitful in the rebirth of man. The fact that a total gift of man concerns both his spirit and his body is
the basis for the inseparability of the two meanings of the conjugal act which is (1) the authentic
expression of conjugal love on the physical level and (2) comes to completion through procreation
during the woman’s fertile phase (Humanae vitae, 12; Familiaris consortio, 32).
88. The mutual gift of man and woman to one another on the level of sexual intimacy is
rendered incomplete through contraception or sterilization. Furthermore, if a technique is used that
does not assist the conjugal act in attaining its goal, but replaces it, and the conception is then
effected through the intervention of a third party, then the child does not originate from the conjugal
act which is the authentic expression of the mutual gift of the parents.
89. In the case of cloning – the production of genetically identical individuals by means of
cleaving of embryos or nuclear transplantation – the child is produced asexually and is in no way to
be regarded as the fruit of a mutual gift of love. Cloning, certainly if it involves the production of a
large number of people from one person, entails an infringement of the identity of the person.
Human community, which as we have seen is also to be conceived as an image of the triune God,
expresses in its variety something of the relations of the three divine Persons in their uniqueness
which, through being of the same nature, marks their mutual differences.
90. Germ line genetic engineering with a therapeutic goal in man would in itself be
acceptable were it not for the fact that is it is hard to imagine how this could be achieved without
disproportionate risks especially in the first experimental stage, such as the huge loss of embryos
and the incidence of mishaps, and without the use of reproductive techniques. A possible alternative
would be the use of gene therapy in the stem cells that produce a man’s sperm, whereby he can
beget healthy offspring with his own seed by means of the conjugal act.
91. Enhancement genetic engineering aims at improving certain specific characteristics. The
idea of man as “co-creator” with God could be used to try to justify the management of human
evolution by means of such genetic engineering. But this would imply that man has full right of
disposal over his own biological nature. Changing the genetic identity of man as a human person
through the production of an infrahuman being is radically immoral. The use of genetic
modification to yield a superhuman or being with essentially new spiritual faculties is unthinkable,
given that the spiritual life principle of man – forming the matter into the body of the human person
– is not a product of human hands and is not subject to genetic engineering. The uniqueness of each
human person, in part constituted by his biogenetic characteristics and developed through nurture
and growth, belongs intrinsically to him and cannot be instrumentalized in order to improve some of
these characteristics. A man can only truly improve by realizing more fully the image of God in him
by uniting himself to Christ and in imitation of him. Such modifications would in any case violate
the freedom of future persons who had no part in decisions that determine his bodily structure and
characteristics in a significant and possibly irreversible way. Gene therapy, directed to the
alleviation of congenital conditions like Down's syndrome, would certainly affect the identity of the
person involved with regard to his appearance and mental gifts, but this modification would help the
individual to give full expression to his real identity which is blocked by a defective gene.
 92. Therapeutic interventions serve to restore the physical, mental and spiritual functions,
placing the person at the center and fully respecting the finality of the various levels in man in
relation to those of the person. Possessing a therapeutic character, medicine that serves man and his
body as ends in themselves respects the image of God in both. According to the principle of
proportionality, extraordinary life-prolonging therapies must be used when there is a just proportion
between the positive results that attend these therapies and possible damage to the patient himself.
Therapy may be abandoned, even if death is thereby hastened, when this proportion is absent. A
hastening of death in palliative therapy by the administration of analgesics is an indirect effect
which, like all side-effects in medicine, can come under the principle of double effect, provided that
the dosage is geared to the suppression of painful symptoms and not to the active termination of
life.
93. Disposing of death is I n reality the most radical way of disposing of life. In assisted
suicide, direct euthanasia, and direct abortion - however tragic and complex personal situations may
be - physical life is sacrificed for a self-selected finality. In the same category is the
instrumentalization of the embryo through non-therapeutic experimentation on embryos, as well as
by pre-implantation diagnostics.
94. Our ontological status as creatures made in the image of God imposes certain limits on
our ability to dispose of ourselves. The sovereignty we enjoy is not an unlimited one: we exercise a
certain participated sovereignty over the created world and, in the end, we must render an account
of our stewardship to the Lord of the Universe. Man is created in the image of God, but he is not
God himself.

CONCLUSION
95. Throughout these reflections, the theme of the imago Dei has demonstrated its
systematic power in clarifying many truths of the Christian faith. It helps us to present a relational -
and indeed personal - conception of human beings. It is precisely this relationship with God which
defines human beings and founds their relationships with other creatures. Nonetheless, as we have
seen, the mystery of the human is made fully clear only in the light of Christ who is the perfect
image of the Father and who introduces us, through the Holy Spirit, to a participation in the mystery
of the triune God. It is within this communion of love that the mystery of all being, as embraced by
God, finds its fullest meaning. At one and the same time grand and humble, this conception of
human being as the image of God constitutes a charter for human relations with the created world
and a basis upon which to assess the legitimacy of scientific and technical progress that has a direct
impact on human life and the environment. In these areas, just as human persons are called to give
witness to their participation in the divine creativity, they are also required to acknowledge their
position as creatures to whom God has confided a precious responsibility for the stewardship of the
physical universe.
 * Preliminary Note
The theme of “man created in the image of God” was submitted for study to the
International Theological Commission. The preparation of this study was entrusted to a
subcommission whose members included: Very Rev. J. Augustine Di Noia, O.P., Most Reverend
Jean-Louis Bruguès, Msgr. Anton Strukelj, Rev. Tanios Bou Mansour, O.L.M., Rev. Adolpe Gesché,
Most Reverend Willem Jacobus Eijk, Rev. Fadel Sidarouss, S.J., and Rev. Shun ichi Takayanagi,
S.J.
As the text developed, it was discussed at numerous meetings of the subcommission and
several plenary sessions of the International Theological Commission held at Rome during the
period 2000-2002. The present text was approved in forma specifica, by the written ballots of the
International Theological Commission. It was then submitted to Joseph Cardinal Ratzinger, the
President of the Commission, who has give his permission for its publication.

W000
L’Église et l’évolution
Message du pape Jean Paul II à l’Académie Pontificale des
sciences

AUX MEMBRES DE L’ACADÉMIE PONTIFICALE DES SCIENCES, RÉUNIS EN

ASSEMBLÉE PLÉNIÈRE.

C’est avec un grand plaisir que je vous adresse un cordial salut, à vous, Monsieur le
Président, et à vous tous qui constituez l’Académie pontificale des Sciences, à l’occasion
de votre Assemblée plénière.
J’adresse en particulier mes vœux aux nouveaux Académiciens (1), venus prendre part à
vos travaux pour la première fois. Je tiens aussi à évoquer Académiciens décédés au
cours de l’année écoulée, que je confie au Maître de la vie.

1. En célébrant le soixantième anniversaire de la refondation de l’Académie, il me plaît de

rappeler les intentions de mon prédécesseur Pie XI, qui voulut s’entourer d’un groupe
choisi de savants en attendant d’eux qu’ils informent le Saint-Siège en toute liberté sur les
développements de la recherche scientifique et qu’ils l’aident ainsi dans ses réflexions. À
ceux qu’il aimait appeler le Senatus scientificus de l’Église, il demanda de servir la vérité.
C’est la même intention que je vous renouvelle aujourd’hui, avec la certitude que nous
pourrons tous tirer profit de la " fécondité d’un dialogue confiant entre l’Eglise et la science
" (Discours à l’Académie des Sciences, 28 octobre 1986, n. 1).

2. Je me réjouis du premier thème que vous avez choisi, celui de l’origine de la vie et de
l’évolution, un thème essentiel qui intéresse vivement l’Eglise, puisque la Révélation
contient, de son côté, des enseignements concernant la nature et les origines de l’homme.
Comment les conclusions auxquelles aboutissent les diverses disciplines scientifiques et
celles qui sont contenues dans le message de la Révélation se rencontrent-elles ? Et si, à
première vue, il peut sembler que l’on se heurte à des oppositions, dans quelle direction
chercher leur solution ? Nous savons en effet que la vérité ne peut pas contredire la vérité
(cf. Léon XIII, Encyclique Providentissimus Deus).D’ailleurs, pour mieux éclairer la vérité
historique, vos recherches sur les rapports de l'Eglise avec la science entre le XVI e et le
XVIII e siècles sont d’une grande importance.

Au cours de cette session plénière, vous menez une " réflexion sur la science à l’aube du
troisième millénaire ", en commençant par déterminer les principaux problèmes engendrés
par les sciences, qui ont une incidence sur avenir de l’humanité. Par votre démarche, vous
jalonnez les voies de solutions qui seront bénéfiques pour toute la communauté humaine.
Dans le domaine de la nature inanimée et animée, l’évolution de la science et de ses
applications fait naître des interrogations nouvelles. L’Eglise pourra en saisir la portée
d’autant mieux qu’elle en connaîtra les aspects essentiels. Ainsi, selon sa mission
spécifique, elle pourra offrir des critères pour discerner les comportements moraux
auxquels tout homme est appelé en vue de son salut intégral.

3. Avant de vous proposer quelques réflexions plus spécialement sur le thème de l’origine
de la vie et de l’évolution, je voudrais rappeler que le Magistère de l’Eglise a déjà été
amené à se prononcer sur ces matières dans le cadre de sa propre compétence. Je citerai
ici deux interventions. Dans son Encyclique Humani generis (1950), mon prédécesseur
Pie XII avait déjà affirmé qu’il n’y avait pas d’opposition entre l’évolution et la doctrine de la
foi sur l’homme et sur sa vocation, à condition de ne pas perdre de vue quelques points
fermes (cf. AAS 42 [1950], p. 575-576).

Pour ma part, en recevant le 31 octobre 1992 les participants à l’Assemblée plénière de

votre Académie, j’ai eu l’occasion, à propos de Galilée, d’attirer l’attention sur la nécessité,
pour l’interprétation correcte de la Parole inspirée, d’une herméneutique rigoureuse. Il
convient de bien délimiter le sens propre de l’Écriture, en écartant des interprétations
indues qui lui font dire ce qu’il n’est pas dans son intention de dire. Pour bien marquer le
champ de leur objet propre, l’exégète et le théologien doivent se tenir informés des
résultats auxquels conduisent les sciences de la nature (cf. AAS 85 [1993], p. 764-772 ;
Discours à la Commission biblique pontificale, annonçant le document sur l’Interprétation
de la Bible dans l’Église : AAS 86 [1994], p. 232-243).

4. Compte tenu de l’état des recherches à l’époque et aussi des exigences propres de la
théologie, l’Encyclique Humani generis considérait la doctrine de l’" évolutionnisme "
comme une hypothèse sérieuse, digne d’une investigation et d’une réflexion approfondies
à l’égal de l’hypothèse opposée. Pie XII ajoutait deux conditions d’ordre méthodologique
:qu’on n’adopte pas cette opinion comme s’il s’agissait d’une doctrine certaine et
démontrée, et comme si on pouvait faire totalement abstraction de la Révélation à propos
des questions qu’elle soulève.

Il énonçait également la condition à laquelle cette opinion était compatible avec la foi
chrétienne, point sur lequel je reviendrai.
Aujourd’hui, près d’un demi-siècle après la parution de l’Encyclique, de nouvelles
connaissances conduisent à reconnaître dans la théorie de l’évolution plus qu’une
hypothèse. Il est en effet remarquable que cette théorie se soit progressivement imposée
à l’esprit des chercheurs, à la suite d’une série de découvertes faites dans diverses
disciplines du savoir. La convergence, nullement recherchée ou provoquée, des résultats
de travaux menés indépendamment les uns des autres, constitue par elle-même un
argument significatif en faveur de cette théorie.

Quelle est la portée d’une semblable théorie ?

Aborder cette question, c’est entrer dans le champ de l’épistémologie. Une théorie est une
élaboration métascientifique, distincte des résultats de l’observation, mais qui leur est
homogène. Grâce à elle, un ensemble de données et de faits indépendants entre eux
peuvent être reliés et interprétés dans une explication unitive. La théorie prouve sa validité
dans la mesure où elle est susceptible d’être vérifiée ; elle est constamment mesurée à
l’étiage des faits ; là où elle cesse de pouvoir rendre compte de ceux-ci, elle manifeste ses
limites et son inadaptation. Elle doit alors être repensée.

En outre, l’élaboration d’une théorie comme celle de l’évolution, tout en obéissant à

l’exigence d’homogénéité avec les données de l’observation, emprunte certaines notions à
la philosophie de la nature. Et, à vrai dire, plus que de la théorie de l’évolution, il convient
de parler des théories de l’évolution. Cette pluralité tient, d’une part, à la diversité des
explications qui ont été proposées du mécanisme de l’évolution et, d’autre part, aux
diverses philosophies auxquelles on se réfère. Il existe ainsi des lectures matérialistes et
réductionnistes, et des lectures spiritualistes. Le jugement ici est de la compétence propre
de la philosophie et, au-delà, de la théologie.

5. Le Magistère de l’Église est directement intéressé par la question de l’évolution, car

celle-ci touche la conception de l’homme, dont la Révélation nous apprend qu’il a été créé
à l’image et à la ressemblance de Dieu (cf. Gn 1, 28-29). La Constitution conciliaire
Gaudium et spes a magnifiquement exposé cette doctrine, qui est un des axes de la
pensée chrétienne.

Elle a rappelé que l’homme est " la seule créature sur terre que Dieu a voulue pour elle-
même "(n. 24).

En d’autres termes, l’individu humain ne saurait être subordonné comme un pur moyen ou
un pur instrument ni à l’espèce ni à la société ; il a valeur pour lui-même. Il est une
personne. Par son intelligence et sa volonté, il est capable d’entrer en relation de
communion, de solidarité et de don de soi avec son semblable. Saint Thomas observe que
la ressemblance de l’homme avec Dieu réside spécialement dans son intelligence
spéculative, car sa relation avec l’objet de sa connaissance ressemble à la relation que
Dieu entretient avec son œuvre (Somme théologique, I-II, q. 3, a. 5, ad 1m).
Mais, plus encore, l’homme est appelé à entrer dans une relation de connaissance et
d’amour avec Dieu lui-même, relation qui trouvera son plein épanouissement au-delà du
temps, dans l’éternité.

Dans le mystère du Christ ressuscité, nous sont révélées toute la profondeur et toute la
grandeur de cette vocation (cf. Gaudium et spes, 22). C’est en vertu de son âme spirituelle
que la personne tout entière jusque dans son corps possède une telle dignité. Pie XII avait
souligné ce point essentiel : si le corps humain tient son origine de la matière vivante qui
lui préexiste, l’âme spirituelle est immédiatement créée par Dieu (" Animas enim a Deo
immediate creari catholica fides nos retinere jubet ")(Enc. Humani generis, AAS 42 [ 1950],
p. 575).

En conséquence, les théories de l’évolution qui, en fonction des philosophies qui les
inspirent, considèrent l’esprit comme émergeant des forces de la matière vivante ou
comme un simple épiphénomène de cette matière, sont incompatibles avec la vérité de
l’homme. Elles sont d’ailleurs incapables de fonder la dignité de la personne.

6. Avec l’homme, nous nous trouvons donc devant une différence d’ordre ontologique,
devant un saut ontologique, pourrait-on dire. Mais poser une telle discontinuité
ontologique, n’est-ce pas aller à l’encontre de cette continuité physique qui semble être
comme le fil conducteur des recherches sur l’évolution, et cela dès le plan de la physique
et de la chimie ?

La considération de la méthode utilisée dans les divers ordres du savoir permet de mettre
en accord deux points de vue qui sembleraient inconciliables. Les sciences de
l’observation décrivent et mesurent avec toujours plus de précisions les multiples
manifestations de la vie et les inscrivent sur la ligne du temps.

Le moment du passage au spirituel n’est pas objet d’une observation de ce type, qui peut
néanmoins déceler, au niveau expérimental, une série de signes très précieux de la
spécificité de l’être humain.

Mais l’expérience du savoir métaphysique, de la conscience de soi et de sa réflexivité,

celle de la conscience morale, celle de la liberté, ou encore l’expérience esthétique et
religieuse, sont du ressort de l’analyse et de la réflexion philosophiques, alors que la
théologie en dégage le sens ultime selon les desseins du Créateur.

7. En terminant, je voudrais évoquer une vérité évangélique susceptible d’apporter une

lumière supérieure à l’horizon de vos recherches sur les origines et le déploiement de la
matière vivante.
La Bible, en effet, est porteuse d’un extraordinaire message de vie. Elle nous donne sur la
vie, en tant qu’elle caractérise les formes les plus hautes de l’existence, une vision de
sagesse.

Cette vision m’a guidé dans l’Encyclique que j’ai consacrée au respect de la vie humaine
et que j’ai intitulée précisément Evangelium vitae. Il est significatif que, dans l’Évangile de
saint Jean, la vie désigne la lumière divine que le Christ nous communique. Nous sommes
appelés à entrer dans la vie éternelle, c’est-à-dire dans l’éternité de la béatitude divine.
Pour nous mettre en garde contre les tentations majeures qui nous guettent, notre
Seigneur cite la grande parole du Deutéronome : " Ce n’est pas de pain seul que vivra
l’homme, mais de toute parole qui sort de la bouche de Dieu " (8, 3 ; cf. Mt 4, 4).
Bien plus, la vie est un des plus beaux titres que la Bible ait reconnu à Dieu. Il est le Dieu
vivant.

De grand cœur, j’invoque sur vous tous et sur tous ceux qui vous sont proches,
l’abondance des Bénédictions divines.
Du Vatican, le 22 octobre 1996

JEAN-PAUL II
 EVOLUTION
A book of readings from Wikipedia
Vol. 4: Case studies

Khalid Chraibi

w000
 Basic topics in evolutionary biology
Evidence of common descent
Processes of evolution
Adaptation · Macroevolution · Microevolution · Speciation
Population genetic mechanisms
Genetic drift · Gene flow · Mutation · Natural selection
Evolutionary developmental
biology (Evo-devo) concepts
Canalisation · Inversion · Modularity · Phenotypic plasticity
Evolution of organs
and biological processes
Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair · Human intelligence · Modular ·
Muticellular · Sex
Taxa evolution
Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses · Humans · Influenza · Insects ·
Lemur · Life · Molluscs · Plants · Sirenians (sea cows) · Spiders
Modes of speciation
 Anagenesis · Catagenesis · Cladogenesis
History of evolutionary thought
Charles Darwin · On the Origin of Species · Modern evolutionary synthesis · Gene-centered view of
evolution · Life (classification trees)
Other subfields
Ecological genetics · Molecular evolution · Phylogenetics · Systematics
List of evolutionary biology topics · Timeline of evolution

w000

Evolution of birds
 From Wikipedia, the free encyclopedia

Jump to: navigation, search

Archaeopteryx at Paläontologisches Museum München

The evolution of birds is thought to have begun in the Jurassic Period, with the earliest birds derived
from theropod dinosaurs. Birds are categorized as a biological class, Aves. The earliest known species of
class Aves is Archaeopteryx lithographica, from the Late Jurassic period, though Archaeopteryx is not
commonly considered to have been a true bird. Modern phylogenies place birds in the dinosaur clade
Theropoda. According to the current consensus, Aves and a sister group, the order Crocodilia, together are
the sole living members of an unranked "reptile" clade, the Archosauria.
Phylogenetically, Aves is usually defined as all descendants of the most recent common ancestor of
a specific modern bird species (such as the House Sparrow, Passer domesticus), and either Archaeopteryx,
[1] or some prehistoric species closer to Neornithes (to avoid the problems caused by the unclear
relationships of Archaeopteryx to other theropods).[2] If the latter classification is used then the larger group
is termed Avialae. Currently, the relationship between dinosaurs, Archaeopteryx, and modern birds is still
under debate.

Contents
[hide]
• 1 Origins
• 2 Adaptive radiation of birds
• 3 Classification of modern species
• 4 Current evolutionary trends in birds
• 5 References

[edit] Origins
Main article: Origin of birds
 See also: Avicephala
There is significant evidence that birds evolved from theropod dinosaurs, specifically, that birds are
members of Maniraptora, a group of theropods which includes dromaeosaurs and oviraptorids, among
others.[3] As more non-avian theropods that are closely related to birds are discovered, the formerly clear
distinction between non-birds and birds becomes less so. Recent discoveries in northeast China (Liaoning
Province), demonstrating that many small theropod dinosaurs had feathers, contribute to this ambiguity.[4]
The basal bird Archaeopteryx, from the Jurassic, is well-known as one of the first "missing links" to
be found in support of evolution in the late 19th century, though it is not considered a direct ancestor of
modern birds. Confuciusornis is another early bird; it lived in the Early Cretaceous. Both may be predated by
Protoavis texensis, though the fragmentary nature of this fossil leaves it open to considerable doubt whether
this was a bird ancestor. Other Mesozoic birds include the Confuciusornis, the Enantiornithes, Yanornis,
Ichthyornis, Gansus, and the Hesperornithiformes - a group of flightless divers resembling grebes and loons.
The recently (2002) discovered dromaeosaur Cryptovolans (which may be a Microraptor) was capable of
powered flight, possessed a sternal keel and had ribs with uncinate processes. In fact, Cryptovolans makes a
better "bird" than Archaeopteryx which lacks some of these modern bird features. Because of this, some
paleontologists have suggested that dromaeosaurs are actually basal birds whose larger members are
secondarily flightless, i.e. that dromaeosaurs evolved from birds and not the other way around. Evidence for
this theory is currently inconclusive, but digs continue to unearth fossils (especially in China) of the strange
feathered dromaeosaurs. At any rate, it is fairly certain that flight utilizing feathered wings existed in the mid-
Jurassic theropods and was "tried out" in several lineages and variants by the mid-Cretaceous, such as in
Confuciusornis. This latter species had some peculiar features. For example, its vestigial tail was unfit for
steering, and its wing shape seems rather specialized although the arm skeleton was still quite
"dinosaurian").
 Illustration of Confuciusornis, a Cretaceous bird from China. Note wing claws and lack of tail.
Although ornithischian (bird-hipped) dinosaurs share the same hip structure as birds, birds actually
originated from the saurischian (lizard-hipped) dinosaurs if the dinosaurian origin theory is correct. They thus
arrived at their hip structure condition independently. In fact, a bird-like hip structure also developed a third
time among a peculiar group of theropods, the Therizinosauridae.
An alternate theory to the dinosaurian origin of birds, espoused by a few scientists, notably Larry
Martin and Alan Feduccia, states that birds (including maniraptoran "dinosaurs") evolved from early
archosaurs like Longisquama.[5] This theory is contested by most other paleontologists and experts in
feather development and evolution.[6]
 In a study published in 2010, P. Senter identified what he called a "Birdlike cluster" of
coelurosaurians and a "Tyrannosaur cluster". He included in the Birdlike cluster: Archaeopteryx,
Confuciusornis, Sapeornis, Dromaeosauridae, Troodontidae, Epidendrosaurus + Epidexipteryx,
Protarchaeopteryx + Incisivosaurus and Falcarius. He included in the Tyrannosaur cluster:
Tyrannosauroidea, Compsognathidae and Ornitholestes. Using classic multidimensional scaling (CMDS) he
shows that the two clusters are separate.

[edit] Adaptive radiation of birds

Modern birds are classified in Neornithes, which are now known to have evolved into some basic
lineages by the end of the Cretaceous (see Vegavis). The Neornithes are split into the paleognaths and
neognaths.
Paleognathae
The paleognaths include the tinamous (found only in Central and South America) and the ratites which
nowadays are found almost exclusively on the Southern Hemisphere. The ratites are large flightless birds,
and include ostriches, rheas, cassowaries, kiwis and emus. A few scientists propose that the ratites
represent an artificial grouping of birds which have independently lost the ability to fly in a number of
unrelated lineages[citation needed]; in any case, the available data regarding their evolution is still very
confusing.
Neognathae
The basal divergence from the remaining Neognathes was that of the Galloanserae, the superorder
containing the Anseriformes (ducks, geese and swans), and the Galliformes (chickens, turkeys, pheasants,
and their allies).
 The dates for the splits are a matter of considerable debate amongst scientists. It is agreed that the
Neornithes evolved in the Cretaceous and that the split between the Galloanserae and the other neognaths -
the Neoaves - occurred before the K-T extinction event, but there are different opinions about whether the
radiation of the remaining neognaths occurred before or after the extinction of the other dinosaurs.[7] This
disagreement is in part caused by a divergence in the evidence, with molecular dating suggesting a
Cretaceous radiation, a small and equivocal neoavian fossil record from Cretaceous, and most living families
turning up during the Paleogene. Attempts made to reconcile the molecular and fossil evidence have proved
controversial.[7][8]
On the other hand, two factors must be considered: First, molecular clocks cannot be considered
reliable in the absence of robust fossil calibration, whereas the fossil record is naturally incomplete. Second,
in reconstructed phylogenetic trees, the time and pattern of lineage separation corresponds to the evolution
of the characters (such as DNA sequences, morphological traits etc.) studied, not to the actual evolutionary
pattern of the lineages; these ideally should not differ by much, but may well do so in practice.
Considering this, it is easy to see that fossil data, compared to molecular data, tends to be more
accurate in general, but also to underestimate divergence times: morphological traits, being the product of
entire developmental genetics networks, usually only start to diverge some time after a lineage split would
become apparent in DNA sequence comparison - especially if the sequences used contain many silent
mutations.

[edit] Classification of modern species

See also: Sibley-Ahlquist taxonomy and dinosaur classification
 The phylogenetic classification of birds is a contentious issue. Sibley & Ahlquist's Phylogeny and
Classification of Birds (1990) is a landmark work on the classification of birds (although frequently debated
and constantly revised). A preponderance of evidence suggests that most modern bird orders constitute
good clades. However, scientists are not in agreement as to the precise relationships between the orders;
evidence from modern bird anatomy, fossils and DNA have all been brought to bear on the problem but no
strong consensus has emerged. As of the mid-2000s, new fossil and molecular data provide an increasingly
clear picture of the evolution of modern bird orders, and their relationships. For example, the Charadriiformes
seem to consititute an ancient and distinct lineage, while the Mirandornithes and Cypselomorphae are
supported by a wealth of anatomical and molecular evidence. Our understanding of the interrelationships of
lower level taxa also continues to increase, particularly in the massively diverse perching bird order
Passeriformes.
On June 27, 2008, the largest study of bird genetics was published. It overturns several hypothesized
relationships, and will likely necessitate a wholesale restructuring of the avian phylogenetic tree.[9]

[edit] Current evolutionary trends in birds

See also: Bird conservation
Evolution generally occurs at a scale far too slow to be witnessed by humans. However, bird species
are currently going extinct at a far greater rate than any possible speciation or other generation of new
species. The disappearance of a population, subspecies, or species represents the permanent loss of a
range of genes.
Another concern with evolutionary implications is a suspected increase in hybridization. This may
arise from human alteration of habitats enabling related allopatric species to overlap. Forest fragmentation
can create extensive open areas, connecting previously isolated patches of open habitat. Populations that
were isolated for sufficient time to diverge significantly, but not sufficient to be incapable of producing fertile
offspring may now be interbreeding so broadly that the integrity of the original species may be compromised.
For example, the many hybrid hummingbirds found in northwest South America may represent a threat to the
conservation of the distinct species involved.[10]
Several species of birds have been bred in captivity to create variations on wild species. In some
birds this is limited to color variations, while others are bred for larger egg or meat production, for
flightlessness or other characteristics.
Some species, like the rock pigeon or several species of crows have been successful living in man
made environments. Because these new habitats are different from their far less numerous "natural" habitats
these species are to a certain extent evolutionary adapting to living close to man.

[edit] References
1. ^ Padian K & Chiappe LM (1997). "Bird Origins". in Currie PJ & Padian K. Encyclopedia of
Dinosaurs. San Diego: Academic Press. pp. 41–96.
2. ^ Gauthier, J (1986). "Saurischian Monophyly and the origin of birds". in Padian K. The
Origin of Birds and the Evolution of Flight. Mem. California Acad. Sci 8 . pp. 1–55.
3. ^ Hou L,Martin M, Zhou Z & Feduccia A, (1996) "Early Adaptive Radiation of Birds: Evidence
from Fossils from Northeastern China" Science 274(5290): 1164-1167 Abstract
4. ^ Norell, M & Ellison M (2005) Unearthing the Dragon, The Great Feathered Dinosaur
Discovery Pi Press, New York, ISBN 0-13-186266-9
 5. ^ Feduccia A, Lingham-Soliar T, Hinchliffe JR (2005) "Do feathered dinosaurs exist? Testing
the hypothesis on neontological and paleontological evidence" Journal of Morphology 266(2): 125-
166
6. ^ Prum R (2003) "Are Current Critiques Of The Theropod Origin Of Birds Science? Rebuttal
To Feduccia 2002" Auk 120(2) 550-561
7. ^ a b Ericson PGP, Anderson CL, Britton T, Elzanowski A, Johansson US, Kallersjo M,
Ohlson JI, Parsons TJ, Zuccon D, Mayr G (2006)"Diversification of Neoaves: integration of molecular
sequence data and fossils" Biology Letters 2(4): 543-547
8. ^ Brown JW, Payne RB, & Mindell DP (2007) "Nuclear DNA does not reconcile ‘rocks’ and
‘clocks’ in Neoaves: a comment on Ericson et al. Biology Letters 3(3): 257-259
9. ^ Science Daily, June 27, 2008
10.^ Fjeldså, Jon; Niels Krabbe. (1990). Birds of the High Andes: A Manual to the Birds of the
Temperate Zone of the Andes and Patagonia, South America . Apollo Books. ISBN 8788757161.
 [hide]
v•d•e
Birds (class: Aves)

Anatomy Bird anatomy • Flight • Eggs • Feathers • Plumage • Beak • Vision • Dactyly •
Preen gland

Behaviour Singing • Intelligence • Migration • Incubation • Brood parasites • Nesting •

Hybrids

Evolution Evolution of birds • Origin of birds • Darwin's finches • Seabirds

Superorder: Palaeognathae
Subclass:
Neornithes Struthioniformes (ratites) • Tinamiformes (tinamous)
(29 orders of
modern birds)
Superorder: Neognathae

Accipitriformes • Anseriformes (waterfowl) • Apodiformes (swifts and

hummingbirds) • Caprimulgiformes (nightjars and relatives) • Cariamae (seriemas and
relatives) • Charadriiformes (gulls and relatives) • Ciconiiformes (storks) • Coliiformes
(mousebirds) • Columbiformes (doves and pigeons) • Coraciiformes (kingfishers and
 relatives) • Cuculiformes (cuckoos and relatives) • Falconiformes (falcons and
relatives) • Galliformes (gamebirds) • Gaviiformes (loons or divers) • Gruiformes
(cranes and relatives) • Passeriformes (perching birds) • Pelecaniformes (pelicans
and relatives) • Phaethontiformes (tropicbirds) • Phoenicopteriformes (flamingos) •
Piciformes (woodpeckers and relatives) • Podicipediformes (grebes) •
Procellariiformes (albatrosses and petrels) • Psittaciformes (parrots) • Pteroclidiformes
(sandgrouses) • Sphenisciformes (penguins) • Strigiformes (owls) • Trogoniformes
(trogons and quetzals)

Fossil birds Archaeopteryx • Enantiornithes • Hesperornithes

Birds and Ringing • Ornithology • Bird collections • Birdwatching • Bird feeding •

humans Conservation • Aviculture • Waterfowl hunting • Pigeon racing

Lists Families and orders • Genera • Lists by region • Extinct birds • Late Quaternary
prehistoric birds • Sibley-Ahlquist taxonomy
 W000

Histoire évolutive des oiseaux

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La forme ou le fond de cet article est à vérifier.
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Les oiseaux (Aves) sont des animaux bipèdes, ovipares, à sang chaud, en principe dotés de plumes.
Très simplement reconnaissables, les oiseaux ont longtemps été placés dans le règne animal comme
intermédiaires entre les reptiles et les mammifères. Cependant la paléontologie et les études cladistiques
montrent que les oiseaux sont des dinosaures théropodes, et la génétique confirme que les Crocodilia
forment le groupe actuel vivant le plus proche de celui des oiseaux. Les oiseaux ne dérivent donc pas des
reptiles volants (les Ptérosaures). Un consensus définit ce groupe comme monophylétique.

Le groupe des "reptiles" est en bleu.

Sommaire
[masquer]
• 1 Origines
• 1.1 Généralités
• 1.2 Les théropodes
• 1.3 Apparition des oiseaux
• 1.4 Origine de la plume
• 2 Période d'affirmation
• 2.1 La radiation adaptative
• 3 Menace actuelle
• 4 Genèse de la théorie de l'origine des oiseaux
• 4.1 Classification classique
• 4.2 Classification de Sibley
• 4.3 Cladogramme
• 4.4 Nomenclature des ordres d'Oiseaux modernes
• 5 Notes et références
• 6 Voir aussi
• 6.1 Articles connexes
• 6.2 Liens externes
Origines [modifier]
Généralités [modifier]
Il a existé plusieurs théories quant à l'apparition des oiseaux. La plus communément admise fait des
oiseaux les descendants de petits dinosaures théropodes. Une autre les présente comme descendants de
reptiles du Permien, les Thecodontia (taxon considéré aujourd'hui comme désuet car paraphylétique).
L'antithèse la plus sérieuse est une différentiation très tôt des oiseaux les excluant des dinosaures (soutenue
par Feduccia par exemple[1]).
Les études de Jacques Gauthier et d'autres paléontologues démontrent que de nombreux caractères
qui étaient classiquement considérés comme spécifiques aux oiseaux sont apparus avant ceux-ci, chez les
théropodes, voire plus tôt encore.
L'hypothèse des théropodes sous-tend que certains des caractères adaptatifs propres aux dinosaures
terrestres, furent conservés chez les oiseaux. D'autres ont été modifiés comme les écailles appelées
scutelles sur les pattes des gallinacés ou les plumes. Les modifications permirent le vol et un mode de vie
arboricole. Naturellement, l'apparition des caractéristiques des théropodes qui précédèrent les oiseaux ne fut
pas simultanée. De la même façon que l'œuf ne fait pas l'oiseau, la plume est présente chez de très
nombreux théropodes. Bon nombre de caractéristiques propres aux dinosaures ne se retrouvent plus
aujourd'hui que chez les oiseaux.
Position des doigts et longueur des poignets aux doigts
Rhamphorhynchus, un ptérosaure Archéoptéryx
 Le second doigt est le plus long et porte l'aile
Le quatrième doigt porte l'aile, le poignet est long.
pas de tarsométatarsien.
L'anatomie comparée montre notamment que les ptérosaures ne peuvent pas être les ancêtres
d'Archéoptéryx pour des raisons morphologiques.
└─o Archosauria
├─o Crocodylotarsi (ie Crocodylia, les crocodiles)
└─o Ornithodira
├─o
│ └─o Dinosauria (les dinosaures, dont les théropodes, puis les
oiseaux)
└─o
└─o Pterosauria (éteint) (les ptérosaures)

Les théropodes [modifier]

L'ancêtre reptilien direct des dinosaures théropodes est un petit carnivore bipède qui marche comme
un oiseau.
• Ses pattes avant sont libres pour saisir des objets ou des proies, et le plus long doigt est le
deuxième, non le troisième comme chez les autres Sauropsides. Il a toutefois cinq doigts par main,
alors que presque tous les théropodes primitifs et les oiseaux en ont trois. Chez cet animal,
l'articulation de la cheville est déjà en charnière, et les métatarses sont allongés, relevés : l'animal
marche, comme les dinosaures après lui. Les nombreuses modifications ultérieures du pied
permettent probablement une foulée plus longue et plus rapide, qualité qui aidera ultérieurement les
théropodes à développer le vol.
• Les premiers théropodes présentent des adaptations qui allègent leur squelette: des os
creux et des cavités dans le crâne.

Doigts de Dromaeosaurus.
 • Comme les oiseaux actuels, leur cou est long et leur dos horizontal.
Le quatrième doigt de la main des premiers dinosaures est déjà réduit, et le cinquième doigt a déjà
presque disparu. Le quatrième doigt devient bientôt lui-aussi un moignon, puis ces deux doigts
réduits disparaissent ensemble chez les théropodes tétanoures. Les trois doigts restants fusionnent
au cours de l'évolution des oiseaux postérieurs à Archaeopteryx.
• Les membres postérieurs des premiers théropodes se transforment aussi. Ils s'allongent : le
fémur devient plus long que le tibia, et la fibula se réduit.

Pied de Tyrannosaurus rex.

 • Ces dinosaures marchent sur les trois doigts du milieu, ceux qu'utilisent toujours les oiseaux
actuels. Le cinquième orteil, sans articulation, est raccourci et effilé. Le premier orteil, qui dépasse
sur le côté du deuxième orteil, est formé d'un os raccourci, d'une articulation et d'une griffe. Il est
placé au-dessus des autres orteils, tel un ergot.
• Les plumes (ou un protoplumage) sont déjà présentes chez de nombreux théropodes et
devaient avoir d'autres utilités. Ainsi, déjà présentes chez les ancêtres des oiseaux, elles ont ensuite
permis à ceux-ci de voler grâce à toute une autre série de caractéristiques.

o Tetanurae
└─o Coelurosauria
└─o
├─o Ornithomimosauria (éteint)
└─o Maniraptora
├─o Ornitholestes, Compsognathidae, Alvarezsauridae (éteint)
└─o
├─o Oviraptoriformes (éteint)
│ ├─o Therizinosauria (éteint)
│ └─o Oviraptorosauria (éteint)
 └─o Paraves

Apparition des oiseaux [modifier]

• Au cours de l'évolution des théropodes, d'autres caractères qui semblaient être l'apanage
des oiseaux apparaissent. Par exemple, le membre antérieur et la ceinture pectorale se modifient,
favorisant d'abord la capture de proies, puis l'envol.
• Les bras s'allongent, sauf chez les carnivores géants, tels Carnotaurus, Allosaurus et
Tyrannosaurus : chez les théropodes primitifs, le membre antérieur est deux fois plus court que le
membre postérieur, mais, pour Archaeopteryx, le membre antérieur est plus long que le membre
postérieur, et cette différence s'accentue encore chez les oiseaux plus récents, auxquels cet
allongement confère un battement d'ailes plus puissant.

Serre de Centropus.
 • Lorsque Archéoptéryx apparait, le premier orteil orienté situé plus haut a migré pour se
placer derrière les autres.
Chez certains oiseaux plus récents, il sera plus bas sur la patte, opposable aux autres orteils, et il
permettra aux pieds d'oiseaux arboricoles d'agripper fortement les branches ou de se transformer en
serre.

Aile de Confuciusornis. En rouge, l'ouverture de l'humérus et la soudure du Carpométacarpe,

Chiappe et al. 1999).
• Les doigts des ailes aussi s'allongent, occupant progressivement une plus grande proportion
du membre antérieur, et le poignet est remodelé. Les théropodes primitifs ont un os du poignet
aplati, qui chevauche la base des premier et second os de la paume et les doigts. Chez les
maniraptoriens, cet os a une forme de demi-lune le long de la surface de contact avec les os du bras.
Cette forme en demi-lune est importante : elle permet à ces animaux de plier le poignet latéralement,
et plus seulement verticalement. Ainsi, ils peuvent presque replier leur longue main comme le font
les oiseaux actuels avec leurs ailes. Cette main allongée peut aussi être tournée et brusquement
projetée vers l'avant pour la capture d'une proie. Seul les Hoazins ont gardés des griffes sur leurs
ailes.
• Le sternum des tétanoures s'ossifie en deux plaques qui fusionnent, à l'origine il est
cartilagineux et il devient le bréchet. L'ensemble de ces modifications consolident le squelette. Plus
tard, ce renforcement soutient les muscles du vol ; la furcula devient un point d'ancrage pour les
muscles du membre antérieur, permettant d'abord de saisir, puis de voler.
• Dans la région du bassin, au cours de l'évolution, des vertèbres s'ajoutent à la ceinture
pelvienne, et l'orientation du pubis change, le pubis pointe vers l'avant et vers le bas ; puis il recule
et, chez les oiseaux plus récents qu'Archéoptéryx, devient parallèle à l'ischion. Nous ignorons ce qui
a favorisé cette évolution, mais le partage de ces caractéristiques par les oiseaux et par d'autres
Maniraptora témoignent de leur origine commune.
 De gauche à droite, les bassins de :
-Coélophysis
-Allosaurus
-Vélociraptor
-Archéoptéryx
-Columba
• Leur queue raccourcit et se rigidifie, devenant un balancier plus efficace pendant la course.
Steven Gatesy, de l'Université Brown, a montré que cette transformation est associée à un
changement de fonction : l'ancrage des muscles des membres postérieurs passe de la queue au
bassin. Chez les maniraptoriens, les muscles qui tiraient les pattes vers l'arrière peuvent alors
contrôler la queue. Chez les oiseaux qui évoluent après Archéoptéryx, ces muscles semblent être
utilisés pour diriger la queue, couverte de plumes, pendant le vol.
 Le vol engendre de nombreuses contraintes physiologiques, ayant nécessité des adaptations par
rapport à un ancêtre reptilien:
• L'appareil respiratoire des oiseaux présente une disposition unique de plusieurs sacs aériens
séparés des voies respiratoires, l'air circule de façon continue dans leurs poumons. Il s'agit d'un
système très efficace qui permet notamment de répondre aux besoins énergétiques liés au vol.
• Les oiseaux sont endothermes et leur niveau d'énergie ne varie pas avec la température
ambiante, l'isolation thermique étant fournie par les plumes. Il est néanmoins probable que
l'endothermie ait précédé la capacité de voler chez les théropodes.
Tous ces éléments physiologiques sont étudiés difficilement sur les fossiles (organes mous internes
non fossilisés, fragilité du squelette des oiseaux, faiblesse du registre fossile).
└─o Paraves
├─o Deinonychosauria (éteint)
│ ├─o Troodontidae (éteint)
│ └─o Dromaeosauridae (éteint)
└─o Aves
├─o Archaeopterygidae (éteint)
└─o Ornithothoraces

[Dérouler]
Comparaison de caractéristiques ancestrales et dérivées dans l'évolution des oiseaux
Caractéristiques Théropodes Archaeopteryx Confuciusornis oiseaux p
anatomiques (non avaires) développés
 Ornithothoraces à p
220 à 65 Ma 156 à 150 Ma 125 et 110 Ma
du Jurassique mo

Fusion des deux

fenêtres temporales avec - + - +
l'orbite

Beaucoup
Disparition des -
dents nombreuses
- + les
Oiseaux du Créta

La queue à
Pygostyle raccourcie,
vertèbres de queue
en petit nombre - + +
fusionnées

Clavicules
-
+ + +
Accollées Forme en V,
fusionnées (furcula)| Forme de boomerang Forme de boomerang
(forme de boomerang) | à angle aigu

longueur du
Coracoïdes - - + +
long, en forme de ruban
 +
- Seulement les
Sternum
- - Sternum plat avec Neognathae , dispa
avec bréchet
quille très réduite secondaire chez
paléognathes

Côtes liées au
sternum
avec une - + +
Disparition des
Côtes abdominales - - - +
(Gastralia)

avec une côte

côtes avec supplémentaire ou
- + +
Processus unciné présent chez les Plusieurs
Maniraptora

Humérus dilaté
avec crête deltopectoral - - + -
perforé (autoapomorphie)

os de métacarpe - - + (fusion +
 des patte avant I, III
à Carpe-métacarpe après la base)
fusionné

os médian du
pied II-IV fusionné un + + +
(Tarso-métatarse)

os médian du
+ + + -
pied IV present

Plumes de vol
quelques uns + + +
actif, asymétriques

peu
Alula - - +
abondantes

Origine de la plume [modifier]

Article détaillé : Plume#Place de la plume dans l'évolution.
La plume ne dérive pas de l'écaille reptilienne, ce ne sont donc pas des structures homologues.
 Les nombreuses découvertes de dinosaures à plumes ont permis de proposer un scénario de
l'évolution de la plume, d'une protoplume filamenteuse à la plume à double vexille asymétrique moderne. Le
premier stade correspond à des protoplumes, décrites par exemple chez des Coelurosaures comme le
dinosaure Sinosauropteryx. Le deuxième stade correspond à une plume ramifiée, à rachis court et à barbes
libres, décrite par exemple chez Shuvuuia ou des Thérizinosaures. Le stade 3 correspond à des plumes dont
les barbes libres sont reliées au rachis. Au stade 4, les plumes possèdent des barbes avec barbules reliées
par des crochets, ce qui assure l'étanchéité du plumage (comme Caudipteryx ou Sinornithosaurus). Dans le
cinquième et dernier stade, la plume devient asymétrique, adaptation au vol ( Microraptor, Archaeopteryx).
De nombreux dinosaures à plumes sont des théropodes coureurs sans doute incapables de
voler[réf. nécessaire], ce qui tend à démontrer que l'apparition de la plume est indépendante du vol. Elle
pourrait être liée à un rôle d'isolation thermique, de communication notamment sexuelle dans la
reproduction. L'importance de la sélection sexuelle chez les oiseaux (parades nuptiales, ornementations,
etc.) a été à l'origine d'une théorie selon laquelle des productions cutanées se seraient chargées de
pigments colorés (comme les plumes actuelles) et auraient pris de l'importance suite à une sélection
sexuelle pour exhiber leurs couleurs, devenant plus larges et plus longues, d'où les plumes qui aurait conduit
aux ailes et en recouvrant les griffes auraient permis leur disparition[réf. nécessaire]. La régression de la
main étant liée à la course bipède, la réduction des doigts aurait été compensée par l'allongement des
plumes devenant des rémiges.

Période d'affirmation [modifier]

Articles détaillés : Extinction KT et Extinction JK.
 Un arbre évolutif des oiseaux, depuis les théropodes.
Certains auteurs défendent la thèse que les oiseaux pré-tertiaires appartiendraient à des groupes
modernes de charadriiformes ou de gaviiformes. D’autres assurent que peu de lignées d’oiseaux ont survécu
à la fin du Crétacé.
Les fossiles d'oiseaux de mer, sont plus nombreux car ceux-ci vivaient dans un milieu ou des zones
de sédimentation sont présentes[2].

Hesperornis regalis au National Museum of Natural History.

Les plus anciens identifiés sont les Hesperornithiformes, comme Hesperornis regalis et vivaient au
Crétacé. Ils avaient de petites dents pointues et ne pouvaient certainement pas voler. Les oiseaux marins
d'aujourd'hui descendent plutôt d'un groupe apparenté aux Tytthostonyx glauconiticus.
──o Pygostylia
├─o Confuciusornithidae
 └─o Ornithothoraces
├─o Enantiornithomorpha (éteint)
└─o Ornithurae
├─o Yanornis (éteint)
└─o
├─o Patagopteryx (éteint)
└─o
├─o Hesperornithiformes (éteint)
│ ├─o Enaliornithidae (éteint)
│ ├─o Baptornithidae (éteint)
│ └─o
│ └─o Hesperornithidae (éteint)
└─o Carinatae
├─o Ichthyornithiformes (éteint)
└─o Neornithes

Durant le paléogène, les oiseaux de mer les plus fréquents sont classés dans le groupe des
Procellariiformes parmi lequel on classe plusieurs espèces éteintes d'oiseaux de grande taille ressemblants
à des pingouins[3].
C'est à la fin du Miocène et au cours du Pliocène que les genres modernes deviennent
prépondérants, bien que le genre Puffinus remonte lui à l'Oligocène[2]. Leur diversité disparait
soudainement, et, un peu plus tard, les mammifères marins semblent leur ravir la place. Ceci est peut être dû
à un évènement astronomique qui a pu diminuer la biomasse des océans il y a 2 Ma[4], la nouvelle présence
des Pinnipedia empêchant dès lors les oiseaux de retrouver leur diversité d'autrefois[5].

La radiation adaptative [modifier]

Il existe aujourd'hui près de 10 000 espèces d'oiseaux.
 Les Pinsons de Darwin des Galápagos illustrent une radiation évolutive :
d'une espèce originale, il est apparu 4 types de bec pour 13 espèces au total.
└─o Carinatae
└─o Neornithes
├─o Paleognathae (autruche, émeu...)
└─o Neognathae (les autres oiseaux.)

Les Paléognathes qui incluent les Tinamidae que l'on ne retrouve qu'en Amérique du Sud et
Centrale et les Struthioniformes. Ils vivent aujourd'hui exclusivement sur l'hémisphère Sud. Ces oiseaux ne
peuvent voler. Certains scientifiques pensent qu'ils représentent un groupement artificiel d'oiseaux qui ont
perdu leur capacité de voler indépendamment ; en tout cas, les données disponibles quant à leur évolution
ne sont pas très claires.

La ceinture scapulaire de Neognathae

H : Humérus ;
S : Scapula ;
C : Coracoïde ;
F : Furcula ;
T : Canal

Les dates de différentiation entre les diverses espèces de Neognathae sont très disputées.
On observe cependant que la scapula s'allonge ensuite et s'affine, comme le coracoïde, qui s'étire ainsi vers
le sternum. Les clavicules fusionnent sur leur ligne médiane et s'élargissent jusqu'à former une furcula en
forme de boomerang.

Les analyses génétiques, pour être précises, doivent être corroborées puis étalonnées par des
fossiles et doivent porter sur des gènes significatifs qui témoignent de l'évolution.
De nombreuses discussions entre scientifiques portent sur cette question.
Voir aussi: Liste d'oiseaux préhistoriques
Menace actuelle [modifier]
Articles détaillés : Extinction de l'Holocène et Protection des oiseaux.
L'évolution se produit généralement à une échelle trop lente pour être perçue par les humains.
Cependant on note une forte diminution des espèces, ainsi actuellement 12 % des espèces sont reconnues
en danger par l'UICN. Outre leur disparition par la prédation, l'empoisonnement par les pesticides, la
destruction de leur environnement, on observe pour certaines espèces qui cherchent à s'adapter, une
augmentation insoupçonnée de l'hybridation. En effet, certaines espèces proches, voyant leur habitat se
morceler, se croisent en produisant même des hybrides stériles en quantité si importante que cela peut
mettre ces espèces en péril. Par exemple, les nombreux oiseaux-mouches hybrides trouvés dans le nord-
ouest de l'Amérique du Sud peuvent représenter une menace pour la conservation de ces espèces en
espèces distinctes.
Par conséquent on observe donc une perte manifeste du capital génétique.

Genèse de la théorie de l'origine des oiseaux [modifier]

Article détaillé : Chronologie de l'ornithologie.
Le premier fossile de ptérosaure a été décrit en 1784 et les paléontologues ne savaient s'il s'agissait
d'une créature aquatique, d'une chauve-souris ou d'un oiseau. Georges Cuvier, en se basant sur les
principes de l'anatomie comparée, annonce en 1801 qu'il s'agit d'un reptile volant et qu'ils ne sont pas les
ancêtres des oiseaux.
Charles Darwin publie en 1859 L'Origine des espèces. La première découverte, qui allait être suivie
de neuf autres, l'est sous la forme d’une plume fossilisée d' Archaeopteryx en 1861. Tous ces fossiles
proviennent de la Bavière, conservés dans le calcaire de Solnhofen. La particularité de ce matériau est qu’il
s’est formé dans un ancien lagon, sa composition ayant permis de protéger les moindres détails des êtres
vivants (dont les plumes) qui s’y sont égarés lors de l’ère Mésozoïque. Des squelettes d’Archaeopteryx sont
exposés au Muséum d'histoire naturelle de Londres et au Humboldt Museum de Berlin.
En 1862, est publié une étude d’Johann Andreas Wagner, un opposant à la théorie de Darwin,
directeur du Musée de Paléontologie de Munich qui identifie un reptile, le Griphosaurus. En 1863, Richard
Owen du British Museum y voit un oiseau. Le naturaliste britannique Thomas Huxley, est le premier à
affirmer que c'est bien une forme transitoire. Il se demande également si les dinosaures n'auraient pas été
des animaux à sang chaud (comme les oiseaux) plutôt qu'à sang froid (comme les reptiles) dans son
ouvrage On the Animals which are Most Nearly Intermediate between Birds and Reptiles publié auprès de la
Société géologique de Londres. En comparant l'anatomie de l'archéoptéryx et celle d'un autre fossile
bavarois, le petit Compsognathus, il recense quatorze caractéristiques communes et voit entre eux un lien de
parenté. Son hypothèse tomba dans l'oubli.
En 1870, le premier Hesperornithes est découvert par Othniel Charles Marsh et correctement décrit
par lui-même en 1877. En 1887, Harry Govier Seeley publie sa théorie selon la laquelle les dinosaures se
sont séparés en deux groupes qui se différencient selon leur ischion. Ironiquement, les oiseaux, qui sont,
selon la théorie la plus admise, descendant des Saurischia (terme qui dérive de lézard à ischion) et non pas
des Ornithischia (terme qui dérive du grec oiseau à ischion).
En 1924, la publication d'un dessin anatomique d'Oviraptor montrant une fourchette passe
inaperçue, puis en 1936, à l'Université de Berkeley, Charles Camp découvre le squelette complet d'un petit
théropode du Jurassique inférieur qui possède des clavicules. Des travaux plus récents démontrent, plus
généralement, la présence de clavicules chez de nombreux théropodes apparentés aux oiseaux.
 À la fin des années 1960, un siècle après la communication controversée de Huxley , John Ostrom,
de l'Université Yale, décrit le squelette du théropode du groupe des maniraptoriens Deinonychus, prédateur
aux griffes en faucille et grand comme un adolescent humain, qui rôdait dans le Montana il y a 115 Ma.
Ostrom identifie un ensemble de caractères communs aux oiseaux, dont Archaeopteryx, Oviraptor et
Deinonychus, et à d'autres théropodes, mais que ne possèdent pas d'autres Sauropsida. Il en déduit que les
oiseaux descendent directement des théropodes.
En 1975, l'américain Robert Bakker jeta finalement le pavé dans la mare en affirmant que les
dinosaures étaient les ancêtres directs des oiseaux contemporains. À l'Université de Berkeley, Jacques
Gauthier entreprend une vaste étude cladistique des oiseaux, des dinosaures et de leurs parents reptiliens,
en se fondant sur les comparaisons de J. Ostrom et sur bien d'autres caractères. Il confirme que les oiseaux
descendent de petits dinosaures théropodes, et le maniraptorien aux griffes en faucille décrit par J. Ostrom,
Deinonychus, est bien l'un des plus proches parents des oiseaux.
Vers la fin des années 1970 et durant toute la décennie suivante, Charles Gald Sibley et Jon Edward
Ahlquist mènent des études basées sur des méthodes d'hybridation de l'ADN, ce qui modifie profondément
les connaissances sur la phylogénie des oiseaux. Cependant les datations obtenues, en tenant compte du
taux de mutations naturelles, laissent supposer que la diversification des oiseaux est antérieure au Crétacé.
En 1981, une quarantaine de spécimens de Gansus, les plus anciens membres des Neornithes sont
découverts en Chine.
En 1993, la découverte de Vegavis, daté de 66 à 68 Ma, sont la première preuve de la coexistence
entre thérapodes non-aviaires et oiseaux modernes[6].
Depuis les années 1990, des dizaines d'espèces de dinosaures à plumes, aviens ou non, capables
de voler ou non, ont été découverts principalement en Chine, dans la région de Jehol: Sinosauropteryx,
Protarchaeopteryx, Caudipteryx, Microraptor, Confuciusornis, Protopteryx, Jeholornis... De toutes tailles et
appartenant à des groupes très différents, ils témoignent de manière difficilement contestable d'une
continuité entre dinosaures et oiseaux. Certaines espèces sont de véritables mosaïques de caractères plus
ou moins proches des oiseaux modernes. Toutes ces espèces permettent d'affiner la compréhension de
l'évolution des caractères des dinosaures aux oiseaux et de préciser leurs relations de parenté. Certains sont
ainsi plus proches des oiseaux modernes que d'Archaeopteryx (notamment Confuciusornis, Protopteryx,
Jeholornis).
L'origine dinosaurienne des oiseaux ne fait donc plus de doute à l'heure actuelle.

Classification classique [modifier]

La vision traditionnelle de l'évolution des Neornithes (oiseaux modernes) est de placer à la base de
l'arbre phylogénétique, après les ratites et tinamous, les groupes d'oiseaux marins tels les manchots, les
grèbes, les plongeons et pélicans, etc.

Classification de Sibley [modifier]

Articles détaillés : Dinosauria (classification phylogénétique), Aves (classification phylogénétique) et
Taxinomie Sibley-Ahlquist.
La nouvelle classification a montré que les canards et les gallinacés étaient les taxons les plus
proches de la séparation d'avec les ratites (qui intègrent les tinamous) formant les paléognathes. Les
Galloanserae constituant la lignée de base des Néognathes. Les groupes d'oiseaux marins
traditionnellement considérés comme archaïques ont été placés dans un ordre des Ciconiiformes élargi, qui
comprend aussi les rapaces diurnes (Accipitridae, Sagittariidae (Aves) et Falconidae) et les limicoles
(Scolopacidae, Charadriidae, Laridae, etc.).

Cladogramme [modifier]
Le cladogramme simplifié suivant suit la proposition de Norell et al. 2006, with clade names after the
definitions by Sereno, 2005[7], [8].
└─o Sauria
├─o Lepidosauria (serpents, lézards...)
└─o Archosauria
├─o Suchia (crocodiliens)
└─o Dinosauria
├─o Ornithischia (éteint)
└─o Saurischia
└─o Theropoda
└─o Coelurosauria
└─o Maniraptora
├─o Oviraptoriformes (éteint)
│ ├─o Therizinosauria (éteint)
│ └─o Oviraptorosauria (éteint)
└─o Paraves
├─o Deinonychosauria (éteint)
│ ├─o Troodontidae (éteint)
│ └─o Dromaeosauridae (éteint) (dont les microraptors)
 └─o Aves
├─o Archaeopterygidae (éteint) (les archéoptéryx)
└─o Ornithothoraces
├─o Enantiornithomorpha (éteint)
└─o Ornithuromorpha
├─o Hesperornithiformes (éteint)
└─o Carinatae
├─o Ichthyornithiformes
└─o Neornithes
├─o Paleognathae (autruches, émeux...)
└─o Neognathae (les autres oiseaux)

Nomenclature des ordres d'Oiseaux modernes [modifier]

[Dérouler]
Correspondances entre classifications récentes
Classification
Exemple de
Gill&Wright 06 Clements 07 Howard&Moore 03
nom vernaculaire
Struthioniformes Struthioniformes Struthioniformes Autruche

Rheiformes Rheiformes nandou

Casuariiformes Casuariiformes Casoar

Apterygiformes Apterygiformes kiwi

Tinamiformes Tinamiformes Tinamiformes

Galliformes Galliformes Galliformes

Anseriformes Anseriformes Anseriformes

Sphenisciformes Sphenisciformes Sphenisciformes manchot

Gaviiformes Gaviiformes Gaviiformes

Procellariiformes Procellariiformes Procellariiformes

Podicipediformes Podicipediformes Podicipediformes

Phoenicopteriformes Phoenicopteriformes Phoenicopteriformes flamant

Ciconiiformes Ciconiiformes Ciconiiformes

Pelecaniformes Pelecaniformes Pelecaniformes

Cathartiformes

Falconiformes Falconiformes Falconiformes

Gruiformes Gruiformes Gruiformes

 Turniciformes

Charadriiformes Charadriiformes Charadriiformes

Pteroclidiformes Pteroclidiformes

Columbiformes Columbiformes Columbiformes Pigeon

Psittaciformes Psittaciformes Psittaciformes Perroquet

Opisthocomiformes

Musophagiformes Musophagiformes

Cuculiformes Cuculiformes Cuculiformes

Strigiformes Strigiformes Strigiformes

Caprimulgiformes Caprimulgiformes Caprimulgiformes

Apodiformes Apodiformes Apodiformes Hirondelle

Trochiliformes

Coliiformes Coliiformes Coliiformes

 Trogoniformes Trogoniformes Trogoniformes

Coraciiformes Coraciiformes Coraciiformes

Upupiformes

Bucerotiformes

Galbuliformes

Piciformes Piciformes Piciformes

Passeriformes Passeriformes Passeriformes

Notes et références [modifier]

1. ↑ Hou L. et al. (1996), Early adaptative radiation of birds : Evidence from fossils from
Northrastern China, Science, Vol 274, 1164-1167
2. ↑ a et b Schreiber, Elizabeth A. & Burger, Joanne., Biology of Marine Birds
3. ↑ (en) Goedert, J., « Giant Late Eocene Marine Birds (Pelecaniformes: Pelagornithidae)
from Northwestern Oregon" », dans Journal of Paleontology, vol. 63, no 6, 1989, p. 939–944
[ résumé [archive] ]
 4. ↑ (en)Comins, Niel F.; William J. Kaufmann III, , Discovering the Universe, 7th edition,
New York, NY: Susan Finnemore Brennan, 2005, 359 p. (ISBN 0716775840.)
5. ↑ (en) Olson, S. & Hasegawa, Y., « Fossil Counterparts of Giant Penguins from the North
Pacific », dans Science magazine, vol. 206, no 4419, 1979, p. 688–689 .
6. ↑ (fr)Jean-Luc Goudet, « Après la chute de l'astéroïde, le canard était toujours
vivant [archive] », Futura-Science
7. ↑ (en) Norell, M.A., Clark, J.M., Turner, A.H., Makovicky, P.J., Barsbold, R., et Rowe, T.,
« A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). », dans American
Museum Novitates, vol. 3545, 2006, p. 1-51.
8. ↑ (en)Sereno, P. C., McAllister, S., and Brusatte, S. L., « TaxonSearch: a relational
database for suprageneric taxa and phylogenetic definitions. [archive] », PhyloInformatics

Voir aussi [modifier]

Articles connexes [modifier]
• Anatomie des oiseaux
• Aves (classification phylogénétique)
• La Guerre des os
• Extinction Crétacé-Tertiaire
Liens externes [modifier]
• Gigadino, site sur la paléontologie et les fossiles
• Evolution, dinosaures et Oiseaux
• Archaeopteryx, volatile controversé, Association FABULA
• Dinosaures et oiseaux sur astronomie et d'astronautique.com
• (en)Aves: Overview sur Palæos
• (en)David Swanson, « Zoology 461/561, Ornithology, Summer 1999 (July 6-29) », sur
l'origine et la morphologie des oiseaux.
• (en)Gary Ritchison, « Introduction to Birds », Department of Biological Sciences Eastern
Kentucky University
• (en)Fossil records of avian taxa sur ornitaxa.com, Bird Taxonomy for Birders
• (fr)Une analyse moléculaire confirme le lien entre le Tyrannosaure et les oiseaux sur
ornithomedia.com
• (fr)Une nouvelle étude bouleverse l'arbre phylogénétique des oiseaux sur
ornithomedia.com, 27/6/2008

[Enrouler]
v·d·m
Évolution biologique
[Enrouler]Mécanismes
 Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et
Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Enrouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Mammifères Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire
 évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Enrouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de la paléontologie

• Portail de l’origine et de l’évolution du vivant

• Portail de l’ornithologie

• Portail des dinosaures

• Portail de la zoologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_oiseaux ».

Catégories : Dinosaure | Paléontologie | Oiseau | Histoire évolutive | [+]

 W000

Butterfly evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Butterfly evolution refers to the origin and diversification of butterflies through geologic time and over
a goodly portion of the Earth's surface. Butterflies probably evolved about 150 million years ago. Of the
220,000 species of Lepidoptera, about 45,000 species are butterflies, which probably evolved from moths.
Butterflies are found throughout the world, except in Antarctica, and are especially numerous in the tropics;
they fall into eight different families.
Contents
[hide]
• 1 Phylogeny
• 2 Lines of Evidence
• 2.1 Fossils
• 2.2 Host plants
• 3 Date of origin
• 4 Drivers of speciation
• 5 References

[edit] Phylogeny
The butterflies are the clade Rhopalocera that is composed of three superfamilies: Hedyloidea (the
moth butterfly family Hedylidae), the Hesperioidea (the skipper family Hesperiidae), and the Papilionoidea
(the true butterfly families Papilionidae, Pieridae, Nymphalidae, Lycaenidae, and Riodinidae). All of these
families are monophyletic. The Hedyloidea is the sister group to the other two superfamilies. Within the
Papilionoidea, Papilionidae is the sister group to the other families, and Pieridae is the sister group to
(Nymphalidae+(Lycaenidae+Riodinidae)). Phylogenetic hypotheses within the Nymphalidae are still under
discussion. Current research is concentrated on subfamilial and tribal relationships, especially in the
Nymphalidae.
 Rhopalocera
Hedyli
(position uncer

Hesperioidea
Hesperiidae

Papilionoidea

Papilionidae

Pieridae

Nymphalidae

Lycaenidae

Riodinidae
Phylogenetic relationships of butterfly families[1]
[edit] Lines of Evidence
The modern study of butterfly higher classification began with Ehrlich's phenetic use of hundreds of
previously overlooked morphological characters in tabular form, across families and major groups (Ehrlich,
1958). Scoble (1995) and others continued the search for new characters, but with their application to
cladism. Larval characters are now commonly integrated with those from adult butterflies. The addition of
molecular data has allowed researchers to resolve clades in many lineages.
Evidence is gleaned from paleontology where some 50 butterfly fossils have been identified, from
morphology and the study of homologies, from molecular genetics and comparative biochemistry, from
comparative ethology, and from present-day geographical distributions and ecology. Even though butterflies
are among the most-studied insects, new findings are coming to light almost every month, and the prospect
of a stable butterfly classification based on strongly supported clades is seemingly within reach.

[edit] Fossils
Butterfly fossils have been well covered by Grimaldi & Engel (2005), who point out their weakness in
resolving the sister group of the Rhopalocera: butterflies of 45 My are much like their living counterparts.

[edit] Host plants

Some species in the Satyrinae use ferns as larval host plants, and it is not impossible that the
butterflies could have originated before their present-day angiosperm plant hosts.
[edit] Date of origin
Some researchers theorize that butterflies most likely originated in the Cretaceous when the
continents were arrayed differently from their present positions and with climates unlike those of today. That
is when the major angiosperm radiation took place. Thus, butterfly evolution must be studied through out the
elaboration and testing of phylogenetic hypotheses and through historical zoogeography. Researchers who
accept a Cretaceous origin for the butterflies generally favor vicariant zoogeographic hypothesis for how the
major lineages of butterflies came to be distributed over the world, whereas those who favor a Tertiary age
rely on dispersalist hypotheses (Lamas, 2008).

[edit] Drivers of speciation

Mimicry, hybridization, and co-evolution with host plants have probably contributed to speciation in
the butterflies (Lamas, 2008).

[edit] References
1. ^ Lamas, Gerardo. 2008.
• Ehrlich, P. R. (1958). The comparative morphology, phylogeny and higher classification of
the butterflies (Lepidoptera: Papilionidea). Kansas Univ. Sci. Bull., 39, 305-370.
• Grimaldi, D. & Engel, M. S. 2005. Evolution of the insects. Cambridge: Cambridge Univ.
Press.
• Lamas, G. (2008) Systematics of butterflies (Lepidoptera: Hesperioidea and Papilionoidea) in
the world: current state and future perspectives (in Spanish). In: Jorge Llorente-Bousquets and
Analía Lanteri (eds.) Contribuiciones taxonómicas en ordens de insectos hiperdiversos . Mexico City:
UNAM. Pp. 57-70.
• Scoble, M. J. (1995). The Lepidoptera, form, function and diversity. London: The Natural
history Museum & Oxford University Press.
 W000

Evolution of dinosaurs
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article needs additional citations for verification.
Please help improve this article by adding reliable references. Unsourced material may be
challenged and removed. (June 2009)
This article gives an outline and examples of dinosaur evolution. For a detailed list of
interrelationships see Dinosaur classification.
 Evolution of dinosaurs
Dinosaurs evolved from the archosaurs 232-234 Ma (million years ago) in the Ladinian age, the latter
part of the middle Triassic. Dinosauria is a well-supported clade, present in 98% of bootstraps. It is
diagnosed by many features including loss of the postfrontal on the skull and an elongate deltopectoral crest
on the humerus.[1]
Contents
[hide]
• 1 From archosaurs to dinosaurs
• 2 Earliest dinosaurs
• 3 Saurischia
• 3.1 Sauropodomorpha
• 3.2 Theropoda
• 4 Ornithischia
• 4.1 Thyreophorans
• 4.2 Ornithopoda
• 4.3 Marginocephalia
• 5 Fossil record
• 6 Evolutionary trends
• 6.1 Body size
• 6.2 Mobility
• 6.3 Effect of food sources
• 7 Biogeography
• 8 Extinction
• 9 See also
• 10 References
• 11 External links
[edit] From archosaurs to dinosaurs
The process leading up the first dinosaurs can be followed through fossils of the early Archosaurs
such as the Proterosuchidae, Erythrosuchidae and Euparkeria which have fossils dating back to 250 Ma,
through mid-Triassic archosaurs such as Ticinosuchus 232-236 Ma. Crocodiles are also descendants of mid-
Triassic archosaurs.[1]
The dinosaurs can be defined as the last common ancestor of birds (Saurischia) and Triceratops
(Ornithischia) and all the descendants of that ancestor. With that definition, the pterosaurs and several
species of archosaurs narrowly miss out on being classed as dinosaurs. The pterosaurs are famous for flying
through the Mesozoic skies on leathery wings. Archosaur genera that narrowly miss out on being classified
as dinosaurs include Schleromochlus 220-225 Ma, Lagerpeton 230-232 Ma and Marasuchus 230-232 Ma.

[edit] Earliest dinosaurs

The first known dinosaurs were bipedal predators that were one to two metres long.
Spondylosoma may or may not be a dinosaur; the fossils (all postcranial) are tentatively dated at
230-232 Ma.[1]
The earliest confirmed dinosaur fossils include saurischian ('lizard-hipped') dinosaurs Saturnalia 225-
232 Ma, Herrerasaurus 220-230 Ma, Staurikosaurus possibly 225-230 Ma, Eoraptor 220-230 Ma and
Alwalkeria 220-230 Ma. Saturnalia may be a basal saurischian or a prosauropod. The others are basal
saurischians.
 Among the earliest ornithischian ('bird-hipped') dinosaurs is Pisanosaurus 220-230 Ma. Although
Lesothosaurus comes from 195-206 Ma, skeletal features suggest that it branched from the main
Ornithischia line at least as early as Pisanosaurus.
 A. Eoraptor, an early saurischian, B Lesothosaurus, a primitive ornithischian, C Staurikosaurus
(Saurischia) pelvis, D Lesothosaurus pelvis
It is clear from this figure that early saurischians resembled early ornithischians, but not modern
crocodiles. Saurischians are distinguished from the ornithischians by retaining the ancestral configuration of
bones in the pelvis. Another difference is in the skull, the upper skull of the Ornithischia is more solid and the
joint connecting the lower jaw is more flexible; both are adaptations to herbivory and both can already be
seen in Lesothosaurus.

[edit] Saurischia
Setting aside the basal Saurischia, the rest of the Saurischia are split into the Sauropodomorpha and
Theropoda. The Sauropodomorpha is split into Prosauropoda and Sauropoda. The evolutionary paths taken
by the Theropoda are very complicated. The Dinosauria (2004), a major reference work on dinosaurs,[1]
splits the Theropoda into groups Ceratosauria, Basal Tetanurae, Tyrannosauroidea, Ornithomimosauria,
Therizinosauroidea, Oviraptorosauria, Troodontidae, Dromaeosauridae and Basal Avialae in turn. Each
group branches off the main trunk at a later date. See Dinosaur_classification for the detailed
interrelationships between these.

[edit] Sauropodomorpha
The first sauropodomorphs were prosauropods. Prosauropod fossils are known from the late Triassic
to early Jurassic 227-180 Ma.[1] They could be bipedal or quadrupedal and had developed long necks and
tails and relatively small heads. They had lengths of 2.5 to 10 m and were primarily herbivorous. The earliest
prosauropods, such as Thecodontosaurus from 205-220 Ma, still retained the ancestral bipedal stance and
large head to body ratio.
These evolved into the sauropods which became gigantic quadrupedal herbivores, some of which
reached lengths of at least 26 m. Features defining this clade include a forelimb length to hindlimb length
greater than 0.6. Most sauropods still had hindlimbs larger than forelimbs; one notable exception is
Brachiosaurus whose long forelimbs suggest that it had evolved to feed from tall trees like a modern-day
giraffe.
Sauropod fossils are found from the times of the earliest dinosaurs right up to the K-T extinction
event, from 227 to 65 Ma. Most sauropods are known from the Jurassic, to be more precise between 227 and
121 Ma.
The Cretaceous sauropods form two groups. The Diplodocoidea lived from 121 to 65 Ma. The
Titanosauriformes lived from 132 to 65 Ma. The latter clade consists of series of nested subgoups, the
Titanosauria, the Titanosauridae and Saltasauridae. Both the Diplodocoidea and Titanosauriformes are
descended from the Neosauropoda, the earliest of which lived in about 169 Ma.
The sauropods are famous for being the largest land animals that ever lived, and for having relatively
small skulls. The enlargement of prosauropod and sauropod dinosaurs into these giants and the change in
skull length is illustrated in the following charts.
 Dinosaurs used in creating these charts are (in date order): Eo Eoraptor; Prosauropods Ri
Riojasaurus, Pl Plateosaurus, Yu Yunnanosaurus, Ms Massospondylus, Ji Jingshanosaurus, An
Anchisaurus, Lu Lufengosaurus, Yi Yimenosaurus, ; and Sauropods Sh Shunosaurus, Om Omeisaurus, Mm
Mamenchisaurus, Ce Cetiosaurus, Dc Dicraeosaurus, Br Brachiosaurus, Eu Euhelopus, Ap Apatosaurus, Ca
Camarasaurus, Dp Diplodocus,[1] Ha Haplocanthosaurus, Am Amargasaurus, Ar Argentinosaurus (approx),
Bo Bonitasaura, Q Quaesitosaurus, Al Alamosaurus, Sa Saltasaurus, Ra Rapetosaurus, Op
Opisthocoelicaudia, Ne Nemegtosaurus.
With the exception of Argentinosaurus (included to fill a gap in time), these graphs show only the
length of sauropods for whom near-complete fossil skeletons are known. It doesn't show other very large
sauropods (see Dinosaur size#Sauropods) because these are only known from very incomplete skeletons.
The ratio of skull length to body length is much higher in Eoraptor than in sauropods. The longest skull
graphed is of Nemegtosaurus, which is not thought be a particularly large sauropod. The skull of
Nemegtosaurus was found near the headless skeleton of 11 metre long Opisthocoelicaudia, and it has been
suggested that they may be the same species, but see Nemegtosauridae.
The relationship between the evolution of large herbivores and large plants remains uncertain. About
50% of the plants over the time of the dinosaurs were conifers, they increased in number in the Triassic until
stabilising in about 190 Ma. Cycads formed the second largest group until about 120 Ma. Ferns were present
in roughly constant numbers the whole time. Flowering plants began about 120 Ma and by the end of the
period had taken over from the cycads. All dinosaur herbivores appear to have been adversely affected by
the extinction event at the end of the Jurassic.
[edit] Theropoda
By far the earliest fossils of Theropoda (not counting the basal saurischians) are of the
Coelophysoidea, including Coelophysis and others, from late Triassic and early Jurassic 227-180 Ma.[1]
Cladistic analysis sometimes connects these to the group called Ceratosauria. Principal features of both
include changes in the pelvic girdle and hind limb that differ between the sexes. Other ceratosauria first
appear in the late Jurassic of western North America.
These are followed by the basal Tetanurae, of whom fossils have been found from the mid Jurassic
to past the end of the early Cretaceous 180 Ma to 94 Ma. They have a relatively short maxillary tooth row.
They did not all branch off the evolutionary line leading to coelurosaurs at the same time. Basal tetanurans
include Megalosauridae, spinosaurids, a diverse clade of allosaurs, and several genera of less certain
affinities, including Compsognathus. With the exception of Compsognathus they are large-bodied. Allosaurs
form a distinct long-lived clade that share some cranial characters. They include the well known Allosaurus
and Sinraptor among others.
The great radiation of Theropoda into many different clades of Coelurosauria must have happened in
the mid to late Jurassic, because Archaeopteryx was around in about 152-154 Ma, and cladistic analysis has
shown that many other groups of Coelurosauria branched off before that.[2] Fossil evidence from China
suggests that the earliest feathers were found on the primitive Coelurosauria. The most primitive of these,
e.g. on the tyrannosauroid Dilong, were simply hollow-cored fibres that would have been useful for insulation
but useless for flying.
Occasional bones and cladistic analyses point to the Tyrannosauroidea branching off from the other
Theropoda early, in the middle Jurassic, although nearly complete skeletons haven't yet appeared before
Eotyrannus from 121-127 Ma, and the many close relatives of Tyrannosaurus itself don't appear before
84 Ma, near the end of the late Cretaceous.
Ornithomimosauria fossils are known from 127 to 65 Ma. The earliest branch from the main line of
Ornithomimosauria is believed to be Harpymimus.
The Therizinosauroidea are unusual theropods in being almost all vegetarian. Fossil
Therizinosauroidea are known from 127 to 65 Ma.
Maniraptorans include Oviraptorosauria, Deinonychosaurs and birds. They are characterized by an
ulna with a curved shaft.
Oviraptorosaurian fossils are known from 127 to 65 Ma. They have a toothless skull that is extremely
modified. The skeleton has an unusually short tail.
Deinonychosaurs, named after the enlarged sickle-shaped second digit of the hand, are closely
related to birds. They have two distinct families, Troodontidae and Dromaeosauridae. Troodontid fossils are
known from 127 to 65 Ma. They have the more slender build, and longer limbs. The earliest named
troodontid fossil known is Sinornithoides. Dromaeosaurid fossils are known from about 127 to 65 Ma with the
exception of Utahraptor. The skeletal remains of Utahraptor are about 127-144 Ma. This is interesting
because according to a recent cladistic analysis,[2] Utahraptor is about as far from the ancestral Theropoda
as it is possible to get, further than Archaeopteryx. Dromaeosaurids have a larger second digit; this family
includes the well known dinosaurs Dromaeosaurus, Deinonychus and Velociraptor.
Ancient birds (Avialae) include both the Aves, which are defined as descendants of the common
ancestor of modern birds and Archaeopteryx, and the more primitive Epidendrosaurus. Fossil birds stretch
down from 154 Ma through the K-T extinction event at 65 Ma to the present day. Scores of complete
skeletons have now been found of the more recent Confuciusornis, which is an early representative of the
Ornithurae. Ornithurans all have a bony pygostyle, to which tail feathers are anchored. For more details on
the evolution of birds, see Evolution of birds.

[edit] Ornithischia
Ornithischia, as the name indicates, was coined for the birdlike pelvic girdle, although they are not
the ancestors of birds.
The ornithischian skull and dentition was modified very early by a herbivorous diet.[3] Lesothosaurus
separated early, but the skull of Lesothosaurus already shows such adaptations, with broad proportions, a
less flexible upper jaw, and a more mobile connection for the lower jaw.
The major clades were already established by the early Jurassic. The ornithischians divided into
armoured thyreophorans and unarmoured ornithopods and marginocephalians.

[edit] Thyreophorans
Surface body armour (scutes) is the most striking feature of the thyreophorans.[3] Scutellosaurus has
these but otherwise differs little from Lesothosaurus. It has a long tail and combined bipedal-quadrupedal
posture that separates it from all later thyreophorans including Stegosauria and Ankylosauria. These two
clades, although quite different in overall appearance, share many unusual features in the skull and skeleton.
Stegosaurs are easily recognised by the prominent row of plates above the spine and ling spines on
the tail. Most stegosaurs, but not Stegosaurus, also have a spine over each shoulder. These spines and
plates have evolved from the earlier surface scutes. Huayangosaurus is the oldest and most primitive known
stegosaur.
Ankylosaurs are easily recognised by their extensive body armour. The skull is heavily ossified. Early
in their evolution, ankylosaurs split into the Nodosauridae and Ankylosauridae, distinguished by features of
the skull.

[edit] Ornithopoda
Ornithopods fall into one of three distinct clades - Heterodontosauridae, Hypsilophodontidae, and
Iguanodontia.[3]
Heterodontosaurids are very small (body length < 1 m) and lived in the early to late Jurassic. Apart
from Abrictosaurus all have a short upper canine and longer lower canine. The forelimbs in known fossils are
unusually long.
Hypsilophodontids more closely resemble their ancestors than the heterodontosaurids do. The most
distinctive features are short scapula and rod-shaped pre-pubic process. The earliest is Agilisaurus from the
middle Jurassic of China.
Iguanodontians are a diverse but morphologically tight knit array of genera known from fossils of the
late Cretaceous. Significant modifications include the evolution of tooth batteries, a ligament-bound
metacarpus and a digitigrade hand posture. Tenontosaurus is the most basal iguanodontian. Others include
Iguanodon, Camptosaurus and Muttaburrasaurus.
[edit] Marginocephalia
Marginocephalia are named for a shelf that projects over the back of the skull.[3] They include the
pachycephalosaurians and ceratopsians.
Pachycephalosaurs are best known for their thick upper fronts to their skull. The oldest known is
Stenopelix, from the early Cretaceous of Europe.
Ceratopsians, famous for Protoceratops, Triceratops and Styracosaurus illustrate the evolution of
frilled and horned skulls. The frills evolved from the shelf common to all Marginocephalia. Ceratopsians are
separated into basal ceratopsians, including the parrot-beaked Psittacosaurus, and neoceratopsians.
 Diversity of ceratopsian skulls. A) Skeleton of Protoceratops. B) to I) Skulls. B) & C) Psittacosaurus
side & top. D) & E) Protoceratops side & top. F) & G) Triceratops side & top. H) & I) Styracosaurus side
(without lower jaw) & top.
The sequence of ceratopsian evolution in the Cretaceous is roughly from Psittacosaurus (121
-99 Ma) to Protoceratops (83 Ma) to (Triceratops 67 Ma and Styracosaurus 72 Ma). In side view the skull of
Psittacosaurus bears very little resemblance to that of Styracosaurus but in top view a similar pentagonal
arrangement can be seen.

[edit] Fossil record

The first few lines of primitive dinosaurs diversified rapidly through the Triassic period; dinosaur
species quickly evolved the specialised features and range of sizes needed to exploit nearly every terrestrial
ecological niche. During the period of dinosaur predominance, which encompassed the ensuing Jurassic and
Cretaceous periods, nearly every known land animal larger than 1 meter in length was a dinosaur.
One measure of the quality of the fossil record is obtained by comparing the date of first appearance
with the order of branching of a cladogram based on the shape of fossil elements. Close correspondence
exists for ornithiscians, saurischians and subgroups.[3] The cladogram link between coelophysids and
ceratosaurs is an exception, it would place the origin of coelophysids much too late. The simplest explanation
is convergent evolution - ceratosaur bones evolved independently into a shape that resembles that of the
earlier coelophysids. The other possibility is that ceratosaurs evolved much earlier than the fossil record
suggests.
 Most dinosaur fossils have been found in the Norian-Sinemurian, Kimmeridgian-Tithonian, and
Campanian-Maastrichtian periods. Continuity of lineages across the intervening gaps shows that those gaps
are artifacts of preservation rather than any reduction in diversity or abundance.
In many instances, cladistic analysis shows that ancestral lineages of varying durations fall in those
gaps. The length of missing ancestral lineages in 1997[3] range from 25 Ma (Lesothosaurus, Genasauria,
Hadrosauroidea, Sauropoda, Neoceratopsia, Coelurosauria) to 85 Ma (Carcharodontosauridae). Because
the dinosaurian radiation began at small body size, the unrecorded early history may be due to less reliable
fossilization of smaller species. However, some missing lineages, notably of Carcharodontosauridae and
Abelisauridae, require alternative explanations because the missing range extends across stages rich in
fossil material.

[edit] Evolutionary trends

[edit] Body size
Body size is important because of its correlation with metabolism, diet, life history, geographic range
and extinction rate.[3] The modal body mass of dinosaurs lies between 1 and 10 tons throughout the
Mesozoic and across all major continental regions. That said, there was a trend towards increasing body size
within many dinosaur clades, including the Thyreophora, Ornithopoda, Pachycephalosauria, Ceratopsia,
Sauropomorpha, and basal Theropoda. Marked decreases in body size have also occurred in some lineages,
but are more sporadic. The best known example is the decrease in body size leading up to the first birds;
Archaeopteryx was below 10 kg in weight, and later aves Confuciusornis and Sinornis are starling- to pigeon-
sized.
[edit] Mobility
The ancestral dinosaur was a biped. The evolution of a quadrupedal posture occurred four times,
among the ancestors of Euornithopoda, Thyreophora, Ceratopsia and Sauropodomorpha.[3] In all four cases
this was associated with an increase in body size, and in all four cases the trend is unidirectional without
reversal.
Dinosaurs exhibit a pattern of the reduction and loss of fingers on the lateral side of the hand (digits
III, IV and V). The primitive function of the dinosaur hand is grasping with a partially opposable thumb, rather
than weight-bearing.

[edit] Effect of food sources

The ancestral dinosaur was a carnivore. Herbivory among dinosaurs arose three times, at the origin
of the ornithischian, sauropodomorph, and therizinosaurid clades. Individual therizinosaurids are herbivorous
or omnivorous. Herbivory among the ornithischians and sauropodomorphs was never reversed.[3]
The potential co-evolution of plants and herbivorous dinosaurs has been subject to extensive
speculation. The appearance of prosauropods in the late Triassic has been tentatively linked either to the
demise or diversification of types of flora at that time. The rise of ceratopsids and iguanodont and
hadrosaurid ornithopods in the Cretaceous has been tentatively linked to the angiosperm radiation.
Unfortunately, there are still no hard data on dietary preferences of herbivorous dinosaurs, apart from data on
chewing technique and gastroliths.
[edit] Biogeography
Dinosaurian faunas, which were relatively uniform in character when Pangaea began to break up,
became markedly differentiated by the close of the Cretaceous. Biogeography is based on the splitting of an
ancestral species by the emplacement of a geographic barrier. Interpretation is limited by a lack of fossil
evidence for eastern North America, Madagascar, India, Antarctica and Australia. No unequivocal proof of
the biogeographical action on Dinosaur species has been obtained,[3] but some authors have outlined
centres of origin for many dinosaur groups, multiple dispersal routes, and intervals of geographic isolation.
Dinosaurs that have been given as evidence of biogeography include abelisaurid theropods from
South America and possibly else where on Gondwana.
Relationships between dinosaurs show abundant evidence of dispersal from one region of the globe
to another. Tetanuran theropods travelled widely through western North America, Asia, South America, Africa
and Antarctica. Pachycephalosaurs and ceratopsians show clear evidence of multiple bidirectional dispersion
events across Beringa.

[edit] Extinction
The Cretaceous–Tertiary extinction event, which occurred 65.5 million years ago at the end of the
Cretaceous period, caused the extinction of all dinosaurs except for the line that had already given rise to the
first birds.
[edit] See also
Dinosaurs portal

• Dinosaur classification
• Origin of birds
• List of fossil birds

[edit] References
1. ^ a b c d e f g Weishampel, Dodson & Osmolska, 2004, The Dinosauria
2. ^ a b Senter, P. (2007). "A new look at the phylogeny of Coelurosauria (Dinosauria:
Theropoda)." Journal of Systematic Palaeontology, (doi:10.1017/S1477201907002143)
3. ^ a b c d e f g h i j PC Sereno (1997) "The origin and evolution of dinosaurs" Annu. Rev. Earth
Planet. Sci. 25:435-489
Paul C. Sereno (1999) The evolution of dinosaurs, Science, Vol 284, pp. 2137–2146
http://www.sciencemag.org/cgi/content/abstract/284/5423/2137
[edit] External links
[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
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 Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
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Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_dinosaurs"

Categories: Dinosaurs | Evolution by taxon

 W000

Histoire évolutive des dinosaures

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Les dinosaures ont, selon la théorie de l'évolution, évolué à partir d'archosauriens il y a 232 à 234 Ma
c'est-à-dire de la fin du Trias moyen.
Les dinosaures sont bien connus, de nombreux fossiles ont été découverts, principalement ceux des
grandes espèces. Les dinosaures se reconnaissent par de nombreuses caractéristiques comme la présence
de 2 fosses temporales, l'élongation de la crête deltopectorale sur l'humérus. Il forment une lignée différente
de celle des crocodiliens et des ptérosaures, eux aussi issus des archosauriens.
Cette classe d'animaux va disparaitre à la fin du Crétacé, à l'exception des oiseaux.
Sommaire
[masquer]
• 1 Évolution
• 1.1 Les archosaures ancestraux
• 1.2 Présentation d'un arbre évolutif
• 2 Notes et références
• 3 Voir aussi

Évolution [modifier]
Les archosaures ancestraux [modifier]
Les dinosaures semblent être issus d'une série d'archosauriens dont les plus anciens sont les
Proterosuchidae, Erythrosuchidae et Euparkeria dont les fossiles sont datés de 250 Ma, et jusqu'aux fossiles
de Ticinosuchus, datés eux de 232-236 Ma.
Présentation d'un arbre évolutif [modifier]
 Évolution des dinosaures[1].

Notes et références [modifier]

1. ↑ Based on Polish encyclopedia Wielka Encyklopedia PWN, tom 7, str. 190, wydanie 2001-
2005.

Voir aussi [modifier]

[Enrouler]
v·d·m
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive
 Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie
l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation
 • Portail de la paléontologie

• Portail de l’origine et de l’évolution du vivant

• Portail des dinosaures

• Portail de la zoologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_dinosaures ».

Catégories : Histoire évolutive | Dinosaure | [+]

 W000

Evolution of cetaceans
From Wikipedia, the free encyclopedia

Jump to: navigation, search

 The approximately 80 modern species in the order Cetacea.
The cetaceans (whales, dolphins and porpoises) are marine mammal descendants of land mammals.
Their terrestrial origins are indicated by:
• Their need to breathe air from the surface;
• The bones of their fins, which resemble the limbs of land mammals; and
• The vertical movement of their spines, characteristic more of a running mammal than of the
horizontal movement of fish.
 The question of how land animals evolved into ocean-going leviathans had been a mystery for a long
time, owing to gaps in the fossil record. However, recent discoveries in Pakistan have clarified many of these
questions, and revealed several stages in the transition of the cetaceans from land to sea.

Contents
[hide]
• 1 Earliest ancestors
• 2 The earliest cetaceans: Pakicetids or Indohyus?
• 3 Ambulocetids and remingtonocetids
• 4 Protocetids
• 5 Basilosaurids and dorudontids: fully marine cetaceans
• 6 Early echolocation
• 7 Early baleen whales
• 8 Early dolphins
• 9 Skeletal evolution
• 10 See also
• 11 References
• 12 External links
[edit] Earliest ancestors

The family tree, including Ungulates.

The traditional theory of cetacean evolution was that whales were related to the mesonychids, an
extinct order of carnivorous ungulates (hoofed animals), which looked rather like wolves with hooves and
were a sister group of artiodactyls. These animals possessed unusual triangular teeth that are similar to
those of whales. For this reason, scientists had long believed that whales evolved from a form of
mesonychid; however more recent molecular phylogeny data suggest that whales are more closely related to
the artiodactyls, specifically the hippopotamus.[1] The strong evidence for a clade combining cetaceans and
artiodactyls is further discussed under the entry Cetartiodactyla. However, hippos' anthracothere ancestors
do not appear in the fossil record until millions of years after Pakicetus, the first known whale ancestor.
The recent discovery of Pakicetus, the earliest proto-whale (see below) supports the molecular data.
The skeletons of Pakicetus demonstrate that whales did not derive directly from mesonychids. Instead, they
are a form of artiodactyl (another type of ungulate) that began to take to the water after the artiodactyl family
split from the mesonychids. In other words, the proto-whales were early artiodactyls that retained aspects of
their mesonychid ancestry (such as the triangular teeth) which modern artiodactyls have since lost. An
interesting implication is that the earliest ancestors of all hoofed mammals were probably at least partly
carnivorous or scavengers, today's artiodactyls and perissodactyls having switched to a plant diet later in
their evolution. Whales, due to the readier availability of animal prey and their need for higher caloric content,
in order to live as marine endotherms, naturally retained their carnivorous diet, as did mesonychids, who
were however out-competed by better-adapted animals like the Carnivora later on (mesonychids became
specialized carnivores when the overall availability of large animal prey was still low; thus their adaptation
was likely at a disadvantage when new forms had filled the gaps left by the dinosaurs).
[edit] The earliest cetaceans: Pakicetids or Indohyus?

Reconstruction of Pakicetus
The pakicetids are hoofed-mammals that are sometimes classified as the earliest whales.[2][3] They
lived in the early Eocene, around 53 million years ago. They looked rather like dogs with hoofed feet and
long, thick tails. They have been linked to whales by their ears: the structure of the auditory bulla is formed
from the ectotympanic bone only. The shape of the ear region in Pakicetus is highly unusual and only
resembles the skulls of whales. The feature is diagnostic for cetaceans and is found in no other species. It
was initially thought that the ears of Pakicetus were adapted for underwater hearing, but, as would be
expected from the anatomy of the rest of this creature, the ears of Pakicetus are specialized for hearing on
land, and if Pakicetus is related to the ancestors of whales, underwater hearing must have been a later
adaptation.[4] According to Thewissen, the teeth of Pakicetus also resemble the teeth of fossil whales, being
less like a dog's incisors, with a serrated triangular shape, similar to a shark's tooth, which is another link to
more modern whales.[5]
 Reconstruction of Indohyus.
Thewissen has since found the same ear structure in fossils of a small deer-like creature, Indohyus,
which lived about 48 million years ago in Kashmir.[3] About the size of a raccoon or domestic cat, this
herbivorous creature shared some of the traits of whales, and showed signs of adaptations to aquatic life,
including a thick and heavy outer coating to bones which is similar to the bones of modern creatures such as
the hippopotamus,[6][7] and reduces buoyancy so that they can stay underwater. This suggests a similar
survival strategy to the African mousedeer or water chevrotain which, when threatened by a bird of prey,
dives into water and hides beneath the surface for up to four minutes.[8][9][10]
[edit] Ambulocetids and remingtonocetids

Reconstruction of Kutchicetus, a remingtonocetid

Reconstruction of Ambulocetus natans

The most remarkable of the recent discoveries in Pakistan has been Ambulocetus, which looked like
a three-metre long mammalian crocodile. Ambulocetus was clearly amphibious, as its back legs are better
adapted for swimming than for walking on land, and it probably swam by undulating its back vertically, as
otters, seals and whales do. It has been speculated that Ambulocetids hunted like crocodiles, lurking in the
shallows to snatch unsuspecting riparian prey and fish.
A smaller cousin of Ambulocetus was the remingtonocetid family, which had longer snouts than
Ambulocetus, and were slightly better adapted for underwater life. It has been speculated that they lived like
modern sea otters, hunting for fish in the shallows.
In both groups, the nasal openings were at the tip of the snout, like in land-mammals.

[edit] Protocetids

Reconstruction of Protocetus
 Reconstruction of Rodhocetus
The protocetids form a diverse and heterogeneous group known from Asia, Europe, Africa, and North
America. There were many genera, and some of these are very well known (e.g., Rodhocetus). Known
protocetids had large fore- and hindlimbs that could support the body on land, and it is likely that they lived
amphibiously: in the sea and on land. It is unclear at present whether protocetids had flukes (the horizontal
tail fin of modern cetaceans). However, what is clear is that they are adapted even further to an aquatic life-
style. In Rodhocetus, for example, the sacrum – a bone that in land-mammals is a fusion of five vertebrae that
connects the pelvis with the rest of the vertebral column – was divided into loose vertebrae. However, the
pelvis was still connected to one of the sacral vertebrae. Furthermore, the nasal openings are now halfway
up the snout; a first step towards the telescoped condition in modern whales. Their supposed amphibious
nature is supported by the discovery of a pregnant Maiacetus[11], in which the fossilised fetus was positioned
for a head-first delivery, suggesting that Maiacetus gave birth on land. The ungulate ancestry of these early
whales is still underlined by characteristics like the presence of hoofs at the ends of toes in Rodhocetus.
[edit] Basilosaurids and dorudontids: fully marine cetaceans

Reconstruction of Basilosaurus
Basilosaurus (discovered in 1840 and initially mistaken for a reptile, hence its name) and Dorudon
lived around 38 million years ago, and were fully recognizable whales which lived entirely in the ocean.
Basilosaurus was as big as the larger modern whales, up to 18 m (60 ft) long; dorudontids were smaller,
about 5 m (16 ft) long.
 Reconstruction of Dorudon
Although they look very much like modern whales, basilosaurids and dorudontids lacked the 'melon
organ' that allows their descendants to use echolocation as effectively as modern whales. They had small
brains; this suggests they were solitary and did not have the complex social structure of some modern
cetaceans. Basilosaurus had two tiny but well-formed hind legs which were probably used as claspers when
mating; they are a small reminder of the lives of their ancestors. Interestingly, the pelvic bones associated
with these hind limbs was now no longer connected to the vertebral column as it was in protocetids.
Essentially, any sacral vertebrae can no longer be clearly distinguished from the other vertebrae.

[edit] Early echolocation

Reconstruction of Squalodon
Toothed whales (Odontocetes) echolocate by creating a series of clicks emitted at various
frequencies. Sound pulses are emitted through their melon-shaped foreheads, reflected off objects, and
retrieved through the lower jaw. Skulls of Squalodon show evidence for the first hypothesized appearance of
echolocation. Squalodon lived from the early to middle Oligocene to the middle Miocene, around 33-14
million years ago. Squalodon featured several commonalities with modern Odontocetes. The cranium was
well compressed, the rostrum telescoped outward (a characteristic of the modern suborder Odontoceti),
giving Squalodon an appearance similar to that of modern toothed whales. However, it is thought unlikely
that squalodontids are direct ancestors of living dolphins.

[edit] Early baleen whales

Reconstruction of Cetotherium
All modern mysticetes are large filter-feeding or baleen whales, though the exact means by which
baleen is used differs among species (gulp-feeding with balaenopterids, skim-feeding with balaenids, and
bottom ploughing with eschrichtiids). The first members of some modern groups appeared during the middle
Miocene. These changes may have been a result of worldwide environmental change and physical changes
in the oceans. A large scale change in ocean current and temperature could have initiated the radiation of
modern mysticetes, leading to the demise of the archaic forms. Generally it is speculated the four modern
mysticete families have separate origins among the cetotheres. Modern baleen whales, Balaenopteridae
(rorquals and humpback whale, Megaptera novaengliae), Balaenidae (right whales), Eschrichtiidae (gray
whale, Eschrictius robustus), and Neobalaenidae (pygmy right whale, Caperea marginata) all have derived
characteristics presently unknown in any cetothere.

[edit] Early dolphins

Skeleton of Xiphiacetus sp.

During the early Miocene (about 20 Ma), echolocation developed in its modern form. Various extinct
dolphin-like families flourished. Early dolphins include Kentriodon and Hadrodelphis. These belong to
Kentriodontidae, which were small to medium-sized toothed cetaceans with largely symmetrical skulls, and
thought likely to include ancestors of some modern species. Kentriodontids date to the late Oligocene to late
Miocene. Kentriodontines ate small fish and other nectonic organisms; they are thought to have been active
echolocators, and might have formed schools. Diversity, morphology and distribution of fossils appear
parallel to some modern species.

[edit] Skeletal evolution

Today, the whale hind parts are internal and reduced. Occasionally, the genes that code for longer
extremities cause a modern whale to develop miniature legs (known as atavism).
Whereas early cetaceans such as the Pakicetus had the nasal openings at the end of the snout, in
later species such as the Rodhocetus, the openings had begun to drift toward the top of the skull. This is
known as nasal drift.
The nostrils of modern whales have become modified into blowholes that allow them to break to the
surface, inhale, and submerge with convenience. The ears began to move inward as well, and, in the case of
Basilosaurus, the middle ears began to receive vibrations from the lower jaw. Today's modern toothed
whales use the 'melon organ', a pad of fat, for echolocation.

[edit] See also

Cetaceans portal

• Dolphin, Evolution and anatomy

• Archaeoceti
• Aquatic adaptation
• Indohyus
 • Transitional form
• Evolution of sirenians
• Evolution of mammals
• List of extinct cetaceans

[edit] References
1. ^ University Of California, Berkeley (2005, February 7). "UC Berkeley, French Scientists Find
Missing Link Between The Whale And Its Closest Relative, The Hippo". ScienceDaily.
http://www.sciencedaily.com/releases/2005/02/050205103109.htm. Retrieved 2007-12-21.
2. ^ Philip D. Gingerich, D. E. Russell (1981). "Pakicetus inachus, a new archaeocete
(Mammalia, Cetacea) from the early-middle Eocene Kuldana Formation of Kohat (Pakistan)". Univ.
Mich. Contr. Mus. Paleont 25: 235–246.
3. ^ a b Northeastern Ohio Universities Colleges of Medicine and Pharmacy (2007, December
21). "Whales Descended From Tiny Deer-like Ancestors". ScienceDaily.
http://www.sciencedaily.com/releases/2007/12/071220220241.htm. Retrieved 2007-12-21.
4. ^ J. G. M. Thewissen, E. M. Williams, L. J. Roe and S. T. Hussain (2001). "Skeletons of
terrestrial cetaceans and the relationship of whales to artiodactyls". Nature 413 (6853): 277–281.
doi:10.1038/35095005. PMID 11565023.
5. ^ Whale Origins
6. ^ University Of Michigan (2001, September 20). "New Fossils Suggest Whales And Hippos
Are Close Kin". ScienceDaily. http://www.sciencedaily.com/releases/2001/09/010920072245.htm.
Retrieved 2007-12-21.
 7. ^ University Of California, Berkeley (2005, February 7). "UC Berkeley, French Scientists Find
Missing Link Between The Whale And Its Closest Relative, The Hippo". ScienceDaily.
http://www.sciencedaily.com/releases/2005/02/050205103109.htm. Retrieved 2007-12-21.
8. ^ Carl Zimmer (2007-12-19). "The Loom : Whales: From So Humble A Beginning...".
ScienceBlogs. http://scienceblogs.com/loom/2007/12/19/whales_from_so_humble_a_beginn.php.
Retrieved 2007-12-21.
9. ^ Ian Sample (2007-12-19). "Whales may be descended from a small deer-like animal".
Guardian Unlimited. http://www.guardian.co.uk/science/2007/dec/19/whale.deer?
gusrc=rss&feed=networkfront. Retrieved 2007-12-21.
10.^ PZ Myers (2007-12-19). "Pharyngula: Indohyus". Pharyngula. ScienceBlogs.
http://scienceblogs.com/pharyngula/2007/12/indohyus.php. Retrieved 2007-12-21.
11.^ Gingerich PD, ul-Haq M, von Koenigswald W, Sanders WJ, Smith BH, et al.. "New
Protocetid Whale from the Middle Eocene of Pakistan: Birth on Land, Precocial Development, and
Sexual Dimorphism". PLoS one. http://www.plosone.org/article/info%3Adoi
%2F10.1371%2Fjournal.pone.0004366. Retrieved 2009-02-04.

[edit] External links

• Timeline of Whale Evolution - Smithsonian Ocean Portal
• Cetacean Paleobiology - University of Bristol
• BBC: Whale's evolution
• Hooking Leviathan by Its Past by Stephen Jay Gould
• Whale Origins, Thewissen Lab, Northeastern Ohio Universities College of Medicine
• Digital Library of Dolphin Development, Thewissen Lab
 • Research on the Origin and Early Evolution of Whales (Cetacea) , Gingerich, P.D., University
of Michigan
• Evolution of Whales Adapted from National Geographic, November 2001, Revised 2006 Dr.
J.G.M. Thewissen
• Pakicetus inachus, a new archaeocete (Mammalia, Cetacea) from the early-middle Eocene
Kuldana Formation of Kohat (Pakistan). Gingerich, P.D., 1981, Museum of Paleontology, The
University of Michigan
• Skeletons of terrestrial cetaceans and the relationship of whales to artiodactyls , Nature 413,
277-281 (20 September 2001), J. G. M. Thewissen, E. M. Williams, L. J. Roe and S. T. Hussain
• Evolution of Whales segment from the Whales Tohorā Exhibition Minisite of the Museum of
New Zealand Te Papa Tongarewa
 W000

Histoire évolutive des cétacés

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Traduction à relire --- Evolution of cetaceans → Histoire évolutive des cétacés --- (+)
• Statut de la traduction : À relire
• Commentaire : Intéresse le projet du vivant
• Demandeur : Vincnet G 31 mai 2007 à 12:00 (CEST)
 • Intérêt de la traduction : article complet sur un sujet populaire, baleine, dauphin... et est
susceptible d'être intégré dans plusieurs dizaines d'articles zoologiques mais aussi a des articles en
paléontologie, histoire des sciences...
• Projets : OEV , Zoologie

• Traducteur(s) : Trassiorf (d)

• Avancement de la traduction : ██████████100 %
• Version traduite : 21 janvier 2010
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Page de suivi de traduction --- Mettre à jour ces informations

Au IVs siècle av. J.-C., Aristote classait les cétacés parmi les poissons. Or, bon nombre de
caractères anatomiques et physiologiques les rapprochent des mammifères terrestres comme par exemple
leurs poumons, l'agencement des os de leurs nageoires, les articulations des vertèbres lombaires, ainsi que
l'alimentation lactée des nouveau-nés.
Sommaire
[masquer]
• 1 Relations entre cétacés et autres mammifères
• 2 Ambulocétidés et remingtonocétidés
• 3 Protocetus
• 4 Basilosauridés et dorudontidés: des cétacés entièrement
marins
• 5 Les débuts de l'écholocation
• 5.1 Les premiers dauphins
• 5.2 Premières baleines à fanons (mysticètes)
• 5.3 Évolution du squelette
• 5.4 Atavismes ancestraux
• 5.5 Arbre phylogénétique
• 6 La pression humaine
• 7 Notes et références
• 8 Voir aussi
• 8.1 Articles connexes
• 8.2 Liens externes
Relations entre cétacés et autres mammifères [modifier]
XVIIIe
C'est à partir du siècle que la question de l'origine de ces animaux, reconnus comme
mammifères, a été posée. La paléontologie a permis, grâce à la découverte de plusieurs fossiles, de
comprendre comment les ancêtres de ces animaux sont passés d'un milieu terrestre à un milieu totalement
marin. Cette transition est considérée comme très rapide dans la mesure où elle se serait faite en 10 millions
d'années environ. Jussieu est le premier à séparer les cétacés des poissons ; Linné les reconnaît comme
mammifères. Les cétacés vont poser une énigme aux évolutionnistes du milieu du XVIIIe siècle : dans quel
milieu vivait l'ancêtre commun à tous les mammifères, et quand les ancêtres des cétacés sont-ils devenus
marins ? Lacépède publie son Histoire des cétacés en 1804, après son Histoire naturelle des poissons. Au
vu des fossiles de vertèbres du Basilosaurus découverts en Alabama aux États-Unis au début du
XIXe siècle, Richard Harlan pense qu'il s'agit d'un squelette de serpent de mer, c'est pourquoi son nom se
termine par -saurus, c’est-à-dire reptile. Cette erreur est corrigée par Richard Owen qui reconnaît un
mammalien qu'il rebaptise Zeuglodon.
À la fin du XIXe siècle[1], des auteurs suggèrent une parenté avec les ongulés, et que ce taxon de
cétacés est monophylétique. Même si, jusqu'aux années 1960, plusieurs auteurs continuent à penser que les
mysticètes et les ondontocètes sont de lignées différentes[2], la théorie de base sur l'évolution des cétacés
est qu'ils descendent des mésonychidés, un ordre éteint d'ongulés (animaux à sabots) carnivores
charognards, ressemblant en gros à des loups munis de sabots, et proches parents des artiodactyles. La
raison en était que ces animaux possédaient des dents triangulaires inhabituelles, semblables à celles des
dauphins. En outre, la présence de traits synapomorphiques comme plusieurs préestomacs chez les cétacés
et les artiodactyles actuels renforçait cette hypothèse.
 Cependant ces données sont infirmées par des études immunologiques dès 1950[3]. La découverte
de nombreux fossiles dans cette même décennie permet de retracer plus précisément l'histoire des cétacés.
Durant la même période, la position des cétacés par rapport aux Artiodactyla est vivement discutée.
Ces informations se trouvent confirmées par la morphologie, très particulière, des chevilles d'un
fossile du début de l'Éocène découvert au Pakistan. Cet animal fossile appelé Pakicetus, découvert en 1983,
est celui d'un mammifère terrestre carnivore. Son astragale démontre que les cétacés n'ont pas évolué à
partir des Mesonychidae mais après une différenciation au sein d'un groupe formé des ancêtres des
Artiodactyla[4] (Chriacus ?). Les premiers cétacés sont des artiodactyles primitifs, qui ont gardé de leurs
ancêtres mésonychidés des caractères tels que les dents triangulaires, que les artiodactyles modernes ont
perdues depuis en devenant herbivores.
Au cours des années 1990, en outre, l'analyse phylogénétique des gènes codant certaines protéines
mitochondriales les a regroupés au sein d'un même clade avec les Artiodactyles (Ongulés à doigts pairs)[5].
Des données plus récentes en phylogénétique moléculaire suggèrent que les cétacés sont plus
proches, chez les artiodactyles, spécialement des hippopotames[6]. Ces différentes raisons poussent à créer
le nouvel ordre des Cetartiodactyla qui regroupe les Artiodactyla et Cetacea[7].
Ceci a depuis été confirmé par l'étude de caractères dérivés dans leur génome nucléaire (les
Rétrotransposons)[8].
Une étude basée sur la morphologie remet cependant en question cette proche parenté et suggère
une plus grande proximité avec les Anthracothères fossiles et une évolution des hippopotames à partir d'une
autre branche, au sein des Cetartiodactyla à déterminer[9]. Les cétacés auraient donc divergé très tôt des
Anthracothères, à l'instar des suidés et des ruminants. Il faut cependant préciser que cette étude n'a inclus
que des représentants fossiles des cétacés.
 On a découvert ensuite plusieurs groupes d'espèces similaires au Pakicetus inachus, datés de 55 à
34 Ma. De la taille d'un renard à celle d'un loup, elles ont été regroupées dans la famille des Pakicetidae.
Certains auteurs les classent parmi les cétacés les plus primitifs, dans le sous-ordre des Archaeoceti[10],
[11].
Ils ressemblaient à des chiens à sabots, avec de longues queues épaisses. Ils sont reliés aux
cétacés par leurs oreilles : la structure du logement osseux de leur appareil auditif, entièrement situé dans
l'os tympanal. Cette structure est hautement inusuelle, et ne ressemble qu'au crâne des cétacés. En fait c'est
une structure caractéristique des cétacés, que l'on ne retrouve dans aucun autre ordre.
Thewissen a trouvé depuis la même structure d'oreille dans les fossiles d'un petit mammifère
ressemblant à un cerf, l'Indohyus, vivant il y a 48 Ma au Cachemire[11]. De la taille d'un petit chien, cet
animal partageait certains traits des cétacés, mais était herbivore. Sa position dans l'arbre généalogique des
cétacés n'est donc pas claire. Ses os lourds lui servaient probablement de lest pour la plongée, et il avait
sans doute des habitudes aquatiques. Ce caractère se retrouve chez les hippopotames[12],[6]. Ceci peut
aussi correspondre à une stragégie de survie consistant à se cacher dans l'eau en cas de danger[13],[14],
[15].
Les ancêtres anthracothères des hippopotames n'apparaissent parmi les fossiles que des millions
d'années après Pakicetus.
Le tableau qui émerge de ces considérations est que les ancêtres primitifs de tous les mammifères
ongulés étaient probablement en partie carnivores ou charognards. Les artiodactyles et périssodactyles se
sont tournés vers un régime herbivore ultérieurement dans l'évolution, ceci évidemment accompagné d'un
changement de structure de leurs dents. Les cétacés, en raison de la plus grande abondance de proies
animales, et de leurs besoins métaboliques plus élevés pour survivre comme homéothermes dans l'eau, ont
naturellement gardé leur régime carnivore, comme l'avaient fait les mésonichydés, qui ont été éliminés au
profit d'animaux mieux adaptés, comme les carnivores.

Ambulocétidés et remingtonocétidés [modifier]

Reconstruction d’Ambulocetus natans

En 1994, on a fait la plus remarquable découverte, toujours au Pakistan, celle du fossile
d'Ambulocetus clairement plus récent que ceux de Pakicetus[16]. C'est un mammifère d'environ 3 m de long
et aux pattes orientées vers l'extérieur, munies de petits sabots. Quoique clairement amphibie, il devait nager
en ondulant l'arrière verticalement comme les loutres, les phoques et les cétacés, et être mal à l'aise sur la
terre ferme. Alors que les Pakicetidae devaient vivre en eau douce, d'après certains travaux[17] sur la
composition isotopique des os, les Ambulocetus vivaient dans une grande variété de salinités[16]. On a fait
l'hypothèse que les ambulocétidés chassaient comme les crocodiles, tapis dans les lagunes pour attraper les
imprudents, proies sur les bords, ou poissons.
 Reconstruction de Kutchicetus, un remingtonocetidé
Des cousins plus petits de l’ambulocetus formaient la famille des remingtonocétidés, qui possédaient
des museaux plus longs qu’Ambulocetus, et étaient un peu mieux adaptés à la vie sous-marine. On pense
qu'ils devaient avoir une vie analogue à celle des loutres de mer. L'examen des isotopes de leurs fossiles
montre que tous les Remingtonocetidea trouvés vivaient dans un milieu marin[16]. Dans les deux familles,
les narines se trouvaient à l'extrémité du museau, comme chez les mammifères terrestres.
Protocetus [modifier]

Reconstruction de Rodhocetus
Présents il y a 45 Ma, les protocetidae forment une famille abondante et hétérogène dont on trouve
des fossiles sur tous les continents. Le plus connu des genres étant celui des Rodhocetus. Ils disposaient de
pattes plutôt adaptées à la nage mais leur laissant peut-être la possibilité de se déplacer sur le sol. On ne
sait pas s'ils disposaient d'une nageoire caudale (horizontale comme les cétacés modernes).
Les protocétidés connus avaient de solides membres antérieurs et postérieurs, qui pouvaient
soutenir le corps à terre, et il est probable qu'ils avaient une vie amphibie. Il est clair cependant qu'ils étaient
mieux adaptés que leurs ancêtres à un style de vie aquatique. Chez rodhocetus, par example, le sacrum, qui
chez les mammifères terrestres provient de la fusion de cinq vertèbres, reliant le pelvis avec le reste de la
colonne vertébrale pour former le bassin, était divisé en vertèbres indépendantes, le pelvis n'étant plus
connecté à la colonne vertébrale que par une vertèbre. De plus, ses narines étaient à mi-chemin du museau,
première étape vers la position au sommet du crâne chez les cétacés modernes.
 Reconstruction de Protocetus
L'hypothèse d'une vie amphibie est soutenue par la découverte d'une femelle pleine de
Maiacetus[18], chez laquelle le fœtus avait la position pour être mis bas par la tête, ce qui suggère que
maiacetus mettait bas à terre.
L'appartenance des ancêtres de ces cétacés primitifs aux ongulés est rappelée par des caractères
comme la présence de sabots au bout des doigts de Rodhocetus.
Basilosauridés et dorudontidés: des cétacés entièrement marins [modifier]

Reconstruction de basilosaurus
Le Basilosaurus (découvert en 1840 et initialement pris pour un reptile, d'où le nom), un animal
jusqu'à 18 m de long, apparaît il y a 40 Ma, mais on ne connaît pas précisément son lien de parenté avec les
cétacés. Strictement marin, il conservait deux moignons de pattes arrières, que l'on pense avoir joué un rôle
pour l'accouplement, et qui n'étaient plus reliés à la colonne vertébrale par les os pelviens. Pour l'essentiel,
leurs vertèbres sacrées ne sont plus discernables clairement des autres vertèbres.
Les premiers fossiles de dorudontidés dont on dispose sont datés de 38 Ma, ces animaux ne
mesurent eux que 5 à 6 m. Ce sont des cétacés parfaitements typiques, qui vivaient entièrement dans
l'océan.
 Reconstruction de dorudon
On n'a pas trouvé chez ces mammifères de melon ; on les suppose donc incapables de chant ou
d'écholocation. En outre, ils étaient dotés d'un cerveau petit par rapport aux cétacés modernes[19]. Ce
dernier trait suggère qu'ils étaient solitaires, et n'avaient pas la structure sociale complexe de certains
cétacés modernes.
Les études moléculaires conduites sur les baleines à fanon laissent penser que la divergence qui a
vu leur apparition s'est faite à la fin de l’éocène, il y a 35 à 36 Ma. Leur apparition pourrait être liée à un
changement environnemental marin mondial, peut être lié aux changements de courants et de climat.
Article détaillé : paléoclimatologie.
Les Zygorhiza sont les cétacés les plus modernes qui semblent disposer à la fois des traits des
odontocètes et des mysticètes. Ils vivaient il y a 45 Ma. On pense que les deux sous-ordres ont divergé à la
charnière éocène-oligocène.
Les espèces ont ensuite peu évolué pendant une longue période. Une nouvelle phase de spéciation
est alors survenue au milieu du miocène, il y a 12 à 15 Ma.
Les débuts de l'écholocation [modifier]

Reconstruction d'un squalodon

Les crânes de squalodon sont les plus anciens à montrer la première apparition probable
d'écholocation. Ce cétacé vivait de l'oligocène moyen au miocène moyen soit il y a de 33 à 14 Ma. Le
squalodon montre plusieurs similitudes avc les odontocètes : le crâne est bien comprimé, le museau
télescopé en-dehors (une caractéristique des odontocètes modernes), ce qui lui donnait une apparence de
cétacé à dents. Ces animaux se font un panorama de leur environnement par écholocation sur le principe du
sonar, c'est à dire en émettant une série de clics à diverses fréquences. Ces impulsions sonores sont émises
par leur front en forme de melon, réfléchies par les objets, et recueillies, pour les hautes fréquences, par la
mâchoire inférieure, d'où elles se propagent vers l'oreille.
Cependant il est improbable que ce soient des ancêtres des dauphins modernes.
Les premiers dauphins [modifier]

Squelette d'un Xiphiacetus sp.

Au début du Miocène de nombreuses espèces d'odontocètes ressemblant aux dauphins se sont
développées comme les kentriodons et hadrodelphis. La plupart sont maintenant éteintes. On les classe
dans les Kentriodontidae et ils ont la particularité d'avoir des crânes symétriques. Ils pourraient s'être déjà
organisés en troupeaux. Ils consommaient des petits poissons et leur système d'écholocation devait être
parfaitement opérationnel. La morphologie et la distribution de ces fossiles sont très semblables aux
odontocètes modernes, qui pourraient y trouver leurs ancêtres. Les Odobenocetops sont des exemples de
Delphinoidea du Pliocène.
Premières baleines à fanons (mysticètes) [modifier]
On compte actuellement parmi les mysticètes 4 familles qui pourraient avoir des origines différentes
chez les cétothères :
Tous les mysticètes modernes sont de grands cétacés à fanons, et se nourrissent en filtrant, bien
que la manière exacte d'utiliser les fanons diffère entre les espèces (consommation par goulées chez les
balénoptéridés, par écumage chez les baleines franches, et par aspiration du fond pour les baleines grises).
Les premiers représentants de certains groupes modernes apparaissent au miocène moyen. Ceci peut avoir
été dû à des changements environnementaux et physiques de l'océan. Un changement à grande échelle des
courants et températures de l'océan peut avoir déclenché la radiation des mysticètes modernes, en éliminant
les formes archaïques. En général, on pense que les quatre familles modernes de mysticètes ont des
origines séparées parmi les cétothères : les Balaenopteridae (dont les baleines à bosse et les rorquals), les
Balaenidae (baleines franches), les Eschrichtiidae (baleines grises), les Neobalaenidae (baleines pygmées)
ont tous des caractères dérivés actuellement inconnus chez les cétothères.

Évolution du squelette [modifier]

Aujourd'hui, les membes postérieurs des cétacés sont internes, et réduits. Ils ne servent plus que
comme supports pour les muscles des organes génitaux. Alors que les cétacés primitifs comme le Pakicetus
avaient les orifices nasaux au bout du museau, ces orifices ont commencé à migrer vers le sommet du
crâne. Chez Rodhocetus, ils ont déjà substantiellement amorcé cette migration nasale. Les narines des
cétacés modernes ont atteint le sommet du crâne et se sont modifiés en évents, qui leur permettent
d'effleurer la surface, expirer, inspirer et replonger avec facilité. Les oreilles ont commencé à se déplacer
aussi vers l'intérieur, et déjà chez le Basilosaurus, l'oreille moyenne commençait à recevoir des vibrations de
la mâchoire inférieure. Les odontocètes modernes utilisent leur melon, un amas de graisse jouant le rôle de
lentille acoustique pour l'émission de sons destinés à l'écholocation. Et ils utilisent leur mâchoire inférieure
pour capter les échos de haute fréquence et les focaliser sur l'oreille.

Atavismes ancestraux [modifier]

Article détaillé : Embryologie.
Les cétacés développent toujours certaines caractéristiques ancestrales à l'état embryonnaire ou de
nouveau-nés. Les baleines à fanons développent des bourgeons de dents à l'état embryonnaire, qui
disparaîtront chez l'animal adulte. Les fœtus de certaines espèces portent des poils éphémères qui
disparaissent plus tard. Les embryons des cétacés ont les fosses nasales situées à l'extrémité du museau,
comme chez tout mammifère. Celles-ci migrent durant le développement embryonnaire vers l'arrière, pour se
stabiliser au-dessus du crâne et former l'évent chez le nouveau-né. Certains spécimens adultes disposent de
moignons de pattes arrière. Les embryons de nombreuses espèces en ont également[20]. Ce phénomène
est connu sous le nom d'atavisme.

Arbre phylogénétique [modifier]

Les arguments développés ci-dessus conduisent à une classification phylogénétique plus probable,
encore que comme nous l'avons vu, certains détails restent à clarifier.
On en suivra les développements dans l'article Classification phylogénétique des Cetartiodactyla, qui
est régulièrement mis à jour. Noter qu'en cliquant sur le lien [Dérouler] de l'onglet « Arbre développé », on
peut le développer dans tous ses détails, ce qui n'est pas envisageable dans le présent article.
La pression humaine [modifier]
Bon nombre d'espèces sont en déclin rapide et menacent de disparaître du fait de la chasse à la
baleine. On estime que 90% de l’effectif de grands cétacés a aujourd’hui disparu de la planète. Malgré le
moratoire de 1982, de nombreuses espèces sont toujours menacées d’extinction[21]. On notera, pour
l'échantillon restreint des espèces de cétacés, parmi les espèces disparues récemment, l'extinction,
constatée en 2006, du Dauphin de Chine

Notes et références [modifier]

• (en) Cet article est partiellement ou en totalité issu de l’article de Wikipédia en anglais
intitulé « Evolution of cetaceans » (voir la liste des auteurs)
1. ↑ (en) W.H. Flower, « On whales, present and past and their probable origin », dans Proc.
Zool. Soc., 1883, p. 466-513
2. ↑ (en) A. V. Yablokov, « Convergence or parallelism in the evolution of cetaceans », dans
Int. Geol., vol. 7, no 146, 1965, p. 1-1468
3. ↑ (en) A. Boyden et D. Gemeroy, « The relative position of the Cetacea among the orders
of Mammalia as indicated by precipitin tests. », dans Zoologica, vol. 35, 1950, p. 145-151 .
4. ↑ (en) J. G. M. Thewissen, E. M. Williams, L. J. Roe et S. T. Hussain, « Skeletons of
terrestrial cetaceans and the relationship of whales to artiodactyls », dans Nature, vol. 413, 2001,
p. 277–281 [ résumé [archive] (page consultée le 20 janvier 2010) ] .
 5. ↑ (en) Dan Graur et Desmond G. Higgins, « Molecular evidence for the inclusion of
cetaceans within the order Artiodactyla [archive] », juillet 1993, Mol. Biol. Evol., p. 357-364. Consulté
le 20 janvier 2010
6. ↑ a et b (en) Jean-Renaud Boisserie, « UC Berkeley, French Scientists Find Missing Link
Between The Whale And Its Closest Relative, The Hippo [archive] », février 2005, ScienceDaily.
Consulté le 20 janvier 2010
7. ↑ (en) C. Montgelard et F.M. Catzeflis et E. Douzery, « Phylogenetic Relationships of
Artiodactyls and Cetaceans as Deduced from the Comparison of Cytochrome b and 12s rRNA
Mitochondrial Sequences », dans Mol. Biol. Evol., vol. 14, no 5, 1997, p. 550-559
8. ↑ (en) Mitsuru Shimamura et et al., « Molecular evidence from retroposons that whales
form a clade within even-toed ungulates. », dans Nature, vol. 388, 14 août 1997, p. 666-670
[ résumé [archive] (page consultée le 20 janvier 2010) ]
9. ↑ (en) Jean-Renaud Boisserie et Fabrice Lihoreau et Michel Brunet, « The position of
Hippopotamidae within Cetartiodactyla », dans Proceedings of the National Academy of Sciences,
2005 [ texte intégral [archive] (page consultée le 20 janvier 2010) ]
10.↑ (en) P. D. Gingerich et D. E. Russell, « Pakicetus inachus, a new archaeocete
(Mammalia, Cetacea) from the early-middle Eocene Kuldana Formation of Kohat (Pakistan) », dans
Univ. Mich. Contr. Mus. Paleont., vol. 25, 1981, p. 235–246
11.↑ a et b (en) Northeastern Ohio Universities Colleges of Medicine and Pharmacy, « Whales
Descended From Tiny Deer-like Ancestors [archive] », décembre 2007, ScienceDaily. Consulté le 20
janvier 2010
 12.↑ (en) University Of Michigan, « New Fossils Suggest Whales And Hippos Are Close
Kin [archive] », septembre 2001, ScienceDaily. Consulté le 20 janvier 2010
13.↑ (en) Carl Zimmer, « The Loom : Whales: From So Humble A Beginning... [archive] »,
décembre 2007, ScienceBlogs. Consulté le 20 janvier 2010
14.↑ (en) Ian Sample, « Whales may be descended from a small deer-like animal [archive] »,
19 décembre 2007, Guardian Unlimited. Consulté le 20 janvier 2010
15.↑ (en) PZ Myers, « Pharyngula: Indohyus [archive] », 19 décembre 2007, ScienceBlogs.
Consulté le 20 janvier 2010
16.↑ a, b et c (en)Ambulocetidae [archive]
17.↑ (en) J.G.M. Thewissen (dir.), The Emergence of Whales, Plenum Press, 1998
18.↑ (en) Gingerich PD, ul-Haq M, von Koenigswald W, Sanders WJ, Smith BH, et al., « New
Protocetid Whale from the Middle Eocene of Pakistan: Birth on Land, Precocial Development, and
Sexual Dimorphism [archive] », février 2009, PLoS one. Consulté le 20 janvier 2010
19.↑ (en) Lori Marino, Richard C. Connor, R. Ewan Fordyce, Louis M. Herman, Patrick R.
Hof, Louis Lefebvre, David Lusseau, Brenda McCowan, Esther A. Nimchinsky, Adam A. Pack, Luke
Rendell, Joy S. Reidenberg, Diana Reiss, Mark D. Uhen, Estel Van der Gucht, Hal Whitehead,
« Cetaceans Have Complex Brains for Complex Cognition [archive] », mai 2007, PLoS Biology.
Consulté le 20 janvier 2010
20.↑ (en) Les FOSSILES et les ORGANES VESTIGIAUX [archive], rationalisme.org.
Consulté le 20 janvier 2010
21.↑ (en) Réseau Cétacés - Actions & Pétitions [archive], 2005, Réseau Cétacés. Consulté le
20 janvier 2010
 Voir aussi [modifier]
Articles connexes [modifier]
• Dauphins
• Baleines
• Cétacés
• Siréniens, Otariidés, Odobénidés, Phoques ; les autres espèces de mammifères marins.
• Archaeoceti
• Indohyus
• Histoire évolutive des mammifères
• Histoire évolutive des siréniens
• Histoire évolutive des hippopotamidés
• Évolution

Liens externes [modifier]

• Ancêtres et histoire des Cétacés, Terra Nova.. Consulté le 21 janvier 2010
• Par Jean-Luc Goudet, « Le premier ancêtre nageur des cétacés était-il un daim miniature ? »
sur Futura-sciences.. Consulté le 21 janvier 2010
• Par Christophe Olry, « Prédateur des océans : surprenante baleine à dents de 25 millions
d'années » sur Futura-sciences.. Consulté le 21 janvier 2010
• Éric Marchand, « LES ANCÊTRES DES CÉTACÉS ». Consulté le 21 janvier 2010
 • (en)Michel C. Milinkovitch et Olivier Lambert, « Cetacea » sur Tree of Life web project..
Consulté le 21 janvier 2010
• (en)Cetacea sur Animal Diversity Web (University of Michigan, Museum of Zoology).
(Consulté le = 21 janvier 2010).
• (en) Michel C. Milinkovitch et Axel Meyer et Jeffrey R. Powell, « Phylogeny of All Major
Groups of Cetaceans Based on DNA Sequences from Three Mitochondrial Genes », dans Mol. Biol.
Evol., Oxford Journals, Oxford, vol. 11, 1994, p. 939-948 [ texte intégral (page consultée le 21 janvier
2010) ] </ref>
• (en)CETACEA - Whales, Mikko's Phylogeny Archive.. Consulté le 21 janvier 2010
• (en) J. Geisler et A. Sanders, « Morphological evidence for the phylogeny of Cetacea »,
dans Journal of Mammalian Evolution, vol. 10, no 1 et 2, 2003, p. 23-129 .
• (en) Cetacean Paleobiology, University of Bristol. Consulté le 21 janvier 2010
• (en) How whales learned to swim, mai 2002, BBC News. Consulté le 21 janvier 2010
• (en) Stephen Jay Gould, « Hooking Leviathan by Its Past », 1997. Consulté le 21 janvier
2010
• (en) Thewissen Lab, « Digital Library of Dolphin Development », Northeastern Ohio
Universities Colleges of Medicine and Pharmacy. Consulté le 21 janvier 2010</ref>
• (en) P. D. Gingerich, « Research on the Origin and Early Evolution of Whales (Cetacea) ».
Consulté le 21 janvier 2010</ref>
• (en) Edward T. Babinski, « Evolution of Whales (adapté de National Geographic
Magazine : « The Evolution of Whales » », 2001 rév. 2006 Thewissen. Consulté le 21 janvier
2010</ref>
 • (en) Philip D. Gingerich et Donald E. Russell, « Pakicetus Inachus, a new archaeocete
(mammalia, cetacea) from the early-middle Eocene Kuldana formation of Kohat (Pakistan) », dans
Contributions from the museum of Paleontology, Université du Michigan, Ann Arbor, MI, USA,
vol. 25, no 11, décembre 1981, p. 235-246 [ [pdf]texte intégral (page consultée le 21 janvier
2010) ] </ref>
• (en) J. G. M. Thewissen et E. M. Williams, L. J. Roe & S. T. Hussain, « Skeletons of
terrestrial cetaceans and the relationship of whales to artiodactyls », dans Nature, vol. 413, 20
septembre 2001, p. 277-281 [ résumé, lien DOI (pages consultées le 21 janvier 2010) ] </ref>
• (en) Evolution of Whales segment from the Whales, 2008, Museum of New Zealand Te
Papa Tongarewa, p. Tohorā Exhibition Minisite. Consulté le 21 janvier 2010</ref>

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Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens
 Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des
Deutérostomiens dinosaures · Histoire évolutive des oiseaux

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%C3%A9s ».
Catégories : Cétacé | Histoire évolutive | [+]
 W000

Histoire évolutive des poissons cartilagineux

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Le groupe des poissons cartilagineux, ou Chondrichthyes, est très ancien. Il regroupe des espèces
assez différentes morphologiquement les unes des autres, comme les raies et requins, très variées en taille,
comme le sagre elfe qui mesure 18 centimètres à l'âge adulte et le requin baleine qui, lui, atteint 18 mètres
de long.
Sommaire
[masquer]
• 1 Évolution
• 1.1 Du Paléozoïque au Dévonien
• 1.2 La fin de l'âge d'or
• 1.3 Les Chondrichthyes modernes
• 2 Genèse de cette théorie
• 3 Notes et références
• 4 Voir aussi
• 4.1 Bibliographie
• 4.2 Lien externe

Évolution [modifier]
Article détaillé : Évolution.

Du Paléozoïque au Dévonien [modifier]

Les premiers fossiles découverts de cette classe sont des dents de petite taille datées de 430 ma.
Peu de fossiles sont découverts avant 354 Ma. Leur forme est assez semblable à celle des requins
d'aujourd'hui, on les suppose cependant moins évolués et rapides que ceux d'aujourd'hui. En fait durant
cette période, les placodermes semblent les espèces les plus abondantes. Ces derniers s'éteignent à la fin
du Dévonien[1] et les Chondrichthyes subissent une radiation évolutive, occupant abondamment toutes les
niches écologiques. Certaines espèces vont également grandir, atteignant trois mètres et devenant les
superprédateurs marins, place qu'occupe encore le Grand requin blanc. Certains groupes d'espèces comme
les Xénacanthes vont même conquérir les rivières.

La fin de l'âge d'or [modifier]

À la fin du Permien une extinction de masse se produit faisant disparaître 90 % des espèces marines
dont des chondrichtyens. On suppose que la destruction des environnements par une baisse du niveau des
mers est une raison de la baisse de la diversité de ces espèces. Les Xénacanthes survivent.
Une deuxième extinction à la fin du Trias opère à nouveau une sélection et élimine, par exemple, les
xénacanthes qui avaient vécu pendant 200 Ma. Au Mésozoïque les hybodontes se distinguent par leur
grande taille (au moins 3 mètres) et une mâchoire capable de broyer n'importe quelle carapace[2]. Au
Jurassique, les néosélaciens sont en compétition avec les hybodontes mais l'extinction du Crétacé, il y a 65
Ma, provoqua la fin des hybodontes ainsi que celle de beaucoup d'autres espèces, à commencer par les
dinosaures.

Les Chondrichthyes modernes [modifier]

C'est à la fin du Mésozoïque que les poissons osseux prennent leurs formes actuelles. Les
néosélaciens se diversifiant quelque peu. Par exemple les Torpediniformes apparaissent il y a 62 Ma. Les
Pristidae ou poissons scies apparaissent à leur tour il y a 54 Ma. Et même si le Lamniforme Carcharocles
megalodon, qui est le plus gros des requins ayant jamais existé, avec ses 13 mètres de long apparaît il y 23
Ma, et disparaît définitivement il y a deux Ma, les autres espèces de la classe évoluent peu. Cependant le
nombre des espèces au sein des différentes familles évolue beaucoup et depuis 5 Ma environ c'est la famille
des Carcharhiniformes qui compte le plus de requins. Elle a surclassé celle des Lamniformes qui ne
représente plus que 5% des espèces. Ainsi, très abondant sur les côtes au Crétacé, le Requin lutin ne vit
plus qu'entre 350 et 600 mètres de fond.

Genèse de cette théorie [modifier]

Les dents de requins fossiles ont longtemps été une énigme. C'est Fabio Colonna qui, le premier,
démontre de façon convaincante que les glossopètres sont des dents de requin[3], dans son traité De
glossopetris dissertatio publié en 1616[4]. Les fossiles de requin sont rares puisque les cartilages qui
composent leur squelette se conservent moins bien que les os calcifiés des Ostéichthyens, clade qui inclut
certains poissons et les vertébrés. Cependant les dents des Elasmobranchii dont les requins sont elles
abondantes, du fait que ceux-ci en perdent et en fabriquent tout au long de leur vie. Les dents des chimères
sont, elles, beaucoup plus rares. Certaines épines recouvrant le corps des Chondrichthyes, qui sont de la
même matière que les dents, sont aussi régulièrement découvertes, mais elles aussi plus rares. Ces
hypothèses sont faites à partir de datation des fossiles découverts.
Les squelettes les plus anciens furent découverts à la fin du XIXe siècle dans l'État de l'Ohio aux
États-Unis et sont du genre Cladoselache datant du Dévonien supérieur (360 Ma)[5].

Notes et références [modifier]

1. ↑ (Cuny, 2002, p. 63)
2. ↑ (Cuny, 2002, p. 150)
 3. ↑ Site internet du Centro dei Musei di Scienze Naturali, université de Naples [archive]
consulté le 11 août 2007
4. ↑ Francesco Abbona, Geologia, Dizionario Interdisciplinare di Scienza e Fede, Urbaniana
University Press - Città Nuova Editrice, Roma 2002, http://www.disf.org/Voci/4.asp [archive] article
en ligne consulté le 11 août 2007
5. ↑ (Cuny, 2002, p. 50)

Voir aussi [modifier]

Bibliographie [modifier]
• Gilles Cuny, Les requins sont-ils des fossiles vivants ?, EDP Sciences, coll. « Bulles de
sciences », 2002, (ISBN 2868835384)

Lien externe [modifier]

• Gilles Cuny, « La longue histoire des requins » sur Futura-Sciences, 05/05/2007, dossier

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes
 Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et
Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Mammifères Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire
 évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

• Portail du monde maritime

Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_
%C3%A9volutive_des_poissons_cartilagineux ».

Catégories : Histoire évolutive | Poisson cartilagineux | [+]

 W000

Evolution of fungi
From Wikipedia, the free encyclopedia

Jump to: navigation, search

The evolution of fungi has been going on since fungi diverged from other life around 1,500 million
years ago, (Wang et al., 1999)[1] with the glomaleans branching from the "higher fungi" at ~570 million years
ago, according to DNA analysis. (Schüßler et al., 2001; Tehler et al., 2000)[1] Fungi probably colonised the
land during the Cambrian, over 500 million years ago, (Taylor & Osborn, 1996)[1] but fossils only become
uncontroversial and common during the Devonian, 400 million years ago.[1]
[edit] Early fungi
A rich diversity of fungi is known from the lower Devonian Rhynie chert, an earlier record is absent.
Since fungi do not biomineralise, they do not readily enter the fossil record; there are only three claims of
early fungi. One from the Ordovician[2] has been dismissed on the grounds that it lacks any distinctly fungal
features, and is held by many to be contamination;[3] the position of a "probable" Proterozoic fungus is still
not established,[3] and it may represent a stem group fungus. There is also a case for a fungal affinity for the
enigmatic microfossil Ornatifilum. Since the fungi form a sister group to the animals, the two lineages must
have diverged before the first animal lineages, which are known from fossils as early as the Ediacaran.[4]

[edit] References
Fungi portal

1. ^ a b c d in Brundrett, M.C. (2002). "Coevolution of roots and mycorrhizas of land plants". New
Phytologist 154 (2): 275–304. doi:10.1046/j.1469-8137.2002.00397.x.
2. ^ Redecker, D.; Kodner, R.; Graham, L.E. (2000). "Glomalean Fungi from the Ordovician".
Science 289 (5486): 1920. doi:10.1126/science.289.5486.1920. PMID 10988069.
3. ^ a b Butterfield, N.J. (2005). "Probable Proterozoic fungi". Paleobiology 31 (1): 165–182.
doi:10.1666/0094-8373(2005)031.
4. ^ Miller, A.J. (2004). "A Revised Morphology of Cloudina with Ecological and Phylogenetic
Implications". http://ajm.pioneeringprojects.org/files/CloudinaPaper_Final.pdf. Retrieved 2007-04-24
 W000

Evolution of the horse

From Wikipedia, the free encyclopedia

Jump to: navigation, search

 This image shows a representative sequence but should not be construed to represent a "straight-
line" evolution of the horse. Reconstruction, left forefoot skeleton (third digit emphasized yellow) and
longitudinal section of molars of selected prehistoric horses
The evolution of the horse pertains to the phylogenetic ancestry of the modern horse from the fox-
sized, forest-dwelling Hyracotherium over geologic time scales. Paleozoologists have been able to piece
together a more complete picture of the modern horse's evolutionary lineage than that of any other animal.
The horse belongs to an order known as Perissodactyla, or "odd-toed ungulates", which all share
hoofed feet and an odd number of toes on each foot, as well as mobile upper lips and a similar tooth
structure. This means that horses share a common ancestry with tapirs and rhinoceroses. The perissodactyls
originally arose in the late Paleocene, less than 10 million years after the Cretaceous-Tertiary extinction
event. This group of animals appears to have been originally specialized for life in tropical forests, but
whereas tapirs and, to some extent, rhinoceroses, retained their jungle specializations, modern horses are
adapted to life on drier land in the much-harsher climatic conditions of the steppes. Other species of Equus
are adapted to a variety of intermediate conditions.
The early ancestors of the modern horse walked on several spread-out toes, an accommodation to
life spent walking on the soft, moist grounds of primeval forests. As grass species began to appear and
flourish, the equids' diets shifted from foliage to grasses, leading to larger and more durable teeth. At the
same time, as the steppes began to appear, the horse's predecessors needed to be capable of greater
speeds to outrun predators. This was attained through the lengthening of limbs and the lifting of some toes
from the ground in such a way that the weight of the body was gradually placed on one of the longest toes,
the third.
Contents
[hide]
• 1 History of research
• 2 Eocene and Oligocene: Early equids
• 2.1 Hyracotherium
• 2.2 Orohippus
• 2.3 Epihippus
• 2.4 Mesohippus
• 2.5 Miohippus
• 3 Miocene and Pliocene: True equines
• 3.1 Kalobatippus
• 3.2 Parahippus
• 3.3 Merychippus
• 3.4 Hipparion
• 3.5 Pliohippus
• 3.6 Dinohippus
• 3.7 Plesippus
• 4 Modern horses
• 4.1 Equus
• 4.2 Pleistocene Extinctions
• 4.3 Return to the Americas
[edit] History of research

Restoration of Eurohippus parvulus, Museum für Naturkunde Berlin

Horses were absent from the Americas until the Spanish brought domestic horses from Europe,
beginning in 1493, and escaped horses quickly established large wild herds. The early naturalist Buffon
suggested in the 1760s that this was an indication of inferiority of fauna in the New World, then later
reconsidered this idea.[1] William Clark's 1807 expedition to Big Bone Lick found "leg and foot bones of the
Horses" which were included with other fossils sent to Thomas Jefferson and evaluated by the anatomist
Caspar Wistar, but neither commented on the significance of this find.[2]
The first equid fossil was found in the gypsum quarries in Montmartre, Paris in the 1820s. The tooth
was sent to the Paris Conservatory, where it was identified by Georges Cuvier who identified it as a browsing
equine related to the tapir.[3]. His sketch of the entire animal matched later skeletons found at the site[4].
 During the Beagle survey expedition the young naturalist Charles Darwin had remarkable success
with fossil hunting in Patagonia. On 10 October 1833 at Santa Fe, Argentina, he was "filled with
astonishment" when he found a horse's tooth in the same stratum as fossil giant armadillos, and wondered if
it might have been washed down from a later layer, but concluded that this was "not very probable".[5] After
the expedition returned in 1836, the anatomist Richard Owen confirmed the tooth was from an extinct
species which he subsequently named Equus curvidens, and remarked that "This evidence of the former
existence of a genus, which, as regards South America, had become extinct, and has a second time been
introduced into that Continent, is not one of the least interesting fruits of Mr. Darwin's palæontological
discoveries."[2][6]
In 1848 a study On the fossil horses of America by Joseph Leidy systematically examined
Pleistocene horse fossils from various collections, including that of the Academy of Natural Sciences and
concluded at least two ancient horse species had existed in North America: Equus curvidens and another
which he named Equus americanus. A decade later, however, he found the latter name had already been
taken and renamed it Equus complicatus.[1] In the same year, he visited Europe and was introduced by
Owen to Darwin.[7]
The original sequence of species believed to have evolved into the horse was based on fossils
discovered in North America in the 1870s by paleontologist Othniel Charles Marsh. The sequence, from
Hyracotherium (popularly called Eohippus) to the modern horse (Equus), was popularized by Thomas Huxley
and became one of the most widely-known examples of a clear evolutionary progression. The horse's
evolutionary lineage became a common feature of biology textbooks, and the sequence of transitional fossils
was assembled by the American Museum of Natural History into an exhibit which emphasized the gradual,
"straight-line" evolution of the horse.
 Since then, as the number of equid fossils has increased, the actual evolutionary progression from
Hyracotherium to Equus has been discovered to be much more complex and multi-branched than was
initially supposed. The straight, direct progression from the former to the latter has been replaced by a more
elaborate model with numerous branches in different directions, of which the modern horse is only one of
many. It was first recognized by George Gaylord Simpson in 1951[8] that the modern horse was not the
"goal" of the entire lineage of equids,[9] it is simply the only genus of the many horse lineages that has
survived.
Detailed fossil information on the rate and distribution of new equid species has also revealed the
progression between species was not as smooth and consistent as was once believed. Although some
transitions, such as that of Dinohippus to Equus, were indeed gradual progressions, a number of others,
such as that of Epihippus to Mesohippus, were relatively abrupt and sudden in geologic time, taking place
over only a few million years. Both anagenesis (gradual change in an entire population's gene frequency)
and cladogenesis (a population "splitting" into two distinct evolutionary branches) occurred, and many
species coexisted with "ancestor" species at various times. The change in equids' traits was also not always
a "straight line" from Hyracotherium to Equus: some traits reversed themselves at various points in the
evolution of new equid species, such as size and the presence of facial fossae, and it is only in retrospect
that certain evolutionary trends can be recognized.[10]

[edit] Eocene and Oligocene: Early equids

[edit] Hyracotherium
The earliest animal to bear recognizably horse-like anatomy was the Hyracotherium ("hyrax-like
beast"). Its scientific name is derived from initial confusion over early partial fossils' relationship with living
species: Richard Owen likened early Hyracotherium fossils "to a hare in one passage and to something
between a hog and a hyrax in another".[11] A later name for the Hyracotherium, "eohippus" ("dawn horse"), is
also popular, though the earlier name takes precedence due to scientific naming conventions.[12][13]
Hyracotherium lived in the Ypresian (early Eocene), about 52 mya (million years ago). It was an
animal approximately the size of a fox (250–450 mm in height), with a relatively short head and neck and a
springy, arched back. It had 44 low-crowned teeth, in the typical arrangement of an omnivorous, browsing
mammal: 3 incisors, 1 canine, 4 premolars, and 3 molars on each side of the jaw. Its molars were uneven,
dull, and bumpy, and used primarily for grinding foliage. The cusps of the molars were slightly connected in
low crests. The Hyracotherium browsed on soft foliage and fruit, probably scampering between thickets in the
mode of a modern muntjac; the Hyracotherium had a small brain, and possessed especially small frontal
lobes.[10]

Hyracotherium, with left forefoot (third metacarpal colored) and tooth (a enamel; b dentin; c cement)
detailed.
 Its limbs were decently long relative to its body, already showing the beginnings of adaptations for
running. However, all of the major leg bones were unfused, leaving the legs flexible and rotatable. Its wrist
and hock joints were low to the ground. The forelimbs had developed five toes, out of which only four were
equipped with a small proto-hoof; the large fifth "toe-thumb" was off the ground. The hind limbs had three out
of the five toes equipped with small hooves, while the vestigial first and fifth toes did not touch the ground. Its
feet were padded, much like a dog's, but with the small hooves on each toe in place of claws.
For a span of about 20 million years, the Hyracotherium thrived with few significant evolutionary
changes.[10] The most significant change was in the teeth, which began to adapt to the changing diet of
Hyracotheria, as these early Equidae shifted from a mixed diet of fruits and foliage to one focused
increasingly on browsing foods. During the Eocene, a Hyracotherium species (most likely Hyracotherium
vassacciense) branched out into various new types of Equidae. Thousands of complete, fossilized skeletons
of these animals have been found in the Eocene layers of North American strata, mainly in the Wind River
basin in Wyoming. Similar fossils have also been discovered in Europe, such as Propalaeotherium (which is
not considered ancestral to the modern horse).[14]

[edit] Orohippus
Approximately 50 million years ago, in the early-to-middle Eocene, Hyracotherium smoothly
transitioned into Orohippus over a gradual series of changes.[14] Although its name means "mountain
horse", Orohippus was not a true horse and did not live in the mountains. It resembled Hyracotherium in size,
but had a slimmer body, an elongated head, slimmer forelimbs, and longer hind legs, all of which are
characteristics of a good jumper. Although Orohippus was still pad-footed, the vestigial outer toes of
Hyracotherium were not present in the Orohippus; there were four toes on each forelimb, and three on each
hind leg.
 The most dramatic change between Hyracotherium and Orohippus was in the teeth: the first of the
premolar teeth were dwarfed, the last premolar shifted in shape and function into a molar, and the crests on
the teeth became more pronounced. Both of these factors gave the teeth of Orohippus greater grinding
ability, suggesting that Orohippus ate tougher plant material.

[edit] Epihippus
In the mid-Eocene, about 47 million years ago, Epihippus, a genus which continued the evolutionary
trend of increasingly efficient grinding teeth, evolved from Orohippus. Epihippus had five grinding, low-
crowned cheek teeth with well-formed crests. A late species of Epihippus, sometimes referred to as
Duchesnehippus intermedius, had teeth similar to Oligocene equids, although slightly less developed.
Whether Duchesnehippus was a subgenus of Epihippus or a distinct genus is disputed.

[edit] Mesohippus
In the late Eocene and the early stages of the Oligocene epoch (32–24 mya), the climate of North
America became drier, and the earliest grasses began to evolve. The forests were yielding to flatlands,
[citation needed] home to grasses and various kinds of brush. In a few areas these plains were covered in
sand,[citation needed] creating the type of environment resembling the present-day prairies.
In response to the changing environment, the then-living species of Equidae also began to change.
In the late Eocene, they began developing tougher teeth and becoming slightly larger and leggier, allowing
for faster running speeds in open areas, and thus for evading predators in non-wooded areas. About 40 mya,
Mesohippus ("middle horse") suddenly developed in response to strong new selective pressures to adapt,
beginning with the species Mesohippus celer and soon followed by Mesohippus westoni.
 In the early Oligocene, Mesohippus was one of the more widespread mammals in North America. It
walked on three toes on each of its front and hind feet (the first and fifth toes remained, but were small and
not used in walking). The third toe was stronger than the outer ones, and thus more weighted; the fourth front
toe was diminished to a vestigial nub. Judging by its longer and slimmer limbs, Mesohippus was an agile
animal.
Mesohippus was slightly larger than Epihippus, about 610 mm (24") at the shoulder. Its back was
less arched, and its face, snout, and neck were somewhat longer. It had significantly larger cerebral
hemispheres, and had a small, shallow depression on its skull called a fossa, which in modern horses is quite
detailed. The fossa serves as a useful marker for identifying an equine fossil's species. Mesohippus had six
grinding "cheek teeth", with a single premolar in front—a trait all descendant Equidae would retain.
Mesohippus also had the sharp tooth crests of Epihippus, improving its ability to grind down tough
vegetation.

[edit] Miohippus
Around 36 million years ago, soon after the development of Mesohippus, Miohippus ("lesser horse")
emerged, the earliest species being Miohippus assiniboiensis. Like Mesohippus, Miohippus's evolution was
relatively abrupt, though a few transitional fossils linking the two genera have been found. It was once
believed that Mesohippus had anagenetically evolved into Miohippus by a gradual series of progressions, but
new evidence has shown that Miohippus's evolution was cladogenetic: a Miohippus population split off from
the main Mesohippus genus, coexisted with Mesohippus for around 4 million years, and then over time came
to replace Mesohippus.[15]
 Miohippus was significantly larger than its predecessors, and its ankle joints had subtly changed. Its
facial fossa was larger and deeper, and it also began to show a variable extra crest in its upper cheek teeth,
a trait that became a characteristic feature of equine teeth.
Miohippus ushered in a major new period of diversification in Equidae.[13] While Mesohippus died
out in the mid-Oligocene, Miohippus continued to thrive, and in the early Miocene (24–5.3 mya), it began to
rapidly diversify and speciate. It branched out into two major groups, one of which adjusted to the life in
forests once again, while the other remained suited to life on the prairies.[ citation needed]

[edit] Miocene and Pliocene: True equines

[edit] Kalobatippus
 Fossil Megahippus mckennai
The forest-suited form was Kalobatippus (or Miohippus intermedius, depending on whether it was a
new genus or species), whose second and fourth front toes were long, well-suited travel on the soft forest
floors. Kalobatippus probably gave rise to Anchitherium, which travelled to Asia via the Bering Strait land
bridge, and from there to Europe.[16] In both North America and Eurasia, larger-bodied genera evolved from
Anchitherium; Sinohippus in Eurasia and Hypohippus and Megahippus in North America.[17] Hypohippus
became extinct by the late Miocene.[18]

[edit] Parahippus
The Miohippus population that remained on the steppes is believed to be ancestral to Parahippus, a
North American animal about the size of a small pony, with a prolonged skull and a facial structure
resembling the horses of today. Its third toe was stronger and larger, and carried the main weight of the body.
Its four premolars resembled the molar teeth and the first were small and almost nonexistent. The incisive
teeth of Parahippus, like those of its predecessors, had a crown as humans do; however, the top incisors had
a trace of a shallow crease marking the beginning of the core/cup.
[edit] Merychippus

Merychippus, an effective grazer and runner.

In the middle of the Miocene epoch, the grazer Merychippus flourished. Merychippus had wider
molars than its predecessors, which are believed to have been used for crunching the hard grasses of the
steppes. The hind legs, which were relatively short, had side toes equipped with small hooves, but they
probably only touched the ground when running.[13] Merychippus radiated into at least 19 additional
grassland species.
[edit] Hipparion

Protohippus simus
Three lineages within Equidae are believed to be descended from the numerous varieties of
Merychippus: Hipparion, Protohippus and Pliohippus. The most different from Merychippus was Hipparion.
The main difference was in the structure of tooth enamel: in comparison with other Equidae, the inside, or
tongue side, had a completely isolated parapet. A complete and well-preserved skeleton of the North
American Hipparion shows an animal the size of a small pony. They were very slim, rather like antelopes,
and were adapted to life on dry prairies. On its slim legs, Hipparion had three toes equipped with small
hooves, but the side toes did not touch the ground.
In North America, Hipparion and its relatives (Cormohipparion, Nannippus, Neohipparion, and
Pseudhipparion), proliferated into many kinds of equids, at least one of which managed to migrate to Asia
and Europe during the Miocene epoch.[19] (European Hipparion differs from American Hipparion in its
smaller body size – the best-known discovery of these fossils was near Athens.)

[edit] Pliohippus

Pliohippus pernix
Pliohippus arose from Callippus in the middle Miocene, around 12 mya. It was very similar in
appearance to Equus, though it had two long extra toes on both sides of the hoof, externally barely visible as
callused stubs. The long and slim limbs of Pliohippus reveal a quick-footed steppe animal.
Until recently, Pliohippus was believed to be the ancestor of present-day horses because of its many
anatomical similarities. However, though Pliohippus was clearly a close relative of Equus, its skull had deep
facial fossae, whereas Equus had no fossae at all. Additionally, its teeth were strongly curved, unlike the very
straight teeth of modern horses. Consequently, it is unlikely to be the ancestor of the modern horse; instead,
it is a likely candidate for the ancestor of Astrohippus.[20]

[edit] Dinohippus
Dinohippus was the most common species of Equidae in North America during the late Pliocene. It
was originally thought that Dinohippus was monodactyl, but a 1981 fossil find in Nebraska shows that some
were tridactyl.

[edit] Plesippus

Mounted skeleton of Hagerman Horse (Equus simplicidens)

Plesippus is often considered an intermediate stage between Dinohippus and the extant genus,
Equus.
 The famous fossils found near Hagerman, Idaho were originally thought to be a part of the genus
Plesippus. Hagerman Fossil Beds (Idaho) is a Pliocene site, dating to about 3.5 mya. The fossilized remains
were originally called Plesippus shoshonensis, but further study by paleontologists determined that fossils
represented the oldest remains of the genus Equus.[21] Their estimated average weight was 425 kg, roughly
the size of an Arabian horse.
At the end of the Pliocene, the climate in North America began to cool significantly and most of the
animals were forced to move south. One population of Plesippus moved across the Bering land bridge into
Eurasia around 2.5 Ma ago.[22]

[edit] Modern horses

[edit] Equus
 Skull of a giant extinct horse of the Equus genus, E. eisenmannae
The genus Equus, which includes all extant equines, is believed to have evolved from Dinohippus,
via the intermediate form Plesippus. One of the oldest species is Equus simplicidens, described as zebra-like
with a donkey-shaped head. The oldest material to date is ~3.5 million years old from Idaho, USA. The genus
appears to have spread quickly into the Old World, with the similarly aged Equus livenzovensis documented
from western Europe and Russia.[23]
Molecular phylogenies indicate that the most recent common ancestor of all modern equids
(members of the genus Equus) lived ~5.6 (3.9-7.8) mya. The oldest divergencies are the Asian hemiones
(subgenus E. (Asinus), including the Kulan, Onager, and Kiang), followed by the African zebras (subgenera
E. (Dolichohippus), and E. (Hippotigris)). All other modern forms including the domesticated horse (and many
fossil Pliocene and Pleistocene forms) belong to the subgenus E. (Equus) which diverged ~4.8 (3.2-6.5)
million years ago.[24]
Pleistocene horse fossils have been assigned to a multitude of species, with over 50 species of
equines described from the Pleistocene of North America alone, although the taxonomic validity of most of
these has been called into question.[25] Recent genetic work on fossils has found evidence for only three
genetically divergent equid lineages in Pleistocene North and South America.[24] These results suggest that
all North American fossils of caballine-type horses (which also include the domesticated horse and
Przewalski's Horse of Europe and Asia), as well as South American fossils traditionally placed in the
subgenus E. (Amerhippus)[26] belong to the same species: E. ferus. Remains attributed to a variety of
species and lumped as New World stilt-legged horses (including E. francisci, E. tau, E. quinni and potentially
N. American Pleistocene fossils previously attributed to E. cf. hemiones, and E. (Asinus) cf. kiang) likely all
belong to a second species endemic to N. America, which despite a superficial resemblance to species in the
subgenus E. (Asinus) (and hence occasionally referred to as North American Ass) is closely related to E.
ferus.[24] Surprisingly, the third species, endemic to S. America, and traditionally referred to as Hippidion,
originally believed to be descended from Pliohippus, was shown to be a third species in the genus Equus,
closely related to the New World stilt-legged horse.[24] The temporal and regional variation in body size and
morphological features within each lineage indicates extraordinary intraspecific plasticity. Such environment-
driven adaptative changes would explain why the taxonomic diversity of Pleistocene equids has been
overestimated on morphoanatomical grounds.[26]
According to these results, it appears that the genus Equus evolved from a Dinohippus-like ancestor
~4-7 mya. It rapidly spread into the Old World and there diversified into the various species of asses and
zebras. A North American lineage of the subgenus E. (Equus) evolved into the New World stilt-legged horse
(NWSLH). Subsequently, populations of this species entered South America as part of the Great American
Interchange shortly after the formation of the Isthmus of Panama and evolved into the form currently referred
to as "Hippidion" ~2.5 million years ago. "Hippidion" is thus unrelated to the morphologically similar
Pliohippus, which presumably went extinct during the Miocene. Both the NWSLH and "Hippidium" show
adaptations to dry, barren ground, whereas the shortened legs of " Hippidion" may have been a response to
sloped terrain.[26] In contrast, the geographic origin of the closely related modern E. ferus is not resolved.
However, genetic results on extant[disambiguation needed] and fossil material of Pleistocene age indicate
two clades, potentially subspecies, one of which had a holarctic distribution spanning from Europe through
Asia and across North America and would become the founding stock of the modern domesticated horse.[27]
[28] The other population appears to have been restricted to N. America. One or more N. American
populations of E. ferus entered S. America ~1.0-1.5 million years ago, leading to the forms currently known
as "E. (Amerhippus)", which represent an extinct geographic variant or race of E. ferus, however.
[edit] Pleistocene Extinctions
In Western Canada, there is clear evidence of horses until 12,000 years ago.[29] All Equidae in North
America ultimately became extinct approximately 11,000 years ago. The causes of this extinction
(simultaneous with the extinctions of a variety of other American megafauna) have been a matter of debate.
Given the suddenness of the event and the fact that these mammals had been flourishing for millions of
years previously, something quite unusual must have happened. There are two main hypotheses. The first
attributes extinction to Climate change. Beginning approximately 12,500 years ago, the grasses
characteristic of a steppe ecosystem gave way to shrub tundra, which was covered with unpalatable plants.
[30] Another hypothesis suggests extinction was linked to arrival of and over-hunting by humans. Extinctions
were roughly simultaneous with the end of the most recent glacial advance and the appearance of the big-
game-hunting Clovis culture.[31][32] Findings published in 2006 showed that it was not yet possible to claim
with confidence that the extinction of horses in Alaska preceded the arrival of humans.[33]
Horse fossils disappear from the fossil record from about 10,000 years ago,[ dubious – discuss] but
begin occurring frequently again in archaeological sites in Kazakhstan and the southern Ukraine from 6,000
years ago.[27] From then on, it is probable that domesticated horses as well as the knowledge of capturing,
taming, and rearing wild horses spread relatively quickly, with wild mares from several wild populations being
incorporated en route.[28]

[edit] Return to the Americas

Horses only returned to the Americas with Christopher Columbus in 1493. These were Iberian horses
first brought to Hispaniola and later to Panama, Mexico, Brazil, Peru, Argentina, and, in 1538, Florida.[34]
The first horses to return to the main continent were 16 specifically identified horses brought by Hernan
Cortes. Subsequent explorers, such as Coronado and De Soto brought ever-larger numbers, some from
Spain and other from breeding establishments set up by the Spanish in the Caribbean. Later, as Spanish
missions were founded on the mainland, horses would eventually be lost or stolen, and proliferated into large
herds of feral horses that became known as mustangs.[citation needed]
The indigenous peoples of the Americas did not have a specific word for horses, and came to refer to
them in various languages as a type of dog or deer (in one case, "elk-dog").[ citation needed]

[edit] Details
[edit] Toes
The ancestors of the horse came to walk only on the end of the third toe and both side toes. Skeletal
remnants show obvious wear on the back of both sides of metacarpal and metatarsal bones, commonly
called the “splint bones”. They are the remnants of the second and the fourth toe. Modern horses retain the
splint bones; it is often believed that they are a useless attachment, but they in fact play an important role in
supporting the carpal joints (front knee) and even the tarsal joints (hock).

[edit] Teeth
Throughout the phylogenetic development, the teeth of the horse underwent significant changes. The
type of the original omnivorous teeth with short, "bumpy" molars, with which the prime members of the
evolutionary line distinguished themselves, gradually changed into the teeth common to herbivorous
mammals. They became long (as much as 100 mm), roughly cubical molars equipped with a flat grinding
surface. In conjunction with the teeth, during the horse’s evolution the elongation of the facial part of the skull
is apparent, and can also be observed in the backward set eyeholes. In addition, the relatively short neck of
the equine ancestors became longer with equal elongation of the legs. Finally, the size of the body grew as
well.

[edit] See also

• Evidence of common descent

[edit] Further reading

• MacFadden, Bruce J (2003 digitization of 1999 reprint). Fossil Horses: Systematics,
Paleobiology, and Evolution of the Family Equidae . Cambridge & New York: Cambridge University
Press. ISBN 0-521-47708-5. http://books.google.com/?
id=K1upTamSEW0C&printsec=frontcover&dq=%22fossil+horses%22&cd=1#v=onepage&q.
Retrieved 6 June 2010

[edit] References
1. ^ a b Academy of Natural Sciences - Joseph Leidy - American Horses
2. ^ a b Academy of Natural Sciences - Thomas Jefferson Fossil Collection - Ancient Horse
Fossils
3. ^ Horse breeding and management, James Warren Evans 1992
4. ^ Knell, Simon J.; Suzanne Macleod; Sheila E. R. Watson; Museum revolutions: how
museums and change and are changed Routledge, 2007, 385 pages ISBN 0-415-44467-5,
9780415444675
 5. ^ 'Filled with astonishment': an introduction to the St. Fe Notebook,
Barlow, Nora (ed. 1945) Charles Darwin and the voyage of the Beagle. London: Pilot Press, p. 210
6. ^ Darwin, C. R. (ed. 1840). Fossil Mammalia Part 1 No. 4 of The zoology of the voyage of
H.M.S. Beagle. By Richard Owen. London: Smith Elder and Co. p. 108–109
7. ^ Academy of Natural Sciences - Joseph Leidy - Leidy and Darwin
8. ^ Simpson, George Gaylord (1951): Horses. Oxford University Press; New Impression
edition. ISBN 0-19-500104-4 (1971 reprint)
9. ^ The notion of a goal would contradict modern evolutionary synthesis
10.^ a b c Hunt, Kathleen (1995). Horse Evolution. TalkOrigins Archive.
http://www.talkorigins.org/faqs/horses/horse_evol.html. Retrieved 6 June 2010 See also
downloadable pdf version
11.^ Gould, Stephen Jay (1991). "The Case of the Creeping Fox Terrier Clone" Bully for
Brontosaurus: Reflections in Natural History (pp. 155–167). New York: W.W. Norton & Co.
12.^ Gould, Stephen Jay, op. cit., "Bully for Brontosaurus"
13.^ a b c Fossil Horses In Cyberspace. Florida Museum of Natural History and the National
Science Foundation.
14.^ a b MacFadden, B. J. (1976). "Cladistic analysis of primitive equids with notes on other
perissodactyls." Syst. Zool. 25(1):1-14.
15.^ Prothero, D.R. and Shubin, N. (1989). "The evolution of Oligocene horses." The Evolution
of Perissodactyls (pp. 142–175). New York: Clarendon Press.
16.^ MacFadden, B.J. (2001). "Three-toed browsing horse Anchitherium clarencei from the early
Miocene (Hemingfordian) Thomas Farm, Florida". Bulletin of the Florida Museum of Natural History
43 (3): 79–109.
 17.^ Salesa, M.J., Sánchez, I.M., and Morales, J. (2004). "Presence of the Asian horse
Sinohippus in the Miocene of Europe". Acta Palaeontologica Polonica 49 (2): 189–196.
18.^ Waring, George H (2003). Horse Behavior (2nd ed.). New York: Noyes Publications/William
Andrew Publishing. p. 9. ISBN 0-8155-1484-0. http://books.google.com/?
id=hvy1TRsdtxcC&printsec=frontcover&dq=Horse+behavior&q. Retrieved 6 June 2010
19.^ MacFadden, B.J. (1984). "Systematics and phylogeny of Hipparion, Neohipparion,
Nannippus, and Cormohipparion (Mammalia, Equidae) from the Miocene and Pliocene of the New
World". Bulletin of the American Museum of Natural History 179 (1): 1–195.
http://hdl.handle.net/2246/997.
20.^ MacFadden, B. J. (1984). "Astrohippus and Dinohippus". J. Vert. Paleon. 4(2):273-283.
21.^ equus
22.^ Jens Lorenz Franzen: Die Urpferde der Morgenröte. Elsevier, Spektrum Akademischer
Verlag, München 2007, ISBN 3-8274-1680-9
23.^ Azzaroli, A. (1992). "Ascent and decline of monodactyl equids: a case for prehistoric
overkill". Ann. Zool. Finnici 28: 151–163. http://www.sekj.org/PDF/anzf28/anz28-151-163.pdf.
24.^ a b c d Weinstock, J.; et al. (2005). "Evolution, systematics, and phylogeography of
Pleistocene horses in the New World: a molecular perspective". PLoS Biology 3 (8): e241.
doi:10.1371/journal.pbio.0030241. PMID 15974804. PMC 1159165.
http://biology.plosjournals.org/perlserv/?request=get-
document&doi=10.1371%2Fjournal.pbio.0030241&ct=1. Retrieved 2008-12-19.
25.^ Azzaroli, A. (1998). "The genus Equus in North America". PalaeontographItal 85: 1–60.
26.^ a b c Orlando, L.; et al. (2008). "Ancient DNA Clarifies the Evolutionary History of American
Late Pleistocene Equids". Journal of Molecular Evolution 66 (5): 533–538. doi:10.1007/s00239-008-
9100-x. PMID 18398561.
 27.^ a b Vila, C.; et al. (2001). "Widespread Origins of Domestic Horse Lineages" (PDF).
Science 291. http://www.uky.edu/Ag/Horsemap/Maps/VILA.PDF. Retrieved 2008-12-19.
28.^ a b Jansen, T.; et al. (July 2002). "Mitochondrial DNA and the origins of the domestic
horse". Proceedings of the National Academy of Sciences 99 (16): 10905–10910.
doi:10.1073/pnas.152330099. PMID 12130666. PMC 125071.
http://www.pnas.org/content/99/16/10905.full. Retrieved 2008-12-19.
29.^ Singer, Ben (May 2005). A brief history of the horse in America. Canadian Geographic
Magazine. http://www.canadiangeographic.ca/Magazine/ma05/indepth/#cnd. Retrieved 16 October
2009.
30.^ LeQuire, Elise (2004-01-04). ""No Grass, No Horse"". The Horse, online edition.
http://www.thehorse.com/ViewArticle.aspx?ID=4849. Retrieved 2009-06-08.
31.^ "Ice Age Horses May Have Been Killed Off by Humans" National Geographic News, May 1,
2006.
32.^ Buck, Caitlin E.; Bard, Edouard (2007). "A calendar chronology for Pleistocene mammoth
and horse extinction in North America based on Bayesian radiocarbon calibration". Quaternary
Science Reviews 26 (17-18): 2031. doi:10.1016/j.quascirev.2007.06.013.
33.^ Solow, Andrew; Roberts, David; Robbirt, Karen (May 9, 2006). Haynes, C. Vance. ed. "On
the Pleistocene extinctions of Alaskan mammoths and horses". Proceedings of the National
Academy of Sciences of the United States of America (Proceedings of the National Academy of
Sciences of the United States of America) 103 (19): 7351–3. doi:10.1073/pnas.0509480103.
PMID 16651534. PMC 1464344. http://www.pnas.org/content/103/19/7351.full.
34.^ Luís, Cristina; et al. (2006). "Iberian Origins of New World Horse Breeds". Journal of
Heredity 97 (2): 107–113. doi:10.1093/jhered/esj020. PMID 16489143.
http://jhered.oxfordjournals.org/cgi/content/full/97/2/107.
 [edit] External links
• Horse Evolution Over 55 Million Years. Tufts University. January 10, 1998.
http://chem.tufts.edu/science/evolution/HorseEvolution.htm. Retrieved July 11, 2007. : An excerpted
collection of images of horse fossils

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Categories: Cenozoic | Evolution by taxon | Horse history and evolution

 W000

Horse Evolution
by Kathleen Hunt
Kathleen Hunt
Our apologies, but you must have JavaScript enabled to view author contact information.
Copyright ©1995-2003
[Article last updated: January 4, 1995]

his is a companion file for the Transitional Fossils FAQ and is part of the Fossil Horses FAQs.
In this post I will try to describe the modern view of evolution within the horse family. I apologize in advance
for the length; I didn't want to cut it down any more than this, because horse evolution has been
oversimplified too many times already. I wanted people to see some of the detail and complexity of the fossil
record of a fairly well known vertebrate group. (In fact, even at this length, this post is still only a summary!)
People who are in a hurry may just want to read the intro and summary and look at the tree.

Outline
1. Historical background -- why fossil horses are famous
2. Timescale and horse family tree
3. Small equids of the Eocene
4. Medium-sized browsing equids, late Eocene and Oligocene
5. The Miohippus radiation of browsing equids (24 My)
6. Horses move onto the plains: spring-foot & high-crowned teeth (18 My)
7. The merychippine radiation of the late Miocene (15 My)
8. One-toed grazing horses of the Pliocene & Pleistocene
9. Modern equines
10.Summary
11.References

I. Historical Background
In the 1870's, the paleontologist O.C. Marsh published a description of newly discovered horse
fossils from North America. At the time, very few transitional fossils were known, apart from Archeopteryx.
The sequence of horse fossils that Marsh described (and that T.H. Huxley popularized) was a striking
example of evolution taking place in a single lineage. Here, one could see the fossil species "Eohippus"
transformed into an almost totally different-looking (and very familiar) descendent, Equus, through a series of
clear intermediates. Biologists and interested laypeople were justifiably excited. Some years later, the
American Museum of Natural History assembled a famous exhibit of these fossil horses, designed to show
gradual evolution from "Eohippus" (now called Hyracotherium) to modern Equus. Such exhibits focussed
attention on the horse family not only as evidence for evolution per se, but also specifically as a model of
gradual, straight-line evolution, with Equus being the "goal" of equine evolution. This story of the horse family
was soon included in all biology textbooks.
As new fossils were discovered, though, it became clear that the old model of horse evolution was a
serious oversimplification. The ancestors of the modern horse were roughly what that series showed, and
were clear evidence that evolution had occurred. But it was misleading to portray horse evolution in that
smooth straight line, for two reasons:
1. First, horse evolution didn't proceed in a straight line. We now know of many other branches
of horse evolution. Our familiar Equus is merely one twig on a once-flourishing bush of equine
species. We only have the illusion of straight-line evolution because Equus is the only twig that
survived. (See Gould's essay "Life's Little Joke" in Bully for Brontosaurus for more on this topic.)
2. Second, horse evolution was not smooth and gradual. Different traits evolved at different
rates, didn't always evolve together, and occasionally reversed "direction". Also, horse species did
not always come into being by gradual transformation ("anagenesis") of their ancestors; instead,
sometimes new species "split off" from ancestors ("cladogenesis") and then co-existed with those
ancestors for some time. Some species arose gradually, others suddenly.
Overall, the horse family demonstrates the diversity of evolutionary mechanisms, and it would be
misleading -- and would be a real pity -- to reduce it to an oversimplified straight-line diagram.
With this in mind, I'll take you through a tour of the major genera of the horse family, Equidae.
CAUTION: I will place emphasis on those genera that led to the modern Equus. Do not be misled into
thinking that Equus was the target of evolution! Bear in mind that there are other major branches of the horse
tree that I will mention only in passing. (See the horse tree for a lovely ASCII depiction.)
Small preface: All equids (members of the family Equidae) are perissodactyls -- members of the
order of hoofed animals that bear their weight on the central 3rd toe. (Other perissodactyls are tapirs and
rhinos, and possibly hyraxes.) The most modern equids (descendents of Parahippus) are called "equines".
Strictly speaking, only the very modern genus Equus contains "horses", but I will call all equids "horses"
rather indiscriminately.
Most horse species, including all the ancestors of Equus, arose in North America.

II. Timescale and Horse Family Tree

Recent 10,000 years ago to present

Pleistocene 2.5-0.01 My (million years ago)

Pliocene 5.3-2.5 My

Miocene 24-5.3 My

Oligocene 34-24 My

Eocene 54-34 My
 And here's the tree...note that the timescale is a bit weird (e.g. the Oligocene is compressed almost
to nothing) to keep it from being too long. All the names on the tree are genus names, so recall that each
genus encompasses a cluster of closely related species.
The is a brief description of the tree for those who are visually impaired. Hyracotherium is shown giving rise to
three lineages. Two lineages quickly go extinct. The third branches many times. There are many branches alive
during most times until two million years ago when only the various species of Equus remain. The tree itself is
unreadable to those who are visually impaired so skip the tree graphic.
2My Old & New World Equus
\ | /
\ | /
4My Hippidion Equus Stylohipparion
| | Neohipparion Hipparion Cormohipparion
| | Astrohippus | | |
| | Pliohippus ---------------------------
12My Dinohippus Calippus \ | /
| | Pseudhipparion \ | /
| | | |
------------------------------------------- Sinohippus
15My \ | / |
\ | / Megahippus |
17My Merychippus | |
 | Anchitherium Hypohippus
| | |
23My Parahippus Anchitherium Archeohippus
| | |
(Kalobatippus?)-----------------------------------------
25My \ | /
\ | /
|
35My |
Miohippus Mesohippus
| |
40My Mesohippus
|
|
|
45My Paleotherium |
| Epihippus
| |
Propalaeotherium | Haplohippus
| | |
50My Pachynolophus | Orohippus
| | |
| | |
------------------------------
 \ | /
\ | /
55My Hyracotherium

III. Small Eocene Horses

The first equid was Hyracotherium, a small forest animal of the early Eocene. This little animal (10-
20" at the shoulder) looked nothing at all like a horse. It had a "doggish" look with an arched back, short neck,
short snout, short legs, and long tail. It browsed on fruit and fairly soft foliage, and probably scampered from
thicket to thicket like a modern muntjac deer, only stupider, slower, and not as agile. This famous little equid
was once known by the lovely name "Eohippus", meaning "dawn horse". Some Hyracotherium traits to
notice:
• Legs were flexible and rotatable with all major bones present and unfused.
• 4 toes on each front foot, 3 on hind feet. Vestiges of 1st (& 2nd, behind) toes still present.
Hyracotherium walked on pads; its feet were like a dog's padded feet, except with small "hoofies" on
each toe instead of claws.
• Small brain with especially small frontal lobes.
• Low-crowned teeth with 3 incisors, 1 canine, 4 distinct premolars and 3 "grinding" molars in
each side of each jaw (this is the "primitive mammalian formula" of teeth). The cusps of the molars
were slightly connected in low crests. Typical teeth of an omnivorous browser.
At this point in the early Eocene, equids were not yet very different from the other perissodactyl
groups; the Hyracotherium genus includes some species closely related to (or even ancestral to) rhinos and
tapirs, as well as species that are distinctly equine. [Note: the particular species that probably gave rise to the
rest of the equids, H. vassacciense, may be renamed, perhaps to "Protorohippus".]
Though in retrospect we may consider Hyracotherium to be "primitive", it was a very successful
animal in its time, and seems to have found a nice stable niche for itself. In fact, throughout most of the
Eocene (a good long 20 million years), only minor evolutionary changes took place in Hyracotherium and its
near descendants. The body and feet stayed mostly the same, with slight changes in the toes. The major
change was in the teeth; as Eocene equids started to eat more plant browse and less fruit, they developed
more grinding teeth to deal with the slightly tougher food.

Orohippus
In the early-middle Eocene (approx 50 My), there was a smooth, gradual transition from
Hyracotherium to a close relative, Orohippus (MacFadden, 1976). Overall, Orohippus looked much like
Hyracotherium: 10-20" high at the shoulder, still "doggish" with arched back, short legs, short neck, short
snout, and fairly small brain. Orohippus still had 4 toes on front and 3 behind, with hoofies, and was also
"pad-footed". However, the vestiges of the 1st and 2nd toes vanished.
The most significant change was in the teeth. The last premolar changed in shape to become like a
molar, giving Orohippus one more "grinding tooth". Also, the crests on the teeth were more pronounced,
indicating Orohippus was eating tougher plant material.

Epihippus
Epihippus arose from Orohippus in the middle Eocene (approximately 47 My). Like Orohippus and
Hyracotherium, Epihippus was small, doggish, pad-footed, and small-brained, with 4 toes in front and 3
behind. However, tooth evolution was continuing. Now the last two premolars were like molars, giving
Epihippus five grinding cheek teeth. The crests on the cheek teeth were well-formed, and still low-crowned.
There is a late form of Epihippus sometimes called Duchesnehippus. It's unclear if this is a subgenus
or a species of Epihippus. This animal was basically an Epihippus with teeth similar to, but a bit more
primitive than, later Oligocene horses.

IV. Medium-Sized Browsing Horses (Late Eocene & Oligocene)

As we move toward the Oligocene, horses start to change. The climate of North America was
becoming drier, and grasses were just evolving. The vast forests were starting to shrink. The late Eocene
horses responded by developing tougher teeth and becoming a bit larger and leggier (for better speed out in
the open).

Mesohippus
The species Mesohippus celer appears suddenly in the late Eocene, approx 40 My (such sudden
speciations can occur when a population encounters new selective forces and/or becomes isolated from the
parent species. These speciations are "sudden" only in geological terms, of course, where a few million years
is "sudden".) This animal was slightly larger than Epihippus, 24" at the shoulder. It didn't look as doggish,
either. The back was less arched, the legs a bit longer, the neck a bit longer, and the snout and face
distinctively longer. It had a shallow facial fossa, a depression on the skull. (In later horses these fossae
became complex, and handy for species identification.) Mesohippus had three toes on its hind feet and on its
front feet -- the 4th front toe was reduced to a vestigial nubbin. As before, Mesohippus was pad-footed. Other
significant changes:
 • Cerebral hemispheres notably larger -- has distinctly equine brain now.
• Last three premolars are like the three molars, such that Mesohippus (and all later horses)
had a battery of six similar grinding "cheek teeth", with one lonely little simple premolar in front.
• Has same tooth crests as Epihippus, well-formed and sharp, more suitable for grinding
tougher vegetation.

Miohippus
Soon after Mesohippus celer and its very close relative Mesohippus westoni appeared, a similar
animal called Miohippus assiniboiensis arose (approximately 36 My). This transition also occurred suddenly,
but luckily a few transitional fossils have been found that link the two genera. A typical Miohippus was
distinctly larger than a typical Mesohippus, with a slightly longer skull. The facial fossa was deeper and more
expanded. In addition, the ankle joint had changed subtly.
Miohippus also began to show a variable extra crest on its upper cheek teeth. In later horse species,
this crest became a characteristic feature of the teeth. This is an excellent example of how new traits
originate as variations in the ancestral population.
It was once thought that Mesohippus "transformed" gradually into Miohippus via anagenetic
evolution, so that only Miohippus continued. Recent evidence shows that instead, Miohippus speciated (split
off) from early Mesohippus via cladogenetic evolution, and then Miohippus and Mesohippus overlapped for
some 4 million years. For instance, in one place in modern Wyoming there were three species of late
Mesohippus coexisting with two species of Miohippus. (Prothero & Shubin, 1989)
V. The Miohippus Radiation (Early Miocene, 24 My)
Mesohippus finally died out in the mid-Oligocene. Miohippus continued for a while as it was, and
then, in early Miocene (24 My) began to speciate fairly rapidly. The horse family began to split into at least 2
main lines of evolution and one small side branch:
1. 3-toed browsers called "anchitheres". They were very successful, spread into the Old World,
and thrived for tens of millions of years. They retained the small, simple teeth of Miohippus. Genera
include Anchitherium and the large Hypohippus and Megahippus.
2. A line of small "pygmy horses", e.g. Archeohippus. These horses did not survive long.
3. A line that underwent a transformation from browsing to grazing, taking advantage of the new
grasses. Large grasslands were just beginning to appear, thus creating a new ecological
"opportunity" for grazers. Grass is difficult to chew and wears down teeth rapidly (due to the silica in
the leaves) and thus a grass-eater needs tough teeth with ridges of some sort. Open-country grass
eaters, in addition, often benefit from being swift runners with long legs. The evolution of this line of
horses is described below.

VI. Horses Move Onto the Plains: Spring-Foot & High-Crowned

Teeth (Miocene, 18 My)
As this third line of Miocene horses began to specialize in eating grasses, several changes occurred.
First, the teeth changed to be better suited for chewing harsh, abrasive grass. Small crests on the teeth
enlarged and connected together in a series of ridges for grinding. There was a gradual increase in the
height of the tooth crowns, so that the teeth could grow out of the gum continuously as the tops were worn
down ("hypsodont" teeth). And, in addition, the tooth crowns became harder due to the development of a
cement layer on the teeth.
Second, these horses started to become specialized runners. There was a simultaneous increase in
body size, leg length, and length of the face. The bones of the legs began to fuse together, and the leg bones
and musculature became specialized for efficient forward-and-back strides, with flexible leg rotation being
eliminated. Most significantly, the horses began to stand permanently on tiptoe (another adaptation for
speed); instead of walking on doglike pads, their weight was supported by springy ligaments that ran under
the fetlock to the big central toe. All these changes occurred rapidly, and we are lucky to have a fairly good
fossil record during this time. This was one of the most interesting times in horse evolution. The transitions in
these characters are seen in:

Kalobatippus
This genus is not well known, but its teeth seem to be intermediate between Miohippus and the later
Parahippus (see below).

Parahippus
Arose in early Miocene, 23 My. A typical Parahippus was a little larger than Miohippus, with about the
same size brain and same body form. Parahippus was still three-toed, and was just beginning to develop the
springy ligaments under the foot. Parahippus showed gradual and fluctuating changes in its teeth, including
the permanent establishment of the extra crest that was so variable in Miohippus. In addition, various other
cusps and crests were beginning to join up in a series of strong crests, with slightly taller tooth crowns.
Parahippus evolved rapidly and was quickly transformed into a fully spring- footed, hypsodont grazing horse
called Merychippus gunteri. This burst of evolution took place about 18-17 My. Later fossils of Parahippus
(e.g. the species Parahippus leonensis) are so similar to early Merychippus that it's hard to decide where to
draw the line between the genera.

Merychippus
Arose 17 My ago. A typical Merychippus was about 10 hands (40") tall, the tallest equine yet. The
muzzle became elongated, the jaw became deeper, and the eye moved farther back, to accommodate the
large tooth roots. The brain was notably larger, with a fissured neocortex and a larger cerebellum, making
Merychippus a smarter and more agile equine than the earlier horses. Overall, Merychippus was distinctly
recognizable as a horse, and had a "horsey" head.
Merychippus was still 3-toed, but was fully spring-footed. This animal stood permanently on tiptoe,
supported and propelled by strong, springy ligaments that ran under the fetlock. The side toes were still
complete, but began to be of varying sizes; some Merychippus species had full-size side toes, while others
developed small side toes that only touched the ground during running. The central toe developed a large,
convex, "horsey" hoof, and the legs became longer. The radius and ulna of the forearm fused so that leg
rotation was eliminated. Likewise, the fibula of the shin was greatly reduced. All these changes made
Merychippus' legs specialized for just one function: rapid running over hard ground.
Merychippus' teeth were fully high-crowned, with a thick layer of cement, and with the same
distinctive grazing tooth crests as Parahippus.
Merychippus gunteri evolved into a slightly more advanced form, Merychippus primus, in the
middle/late Miocene.
VII. The Merychippine Radiation (Miocene, 15 My)
By the late Miocene, Merychippus was the one of the first bona-fide speedy plains grazers.
(Simpson, 1961, called Merychippus "the horse with a new look"). Merychippus underwent rapid speciation,
and gave rise to at least 19 new grazing horse species in three major groups. This explosive burst of horse
evolution is often called the "merychippine radiation". The three major groups were:
1. Three-toed grazers known as "hipparions". These were tremendously successful and split
into 4 genera and at least 16 species, eventually covering a variety of niches for small and large
grazers and browsers. They developed large and elaborate facial fossae. Hipparions spread from the
New World into the Old World in several waves of migration.
2. A line of smaller horses including Protohippus and Calippus, collectively called
"protohippines".
3. A line of "true equines" in which the side toes sometimes began to decrease in size. In this
flurry of evolution, Merychippus primus gave rise to two later merychippines called M. sejunctus and
M. isonesus, who had a mixture of "primitive" (Parahippus-like), hipparion, and equine features.
They, in turn, gave rise to M. intermontanus, which begat M. stylodontus and M. carrizoensis. These
last two looked quite "horsey" and gave rise to a set of larger three-toed and one-toed horses known
as the "true equines" (see below). Crystal clear, right?
As this brief list shows, new species arose in rapid succession in all three of these groups. This rapid
speciation makes it hard to determine exactly which species arose from exactly which others.
About 10 My, the horse family reached an apex of diversity (of species and of genera) and sheer
numbers which it has never equalled since. The Old and New Worlds both seemed overrun with a wide
variety of hipparions, protohippines, and "true equines", large and small, forest browsers and plains grazers.
Throughout the evolution of all these related merychippine descendents, the facial fossae got deeper and
more elaborate. With so many equine species overlapping at once, these facial fossae may have housed
species-specific glands of some sort, similar to the scent- marking glands of modern antelopes and deer.

VIII. One-Toed Horses (Late Miocene, Pliocene & Pleistocene)

Let's leave the hipparions and protohippines now, and concentrate on the merychippine line that led
to the "true equines". The late merychippine species of this line, such as M. carrizoensis, were large horses
with small side toes. They gave rise to at least 2 separate groups of horses that independently lost their side
toes. This occurred as side ligaments developed around the fetlock to help stabilize the central toe during
running. These one-toed horses include:

Pliohippus
Arose in middle Miocene (~15 My) as a three-toed horse. Gradual loss of the side toes is seen in
Pliohippus through 3 successive strata of the early Pliocene. Pliohippus was very similar to Equus and until
recently was thought to be the direct ancestor of Equus, except for two significant differences. First,
Pliohippus's skull has deep facial fossae, whereas Equus has no facial fossae at all. Second, Pliohippus's
teeth are strongly curved, and Equus's teeth are very straight. Though Pliohippus is obviously related to
Equus, it probably didn't give rise to Equus.

Astrohippus
Astrohippus (~10My) was another one-toed horse that arose shortly after Pliohippus. Astrohippus
also had large facial fossae, and was probably a descendent of Pliohippus.
Dinohippus
Finally, a third one-toed horse called Dinohippus (recently discovered) arose about 12 My. The exact
ancestor of Dinohippus is not yet known (see Evander, 1989). The earliest known species are D. spectans,
D. interpolatus, and D. leidyanus. They look smashingly like Equus in foot morphology, teeth, and skull. The
teeth were slightly straighter than Merychippus, and the facial fossae were significantly decreased. A slightly
later species was D. mexicanus, that showed even straighter teeth and even smaller fossae. Dinohippus was
the most common horse in North America in the late Pliocene, and almost certainly gave rise to Equus.
(Recall that Equus has very straight teeth and no fossae.)
The Isthmus of Panama arose at this point. Some very early Dinohippus species gave rise to the
"hippidions", stocky, short-legged, one-toed horses with odd boxy skulls (~4 My). They travelled into the
South America and thrived there briefly.
Throughout the end of the Pliocene, Dinohippus showed a gradual decrease in the facial fossae,
straightening of the teeth, and other gradual changes, as Dinohippus smoothly graded into Equus. (Hulbert,
1989)

Equus
Finally we arrive at Equus (4 My), the genus of all modern equines. The first Equus were 13.2 hands
tall (pony size), with a classic "horsey" body -- rigid spine, long neck, long legs, fused leg bones with no
rotation, long nose, flexible muzzle, deep jaw. The brain was a bit larger than in early Dinohippus. Like
Dinohippus, Equus was (and is) one-toed, with side ligaments that prevent twisting of the hoof, and has high-
crowned, straight grazing teeth with strong crests lined with cement.
 Members of Equus still retain the genes for making side toes. Usually these express themselves only
as the vestigial "splint bones" of toes 2 and 4, around the large central 3rd toe. Very rarely, a modern Equus
is born with small but fully-formed side toes. (see Gould, Hen's Teeth and Horses' Toes.)
The earliest known Equus species were a set of three "simple Equus" species collectively known as
the Equus simplicidens group. They still had some primitive traits from Dinohippus, including a slight facial
fossa. They had zebra-like bodies (relatively stocky with a straight shoulder and thick neck), and short,
narrow, donkey-like skulls. They probably had stiff, upright manes, ropy tails, medium-sized ears, striped
legs, and at least some striping on the back (all traits shared by modern equines). They quickly diversified
into at least 12 new species in 4 different groups, in a burst of evolution reminiscent of the great
merychippine radiation. All these Equus species coexisted with other one-toed horses (such as Astrohippus)
and with various successful hipparions and protohippines, which had been merrily evolving on their own
paths.
During the first major glaciations of the late Pliocene (2.6 Ma), certain Equus species crossed to the
Old World. Some entered Africa and diversified into the modern zebras. Others spread across Asia, the
Mideast, & N. Africa as desert-adapted onagers and asses. Still others spread across Asia, the Mideast, and
Europe as the true horse, E. caballus. Other Equus species spread into South America. The Equus genus
was perhaps the most successful perissodactyl genus that ever lived -- even before domestication by
humans.
Compare Equus to Hyracotherium and see how much it has changed. In no way can Equus and
Hyracotherium be considered the same "kind". The change from Hyracotherium to Equus is truly long-term,
large-scale evolution.
IX. Modern Equines (Recent)
The three-toed horses gradually died out, perhaps outcompeted by the phenomenally successful
artiodactyls (or not). Most of the one-toed horses in North America also died out, as the Ice Ages started.
(The causes of these extinctions are unknown.) However, one-toed Equus was very successful. Until about 1
million years ago, there were Equus species all over Africa, Asia, Europe, North America, and South
America, in enormous migrating herds that must easily have equalled the great North American bison herds,
or the huge wildebeest migrations in Africa.
In the late Pleistocene there was a set of devastating extinctions that killed off most of the large
mammals in North and South America. All the horses of North and South America died out (along with the
mammoths and saber-tooth tigers). These extinctions seem to have been caused by a combination of
climatic changes and overhunting by humans, who had just reached the New World. For the first time in tens
of millions of years, there were no equids in the Americas.
The only members of Equus -- and of the entire family Equidae -- that survived to historic times were:
Order Perissodactyla, Family Equidae, Genus Equus
• Equus burchelli: the Plains zebra of Africa, including "Grant's zebra", "Burchell's zebra",
"Chapman's zebra", the half-striped Quagga, and other subspecies. The Plains zebra is what people
usually think of as the "typical zebra", with rather wide vertical stripes, and thick horizontal stripes on
the rump.
• Equus zebra: the Mountain zebra of South Africa. This is the little zebra with the dewlap and
the gridiron pattern on its rump.
 • Equus grevyi: Grevy's zebra, the most horse-like zebra. This is the big zebra with the very
narrow vertical stripes and huge ears.
• Equus caballus, the true horse, which once had several subspecies.
• Equus hemionus: the desert-adapted onagers of Asia & the Mideast, including the kiang
(formerly E. kiang).
• Equus asinus: the true asses & donkeys of northern Africa. (The African wild asses are
sometimes called E. africanus.)
[I have a separate file about the relationships & current status of all surviving wild equines, including
information about captive breeding programs. E-mail for details.]

X. SUMMARY
For many people, the horse family remains the classic example of evolution. As more and more
horse fossils have been found, some ideas about horse evolution have changed, but the horse family
remains a good example of evolution. In fact, we now have enough fossils of enough species in enough
genera to examine subtle details of evolutionary change, such as modes of speciation.
In addition to showing that evolution has occurred, the fossil Equidae also show the following
characteristics of evolution:
1. Evolution does not occur in a straight line toward a goal, like a ladder; rather, evolution is like
a branching bush, with no predetermined goal.
Horse species were constantly branching off the "evolutionary tree" and evolving along various
unrelated routes. There's no discernable "straight line" of horse evolution. Many horse species were
usually present at the same time, with various numbers of toes, adapted to various different diets. In
other words, horse evolution had no inherent direction. We only have the impression of straight-line
evolution because only one genus happens to still be alive, which deceives some people into thinking
that that one genus was somehow the "target" of all the evolution. Instead, that one genus is merely
the last surviving branch of a once mighty and sprawling "bush".
The view of equine evolution as a complex bush with many contemporary species has been around
for several decades, and is commonly recounted in modern biology and evolution textbooks.
2. There are no truly consistent "trends".
Tracing a line of descent from Hyracotherium to Equus reveals several apparant trends: reduction of
toe number, increase in size of cheek teeth, lengthening of the face, increase in body size. But these
trends are not seen in all of the horse lines. On the whole, horses got larger, but some horses
(Archeohippus, Calippus) then got smaller again. Many recent horses evolved complex facial pits,
and then some of their descendants lost them again. Most of the recent (5-10 My) horses were three-
toed, not one-toed, and we see a "trend" to one toe only because all the three-toed lines have
recently become extinct.
Additionally, these traits do not necessarily evolve together, or at a steady rate. The various
morphological characters each evolved in fits and starts, and did not evolve as a suite of characters.
For example, throughout the Eocene, the feet changed little, and only the teeth evolved. Throughout
the Miocene, both feet and teeth evolved rapidly. Rates of evolution depend on the ecological
pressures facing the species.
The "direction" of evolution depends on the ecological challenges facing the individuals of a species
and on the variation in that species, not on an inherent "evolutionary trend".
 3. New species can arise through several different evolutionary mechanisms.
Sometimes, new species split off suddenly from their ancestors (e.g., Miohippus from Mesohippus)
and then co-existed with those ancestors. Other species came into being through anagenetic
transformation of the ancestor, until the ancestor had changed appearance enough to be given a new
name (e.g. Equus from Dinohippus). Sometimes only one or a few species arose; sometimes there
were long periods of stasis (e.g. Hyracotherium throughout the early Eocene); and sometimes there
were enormous bursts of evolution, when new ecological opportunities arose (the merychippine
radiation). Again, evolution proceeds according to the ecological pressures facing the individuals of a
species and on the variation present within that species. Evolution takes place in the real world, with
diverse rates and modes, and cannot be reduced to a single, simple process.
A Question for Creationists: Creationists who wish to deny the evidence of horse evolution should
careful consider this: how else can you explain the sequence of horse fossils? Even if creationists insist on
ignoring the transitional fossils (many of which have been found), again, how can the unmistakable sequence
of these fossils be explained? Did God create Hyracotherium, then kill off Hyracotherium and create some
Hyracotherium-Orohippus intermediates, then kill off the intermediates and create Orohippus, then kill off
Orohippus and create Epihippus, then allow Epihippus to "microevolve" into Duchesnehippus, then kill off
Duchesnehippus and create Mesohippus, then create some Mesohippus-Miohippus intermediates, then
create Miohippus, then kill off Mesohippus, etc.....each species coincidentally similar to the species that
came just before and came just after?
Creationism utterly fails to explain the sequence of known horse fossils from the last 50 million years.
That is, without invoking the "God Created Everything To Look Just Like Evolution Happened" Theory.
 [And I'm not even mentioning all the other evidence for evolution that is totally independent of the
fossil record -- developmental biology, comparative DNA & protein studies, morphological analyses,
biogeography, etc. The fossil record, horses included, is only a small part of the story.]
Truly persistent and/or desperate creationists are thus forced into illogical, unjustified attacks of fossil
dating methods, or irrelevant and usually flat-out wrong proclamations about a supposed "lack" of
"transitional forms". It's sad. To me, the horse fossils tell a magnificent and fascinating story, of millions of
animals living out their lives, in their natural world, through millions of years. I am a dedicated horse rider and
am very happy that the one-toed grazing Equus survived to the present. Evolution in no way impedes my
ability to admire the beauty and nobility of these animals. Instead, it enriches my appreciation and
understanding of modern horses and their rich history.

"All the morphological changes in the history of the Equidae can be accounted for by the neo-
Darwinian theory of microevolution: genetic variation, natural selection, genetic drift, and
speciation." (Futuyma 1986, p.409)

"Because its complications are usually ignored by biology textbooks, creationists have claimed
the horse story is no longer valid. However, the main features of the story have in fact stood the
test of time...." (Futuyma 1982, p. 85)

"When asked to provide evidence of long-term evolution, most scientists turn to the fossil record.
Within this context, fossil horses are among the most frequently cited examples of evolution.
The prominent Finnish paleontologist Bjorn Kurten wrote: 'One's mind inevitably turns to that
inexhaustible textbook example, the horse sequence. This has been cited -- incorrectly more
 often than not -- as evidence for practically every evolutionary principle that has ever been
coined.' This cautionary note notwithstanding, fossil horses do indeed provide compelling
evidence in support of evolutionary theory." (MacFadden 1988, p. 131)

"The fossil record [of horses] provides a lucid story of descent with change for nearly 50 million
years, and we know much about the ancestors of modern horses." (Evander 1989, p. 125)

"It is evolution that gives rhyme and reason to the story of the horse family as it exists today and
as it existed in the past. Our own existence has the same rhyme and reason, and so has the
existence of every other living organism. One of the main points of interest in the horse family is
that it so clearly demonstrates this tremendously important fact." (Simpson, 1961, p. xxxiii)

[Return to Fossil Horses FAQs]

XI. References
I've tried to incorporate all the recent research I could find into this post. For more information, non-
scientists may want to start with Simpson's 1961 book, Horses. This book is a classic, readable account of
horse evolution, and though it's now somewhat outdated, I think it's still the most accessible introduction to
the topic. However, I strongly recommend that Simpson's book be supplemented with newer information from
MacFadden's (1988) nice summary and/or Prothero & Schoch's The Evolution of Perissodactyls (1989).
These and other selected references are listed below.
 Thanks to Larry Moran for the prototype of the ASCII horse tree and other various notes.
Bennett, D.K. 1986? (year not on my xerox! argh.) The origins of breeds. Equus 110:33, 11:37,
112:37. (This is a three-part series in a good-quality trade magazine, written for horse owners who have
some interest in science and evolution. (Further references are in the articles.) The author is a vertebrate
paleontologist who specializes in the evolution, form, and function of modern Equus. Her analysis shows that
E. caballus had at least 5 subspecies before domestication.)
Colbert, E.H. 1980. Evolution of the Vertebrates, 3rd edition. John Wiley & Sons, New York. Carroll,
R.L. 1988. Vertebrate Paleontology and Evolution. WH Freeman & Co., New York. (These are two standard
texts on vertebrate fossils & evolution. Colbert has a 4th edition out now.)
Futuyma, D.J. 1982. Science on Trial: The Case for Evolution. Pantheon Books, New York. (A well-
written book on the evidence for evolution, written for the layperson.)
Futuyma, D.J. 1986. Evolutionary Biology. Sinauer Associates, Sunderland, Mass. (A standard text
covering theories of how evolution occurs -- doesn't stress evidence for evolution per se.)
Gould, S.J. 1983. Hen's Teeth And Horse's Toes. Gould, S.J. 1991. Bully for Brontosaurus.
(Collections of essays written for Natural History magazine. "Hen's Teeth..." has essays on horse side toes
and zebra stripes; "Bully..." contains essays on "fox-terrier size" Hyracotherium and on the fallacy of
perceiving a direction of evolution in the horse family. Other essays are interesting too.)
Hildebrand, M. 1987. The mechanics of horse legs. Amer. Sci. 75:594-601. (Not about evolution, but
interesting & useful nonetheless.)
 Janis, C. 1976. The evolutionary strategy of the Equidae and the origins of rumen and cecal
digestion. Evolution 30:757-774. (An interesting analysis of the significance of hindgut fermentation in equids,
and on why the Equidae tend not to have high species diversity.)
Lowenstein, J.M., and O.A. Ryder. 1985. Immunological systematics of the extinct quagga (Equidae).
Experientia 41:1192-1193. (The authors studied molecules from skins of the extinct quagga, and conclude it
was a subspecies of the plains zebra.)
MacFadden, B.J. 1976. Cladistic analysis of primitive equids with notes on other perissodactyls.
Syst. Zool. 25(1):1-14. (An analysis of the interrelationships of Hyracotherium, Orohippus, Epihippus, the
paleotheres, and other early perissodactyls.)
MacFadden, B.J. 1984. Systematics and phylogeny of the Hipparion, Neohipparion, Nannippus, and
Cormohipparion (Mammalia, Equidae) from the Miocene and Pliocene of the New World. Bull. Am. Mus. Nat.
Hist. 179:1-196. (Extremely detailed analysis of evolution and interrelationships of the hipparions. [Okay,
okay, I didn't read the whole thing.] Relies to a large extent on the distinctive morphology of the facial fossae
in different species.)
MacFadden, B.J. 1984. Astrohippus and Dinohippus from the Yepomera local fauna (Hemphillian,
Mexico) and implications for the phylogeny of one-toed horses. J. Vert. Paleon. 4(2):273-283. (Description &
discussion of Pliohippus, Astrohippus, and Dinohippus. Concludes that Dinohippus was probably the
ancestor of Equus, and Pliohippus was probably the ancestor of Astrohippus.)
MacFadden, B.J. 1985. Patterns of phylogeny and rates of evolution in fossil horses: hipparions from
the Miocene and Pliocene of North America. Paleobiology 1(3):245-257. (Analyzes the evolution of hipparion
species. Of the 16 known species, 6 appear too suddenly for the mode of speciation to be determined. Of the
10 that appeared gradually enough for speciation mode to be determined, 5 have originated by anagenesis
(transformation of an ancestor species into a descendent species, such that the ancestor "disappears") and 5
by cladogenesis (splitting off of a new species from an ongoing ancestor species, such that the 2 species
continue to exist together.))
MacFadden, B.J. 1986. Late Hemphillian monodactyl horses (Mammalia, Equidae) from the Bone
Valley formation of central Florida. J. Paleontology 60(2):466-475. (Description of two recent discovered
advanced one-toed horse species: Astrohippus stocki and Dinohippus mexicanus.)
MacFadden, B.J. 1988. Horses, the fossil record, and evolution: a current perspective. Evol. Biol.
22:131-158. (A useful and readable update on current evidence & theories of horse evolution.)
MacFadden, B.J. 1993. (A new book about horse evolution. I have not read it yet but am trying to get
a copy. Over $70! sheesh.)
MacFadden, B.J., J.D. Bryant, and P.A. Mueller. 1991. Sr-isotopic, paleomagnetic, and
biostratigraphic evidence of horse evolution: evidence from the Miocene of Florida. Geology 19:242-245.
(This is an interesting example of the variety of dating methods paleontologists use to date their finds.
MacFadden et al. dated the Parahippus to Merychippus transition at a Florida site with paleomagnetic data
and Sr/Sr dates, and also by cross-correlation to other sites dated with Sr/Sr, K/Ar, Ar/Ar, zircon fission-track,
and paleomagnetic dating methods. Surprise, surprise, all the dates were consistent at roughly 16 My.)
MacFadden, B.J., & R.C. Hubbert. 1988. Explosive speciation at the base of the adaptive radiation of
Miocene grazing horses. Nature 336:466-468. (An interesting summary of the merychippine radiation. Has a
nice horse tree, too. MacFadden's horse tree is used by almost everyone these days.)
MacFadden, B.J., & M.F. Skinner. 1981. Earliest holarctic hipparion, Cormohipparion goorisi n.sp.
(Mammalia, Equidae) from the Barstovian (medial Miocene) Texas gulf coastal plain. J. Paleontology
55(3):619-627. (Description of a hipparion that was found to have crossed into the Old World from the New
World sooner than previously realized.)
Prothero, D.R., & R.M. Schoch, eds. 1989. The Evolution of Perissodactyls. Clarendon Press, New
York. (A compilation of current research and theories of perissodactyl evolution. The following chapters were
particularly useful:
• Evander, R.L. Phylogeny of the family Equidae. pp. 109-126
• MacFadden, B.J. Dental character variation in paleopopulations and morphospecies of fossil
horses and extant analogs. pp. 128-141
• Hulbert, R.C. Phylogenetic interrelationships and evolution of North) American late Neogene
Equinae. pp. 176-196.
• Prothero, D.R., & R.M. Schoch. Origin and evolution of the perissodactyla: summary and
synthesis. pp. 504-529.
• Prothero, D.R., & N. Shubin. The evolution of Oligocene horses. pp.142-175.
• Winans, M.C. A quantitative study of North American fossil species of the genus Equus. pp.
262-297.)
Radinsky, L. 1983. Allometry and reorganization in horse skull proportions. Science 221 (16
Sept):1189-1191 (Analysis of horse skull changes around the time that horses developed high-crowned
teeth, between 15 and 25 million years ago.)
Renders, E. 1984. The gait of Hipparion sp. from fossil footprints in Laetoli, Tanzania. Nature
308:179-181. (Interesting paper describing fossil hoofprints of an adult female hipparion and her foal. They
were using a gait called a "running walk".)
 Simpson, G.G. 1961. Horses. Doubleday & Co., New York. (An interesting and readable, though
outdated, account of horse evolution. Written for the intelligent non-scientist by a prominent paleontologist.)
Thomason, J.J. 1986. The functional morphology of the manus in the tridactyl equids Merychippus
and Mesohippus: paleontological inferences from neontological models. J. Vert. Pal. 6(2):143-161. (Analysis
of the pad-foot to spring-foot transition.)

"[Fossils] are animals, just as full of life as you are, even though they occur at different points in
the endless stream of time. Within their own segments of this stream, they breathe, eat, drink,
breed, fight, and live their own lives..." (Simpson, 1961, p. xxxiv)

[Return to Fossil Horses FAQs]

 W000

Histoire évolutive des équidés

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

 Présentation fossile d'espèce de la lignée ayant conduit à celle des chevaux.
La famille des Équidés (Equidae) recouvre toutes les espèces de la famille du cheval domestique,
les chevaux sauvages, les ânes et les zèbres.
Cette famille est issue d'une différentiation au sein des Périssodactyles et de laquelle sont issus
également les rhinocérotidés et tapiridés au cours de l'Éocène, il y a 55 Ma. Petit à petit, les membres de la
branche Equidae évoluèrent différemment selon les régions dans lesquelles ils vivaient et les climats
auxquels ils étaient confrontés.
Leur évolution a subi l'influence de leur environnement et très probablement directement de
l'homme. La découverte de cette histoire par les scientifiques a pu donner des arguments solides aux
partisans de la théorie de l'évolution face aux vues des créationnistes.
Sommaire
[masquer]
• 1 Histoire évolutive
• 1.1 Systématique
• 1.2 Équidés précoces
• 1.3 Du Miocène au Pliocène
• 1.4 Les anchithériininés
• 1.5 Equinae
• 1.5.1 Spéciation du genre Equus
• 1.5.1.1 En Eurasie
• 1.5.1.2 En Amérique
• 2 Histoire de cette théorie
• 3 Notes et références
• 4 Voir aussi
• 4.1 Lien externe

Histoire évolutive [modifier]

Au Néogène les continents actuels étaient formés et occupaient à peu de chose près leur
emplacement actuel, le continent eurasiatique n'était cependant pas en contact avec l'Afrique. Des fossiles
des mêmes genres ont été trouvés en Eurasie et en Amérique du nord, certaines de part et d'autre du détroit
de Bering, en effet un passage appelé Béringie a existé jusqu'il y a 11 600 ans environ. Classiquement, on
considère que les équidés modernes seraient apparus il y a 5 Ma[1], en trois étapes successives, les deux
premières au moins étant originaires d'Amérique du nord.

Systématique [modifier]
Les principaux groupes évolutifs sont décrits ci-dessous par phylogénie[2] :
o Hippomorpha (Wood, 1937)
|--o Pachynolophoidea (éteint) (Pavlow, 1888)
| `-- Pachynolophidae (éteint) (Pavlow, 1888)
`--o Equoidea (Gray, 1821 sensu Hay, 1902)
|?- Indolophus (éteint) (Pilgrim, 1925 [Indolophidae Schoch, 1984])
|-- Palaeotheriidae (éteint) (Bonaparte, 1850)
`--o Equidae (Gray, 1821)
|-- Anchitheriinae (éteint) (Leidy, 1869)
|-- Hyracotheriinae (éteint) Cope (1881), peut être paraphylétique
`--o Equinae (Gray, 1821 [non Leidy, 1869, Steinmann & Döderlein,
1890])
|-- Hipparionini (éteint) (Quinn, 1955)
|-- Protohippini (éteint) (Gidley, 1907)
`--o Equini (Gray, 1921)
| “Pliohippus” (éteint) (Marsh, 1874)
|
 `-- Equus

L'évolution dans cette famille s'est faite autant par changement progressif d'une fréquence du gène
dans la population entière (anagénèse) que par différentiations adaptatives et spéciations en branches
distinctes (cladogénèse). Les différentes espèces ancestrales ont pu également bénéficier d'une
convergence évolutive ce qui complique l'analyse des fossiles. Enfin, ponctuellement, les adaptations ont pu
d'une époque à l'autre, être en opposition (taille,…). Ce n'est que sur de longues périodes que l'on peut
identifier clairement les évolutions permanentes.
Équidés précoces [modifier]

Crâne et patte d'un Hyracotherium.

 Les équidés les plus primitifs découverts sont du genre Hyracotherium ou Eohippus. Certains
auteurs ne les considèrent pas encore comme faisant partie des équidés, d'autres différencient ces deux
genres. Ils disposent des caractéristiques de ce qu'allaient devenir les équidés modernes. Dotés de quatre
doigts, ils vivaient dans les forêts, durant l'Éocène, il y a 60 Ma, et ils évoluaient principalement sur ce qui
allait être l'Amérique du Nord, mais certains ont migré vers le Vieux Monde via le détroit de Béring[3].
L'Hyracotherium était petit — environ 20 cm — et trapu. Il se nourrissait de feuilles, menait une vie solitaire
dans les sous-bois. Il va progressivement évoluer en se spécialisant dans la course, grandir et acquérir un
cerveau semblable à celui des équidés modernes. On a appelé Orohippus l'étape évolutive vers 45 à 42 Ma
des sept espèces découvertes.
D'autres genres semblent en être issus. En Amérique du Nord, le Mesohippus, de la taille d'un grand
chien, vivait dans le climat chaud et sec il y a 42 à 33 Ma. Il semble avoir coexisté pour un temps avec le
Miohippus (40 à 25 Ma), encore plus grand que le précédent. Miohippus va ensuite donner naissance à deux
branches différentes sur ce continent dont plusieurs espèces migreront sur le continent eurasien. Ils se
distinguent par leur face allongée.
Du Miocène au Pliocène [modifier]

Évolution du pied des équidés.

De gauche à droite : Mesohippus, Merichippus, Pliohippus.
Les espèces de ces fossiles sont toutes considérées comme d'authentiques équidés.

Les anchithériininés [modifier]

• Kalobatippus
• Anchitherium, en Amérique du Nord et d'Europe à la Chine.
• Hypohippus, entre 17 et 11 Ma.
 • Megahippus, entre 15 et 11 Ma.
• Sinohippus, en Amérique du Nord et Chine.
• Archaeohippus

Equinae [modifier]

Hipparion gracile.
• Parahippus
Puis vint le Merychippus il y a 20 à 17 Ma, animal de 80 cm de haut. Les deux derniers doigts
continuent de s'atrophier et c'est le premier des équidés à ne se nourrir que d'herbe. Vers 13 Ma se
distinguent deux types d'équinés à traits morphologiques nettement distincts, les Hipparions et les
Pliohippus.
 Les Hipparions se distinguent des équidés modernes par leurs membres tridactyles et dessin de
l'émail sur les couronnes des molaires tandis que les Pliohippus, d'il y a 5 à 2 Ma, n'ont plus qu'un doigt. Ces
derniers sont donc considérés de partager un ancêtre avec les Equus plus récents, plutôt qu'avec les
Hipparions. Les Pliohippus sont présents en Amérique du Sud, du Nord et en Eurasie. De cette branche
commune d'Amérique du Nord émerge les Equus et les Dinohippus.

Spéciation du genre Equus [modifier]

Le genre Equus est apparu il y a 4 à 5 Ma[1]. Sa taille atteint 125 à 135 cm. Ses yeux sont disposés
sur les côtés pour détecter les prédateurs sur des grandes distances. Le naseau très ouvert, permet
l'inspiration de grande quantité d'air pour augmenter la vivacité et la rapidité[3].
L'histoire récente des équidés est mal connue. Nous ne savons pas en particulier quand a eu lieu la
spéciation entre les ânes, les chevaux sauvages (Equus ferus), les zèbres et le cheval domestique (Equus
caballus). Nous ne savons pas si les espèces domestiques (cheval ou âne) résultent d'une sélection opérée
par l'homme ou si elles sont le fruit de la sélection naturelle. L'histoire du cheval domestique est beaucoup
mieux connue depuis l'antiquité.
Cependant le plus ancien squelette de cheval, Equus stenonis a été daté de la fin du tertiaire en
Europe de l'ouest. Les scientifiques pensent qu'il est une évolution du Plesippus. Equus stenonis a ensuite
évolué en deux branches, l'une étant plus lourde que l'autre. Equus scotti, une forme similaire est commune
en Amérique, dont certaines sont devenues géantes. Ces espèces américaines se sont éteintes il y a
11 000 ans lors de l'extinction de la mégafaune américaine.
 Évolution du crâne du cheval : de son lointain ancêtre, l'Hyracotherium, de la taille d'un agneau, au
cheval d'aujour'hui

En Eurasie [modifier]
 Peinture de la Grotte Chauvet.
Article détaillé : cheval de Solutré.
Les équidés sont représentés sur les peintures pariétales remontant à plus de 30 000 ans,
notamment dans la Grotte Chauvet. Il est l'un des animaux les plus fréquemment représentés de l'art
préhistorique, comme par exemple dans la Grotte de Lascaux ; il ne s'agit probablement pas de l'ancêtre
direct des chevaux domestiques, mais ceux de l'espèce Equus ferus dans laquelle le Tarpan est classé.
Celui-ci a été chassé pour sa viande et les utilisations de certaines ressources telles que la peau, les
os, etc. La date des premières traces de domestication reste imprécise car les ossements, même plus
récents, ne permettent pas de distinguer, dans le cas du cheval, individus sauvages et individus
domestiques.
Les hypothèses actuelles penchent vers une domestication comme animal de prestige. Le musée
préhistorique d'Île-de-France a produit une exposition avançant des arguments convaincants en ce sens.
En Amérique [modifier]
Plusieurs genres se sont côtoyés, par exemple Hippidion et Equus.
Le site de la grotte de Pendejo est situé à une cinquantaine de kilomètres au sud d'Alamogordo, au
Nouveau-Mexique, et à environ quinze kilomètres au nord-est de l'extrémité méridionale des montagnes de
Sacramento. Une pointe d'alène ou de lance a été découverte dans un os d'une espèce de cheval
aujourd'hui disparue (Equus Amerhippus alaskae). L'ensemble reposait dans une couche stratigraphique
datée de 36 000 ans.
Sur les différents sites archéologiques situés autour du volcan Nevado de Toluca, un des plus
grands volcans de la région de Mexico, tels que les sites de Chimalhuacan ou de Balderas, de nombreux
ossements d'équidés furent mis au jour avec des squelettes humains et d'autres ossements d'animaux tels
que mammouths, glyptodons et camélidés. Tous furent tués il y a 10 500 ans lors de l'éruption plinienne du
volcan Nevado de Toluca.
En 1993, la découverte dans le Yukon canadien d'une peau vieille de 26 000 ans d'un Equus lambei
et de pattes momifiées permirent par une analyse ADN de montrer la très proche parenté avec les actuels
Equus[4].
L'Equus atteignit le continent sud-américain au début du Pléistocène et des espèces se
différencièrent : le cheval nain des Andes (Equus Amerhippus andium), le cheval préhistorique argentin
(Equus Amerhippus curvidens). On retrouve ses ossements dans le site de Pikimachay dans les Andes
péruviennes, il y a 22 000 ans, associés à des pointes de lances[5].
 Au Chili, dans la Caverne du Mylodon, furent mis au jour des indices de présence humaine, des
ossements d'Hippidum (Equus curvidens) et de mylodon répartis en plusieurs strates. La datation au
carbone 14 des ossements et artefacts trouvés donnent des dates comprises entre 12 000 et 13 000 ans.
Les espèces du genre Equus auraient finalement disparu du continent américain, il y a environ
10 000 ans seulement. L'homme, à travers les Amérindiens et Paléoaméricains, les carnassiers comme les
tigres aux dents de sabres, les éléments naturels, en particulier le volcanisme de la région centrale de
Mexico et des épizooties dévastatrices eurent raison de la présence des équidés en Amérique[6]. La
réintroduction d'espèces de la famille s'est faite lors de l'invasion de l'Amérique par les conquistadores.
Certaines espèces devenant marron, comme les mustangs.

Histoire de cette théorie [modifier]

Du fait que les équidés étaient absents des Amériques lorsque les Conquistadors espagnols ont
abordé leurs rivages et que les amérindiens n'avaient pas de mot pour désigner les équidés, on pourrait
s'imaginer que les chevaux sont originaires d'Eurasie, pourtant leur histoire évolutive semble plus
compliquée.
L'expédition de William Clark à Big Bone Lick en 1807 mit au jour de nombreux fossiles parmi
lesquels des os de la jambe et du pied de ce qui semblait être un cheval. Ce fossile a été ensuite pu être
évalué par l'anatomiste Caspar Wistar[7]. Ceci remettait en cause l'idée que les chevaux étaient originaires
uniquement d'Eurasie, mais Clark et Wistar ne soupçonnèrent pas l'importance de cette découverte. En
1833, lors de son voyage à bord du Beagle en Patagonie, Charles Darwin collecte des dents fossiles
d'équidés près de Santa Fe en Argentine. Ces dents, découvertes dans une couche stratigraphique
contenant notamment des os du tatou géant Glyptodon, ont été examinés par Richard Owen en 1840[7]. Ce
dernier confirma que ces dents appartenaient bien à un équidé et nomma l'espèce éteinte Equus curvidens.
Les découvertes de fossiles allaient ensuite se multiplier, entrainant une certaine confusion. En 1848,
Joseph Leidy, un des premiers naturalistes américains à reconnaître la validité de la théorie de l'évolution,
examina d'une manière systématique les fossiles du Pléistocène détenus par l’Academy of Natural Sciences
à Philadelphie, il en conclut qu'au moins une espèce vivait à cette époque en Amérique du Nord et pensa la
nommer Equus americanus. Mais comme ce nom binomial était déjà pris par une autre espèce d'Amérique
du sud, il renomma une dizaine d'année plus tard l'espèce nord-américaine: Equus complicatus[7]. Il décrit
ensuite de nombreux genres d'équidés (qui à l'époque était nommés Solidungulates), comme Anchippus,
Anchitherium, Hipparion, Hippodon, Merychippus, Parahippus et Protohippus. Dès cette époque, il note chez
ces fossiles ce qui semble être une complexification des couronnes dentaires.
Ce travail est repris par Othniel Charles Marsh qui fournit à partir des années 1870 les premiers
scénarios d'évolution des équidés. Ceux-ci sont largement basés sur l'évolution des dents. Ce travail sur les
équidés est le premier à fournir une série fossile complète pour illustrer la théorie de l'évolution (Darwin
comme Lamarck avant lui, s'étaient basés sur des exemples pris dans la biodiversité contemporaine)[7].
Toutefois le scénario, très linéaire et légèrement simpliste de Marsh, mais largement popularisé par Thomas
Huxley, a été bien entendu très critiqué.
Un grand nombre de fossiles a été ensuite découvert, et de nombreuses formes dites
transitionnelles (terme aujourd'hui abandonné, depuis les nouvelles découvertes sur la nature buissonnante
de la spéciation et des relations cladistiques entre espèces), complexifièrent d'autant le schéma initial.
L'existence de nombreuses branches évolutives, qui n'ont pas toutes eu de descendance, apparut. Les
études menées sur la fréquence et la répartition géographique et chronologique des fossiles d'équidés ont
révélé que l'évolution des espèces n'est ni aussi linéaire ni aussi uniforme qu'on l'a cru initialement. Des
"stases" (phases d'évolution lente, graduelle) entrecoupées de "crises" (phases d'évolution rapide,
buissonnante) furent découverts entre Dinohippus et Equus: des extinctions subites furent suivies de
l'apparition de formes différentes, par exemple pour la lignée de Epihippus à Mesohippus. Des "espèces
filles" ont pu coexister avec leurs espèces d'origine restées inchangées, et on a même observé des
inversions évolutives de certaines caractéristiques. Ce sont tous ces phénomènes évolutifs différents qui
expliquent la diversité des fossiles.
Un spécialiste des mammifères américains, George Gaylord Simpson[8], précurseur de la théorie
des équilibres ponctués[précision nécessaire], a contribué à rejeter cette vision linéaire de l'évolution, qui
ferait du cheval un « but » à celle-ci, à la manière de l’Intelligent design. Selon la théorie actuelle, les équidés
d'aujourd'hui ne sont tout simplement l'une des multiples lignées qui sont parvenues à survivre.
En 1958 furent mis au jour près de Lewisville, au Texas, de nombreux ossements de mammouths,
glyptodons, camélidés et équidés ainsi que des objets façonnés par des humains. En 1966, les fouilles
archéologiques du site de Tlapacoya au Mexique, permirent de dégager les ossements de plusieurs équidés,
ainsi que d'autres animaux et le squelette d'un paléoaméricain, montrant que les premiers amérindiens ont
chassé des équidés.
Tout est loin d'être élucidé cependant, et la question de l'origine proche des équidés vivants ne peut
pas être résolue par la paléontologie : en effet l'étude des fossiles ne permet pas de conclure définitivement
sur la différenciation des espèces, et plus particulièrement celle des chevaux actuels. En effet la spéciation
de ceux-ci semble très récente et la domestication y a joué un rôle (c'est d'ailleurs en étudiant et
expérimentant la sélection artificielle dans le cadre de la domestication, que Darwin a trouvé la majorité de
ses arguments). La technique paléontologique usuelle, basée sur la synapomorphie, n'est pas efficace pour
conclure, du fait notamment de la ressemblance des squelettes. Pour les chevaux, deux types de théories
s'affrontent depuis au moins la fin du XXe siècle: la première suppose que les lignées de chevaux actuelles
descendent d'une espèce unique, l'autre qu'ils descendent de plusieurs espèces sauvages. Depuis le début
des années 2000 des études basées sur phylogénie moléculaire essayent de répondre à cette interrogation.
L'origine récente des ânes ou des zèbres est moins étudiée et moins connue.

Notes et références [modifier]

1. ↑ a et b Référence The Paleobiology database : Equus Linnaeus 1758 [archive] (en)
2. ↑ Mikko's Phylogeny Archive [archive]
3. ↑ a et b Sibylle Luise Binder, Gabriele Kärcher (trad. Karine Van Butsele), La vie fascinante
des chevaux, Larousse, Paris, 2002 (ISBN 2035602890)
4. ↑ Yukon Horse [archive]
5. ↑ Flea Cave (Pikimachay), Sam E. Rustan, Minnesota State University [archive] (consulté le
25 février 2007)
6. ↑ « L'origine des chevaux de Hyracotherium à Equus - Partie 2 », dans minéraux & fossiles,
vol. 335, janvier 2005 [ résumé [archive] ]
7. ↑ a, b, c et d Ancient Horse Fossils [archive], Thomas Jefferson Fossil Collection, Academy of
Natural Sciences
8. ↑ Simpson, George Gaylord (1951): Horses. Oxford University Press; New Impression
edition. ISBN 0-19-500104-4 (1971 reprint)

Voir aussi [modifier]

• Histoire des mammifères
 Lien externe [modifier]
• Site de présentation de controverse :(en)Horse evolution by Kathlyn Hunt. Last updated
January 4, 1995

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive
 Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie
l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation
 [Dérouler]
v·d·m
Univers des équidés
Hippologie · Éthologie équine · Chuchoteur
Hippologie
· Pied · Robe · Allure · Écurie · Pansage

Sport hippique · Sport équestre · Centre

équestre · Équitation · Saut d'obstacles · Concours
Sports
complet d'équitation · Dressage · Randonnée
équestre

Évolution Histoire évolutive des équidés ·

et histoire Domestication du cheval · Chevalerie · Cavalerie

Races et Equidae · Cheval · Âne · Zèbre · Bardot ·

espèces Mulet · Liste des races chevalines

Palefrenier · Police montée · Vétérinaire ·

Métiers
Médecine équine · Maréchal-ferrant
 • Portail de l’origine et de l’évolution du vivant

• Portail du monde équestre

• Portail de la zoologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_%C3%A9quid
%C3%A9s ».

Catégories : Équidé préhistorique | Histoire évolutive | [+]

 W000

Histoire évolutive des hippopotamidés

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

 Arsinoitherium (en haut), un ancêtre de l'éléphant, et Bothriogenys fraasi (en bas), une espèce de la
famille des Anthracotheriidae. Ces derniers sont considérés comme des ancêtres des hippopotames.
 L'hippopotame nain de Crête (Hippopotamus creutzburgi) était de la taille d'un hippopotame nain de
l'ouest africain mais il était plus apparenté à l'hippopotame commun.
Il n'existe plus que deux espèces vivantes d'hippopotames, l'hippopotame commun et l'Hippopotame
nain de l'ouest africain. Les scientifiques ont répertorié cependant de très nombreux fossiles qu'ils estiment
être très proches des hippopotamidés actuels[1].
Comme son nom l'indique, les Grecs anciens voyaient dans l'hippopotame une sorte de cheval.
Jusqu'en 1985, les naturalistes regroupaient les hippopotames avec les Suidae (porcs et sangliers) et les
Tayassuidae (pécaris), en se fondant sur les particularités des molaires.
 Pourtant, l'étude des protéines du sang, puis de la systématique moléculaire et enfin plus récemment
de fossiles a bien montré que leurs parents génétiques les plus proches sont les cétacés. Selon cette
théorie, leurs ancêtres communs étaient semi-aquatiques et ceux-ci se sont différenciés des autres
artiodactyles il y a environ 60 Ma[2],[3].
Environ 6 Ma plus tard, ces ancêtres auraient évolués en deux branches[4], celle des cétacés et celle des
Anthracotheriodea. De ce groupe serait issus les Hippopotames et les anthracothères une famille de grands
quadrupèdes, dont les premiers membres, à la fin de l'Eocène, devaient étroitement ressembler aux
hippopotames actuels mais avec une tête relativement plus petite et plus mince[5]. Tous les anthracothères
ont disparu au cours du Pliocène sans laisser de descendance[2].
Au cours de l'Éocène et de l'Oligocène, plusieurs genres d'Anthracotheriodea ont été identifiés, par
exemple Anthracotherium et Elomeryx, au Miocène ce sont Merycopotamus et Libycosaurus tandis que les
derniers sont datés du Pliocène[6]. Libycosaurus étant considéré comme le plus proche des hippopotames
actuels. Leurs ancêtres communs auraient vécu au Miocène, il y a environ 20 Ma.
Le plus ancien hippopotamidé reconnu est du genre Kenyapotamus, il vivait en Afrique entre 16 et 8
Ma. Kenyapotamus n'est connu que par des fragments de fossiles, mais on sait qu'il était de taille similaire
aux hippopotames nains[6]. Les hippopotamidae semblent avoir évolué seulement en Afrique et, à partir d'un
seul groupe, puis s'être propagés dans toute l'Asie et l'Europe, d'ailleurs aucun hippopotame préhistorique
n'a été découvert en Amérique et bien que divers anthracothères aient émigré en Amérique du Nord au
début de l'Oligocène[1]. Il y a entre 7,5 et 1,8 Ma, l'Archaeopotamus, probable ancêtre des genres
Hippopotamus et Hexaprotodon, vivait en Afrique et au Moyen-Orient[7].
Bien que les lignées d'hippopotames préhistoriques soient encore mal connues, tout pousse à croire
que les lignées des deux espèces modernes auraient divergé il y a déjà 8 Ma. La forme ancestrale de
l'hippopotame pygmée africain pourrait être le genre Saotherium. Saotherium et Choeropsis ont des traits
nettement plus basaux que les Hippopotamus et les Hexaprotodons, et sont donc plus proches de l'espèce
ancestrale des hippopotames[7],[6].
De nombreuses espèces ont disparu durant la dernière glaciation européenne, notamment
Hippopotamus antiquus et Hippopotamus gorgops qui vivaient dans la zone concernée. Ces deux espèces
étaient plus grandes que les espèces actuelles, il existait également de nombreuses espèces naines comme
Hippopotamus minor à Chypre. La dernière espèce disparue est Hexaprotodon madagascariensis, elle était
encore présente à Madagascar vers le XIVe siècle.

Notes et références [modifier]

1. ↑ a et b Référence The Paleobiology database : Hippopotamidae Gray 1821 [archive] (en)
2. ↑ a et b (en) « Scientists find missing link between the dolphin, whale and its closest
relative, the hippo », dans Science News Daily, 2005-01-25 [ texte intégral [archive] (page consultée
le 2007-06-18) ]
3. ↑ (en) Gatesy, J, « More DNA support for a Cetacea/Hippopotamidae clade: the blood-
clotting protein gene gamma-fibrinogen », dans Molecular Biology and Evolution, vol. 14, {{{année}}},
p. 537–543 [ texte intégral [archive] ]
4. ↑ (en) Ursing B.M.; U. Arnason, « Analyses of mitochondrial genomes strongly support a
hippopotamus-whale clade », dans Proceedings of the Royal Society, vol. 265, no 1412, 1998,
p. 2251 [ lien DOI [archive] ]
5. ↑ (en) Boisserie Jean-Renaud, Fabrice Lihoreau et Michel Brunet, « The position of
Hippopotamidae within Cetartiodactyla », dans Proceedings of the National Academy of Sciences,
 vol. 102, no 5, février 2005, p. 1537–1541 [ texte intégral [archive], lien DOI [archive] (pages
consultées le 2007-06-09) ]
6. ↑ a, b et c (en) Boisserie Jean-Renaud, Fabrice Lihoreau et Michel Brunet, « Origins of
Hippopotamidae (Mammalia, Cetartiodactyla): towards resolution », dans Zoologica Scripta, vol. 34,
no 2, mars 2005, p. 119–143 [ résumé [archive], lien DOI [archive] (pages consultées le 2007-06-01) ]
7. ↑ a et b (en) Boisserie Jean-Renaud, « The phylogeny and taxonomy of Hippopotamidae
(Mammalia: Artiodactyla): a review based on morphology and cladistic analysis », dans Zoological
Journal of the Linnean Society, vol. 143, 2005, p. 1–26 [ résumé [archive], lien DOI [archive] (pages
consultées le 01 06 2007) ]

Voir aussi [modifier]

Articles connexes [modifier]
• histoire évolutive des mammifères
• histoire évolutive des cétacés

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes
 Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et
Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Mammifères Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire
 évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

• Portail de la zoologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_hippopotamid
%C3%A9s ».

Catégories : Histoire évolutive | Hippopotamidae | [+]

 W000

Human evolution
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article is about the general evolution of humans. For a timeline see Timeline of human evolution.
Human evolution, or anthropogenesis, is the origin and evolution of Homo sapiens as a distinct
species from other hominids, great apes and placental mammals. The study of human evolution
encompasses many scientific disciplines, including physical anthropology, primatology, archaeology,
linguistics and genetics.[1]
 The term "human" in the context of human evolution refers to the genus Homo, but studies of human
evolution usually include other hominids, such as the Australopithecines, from which the genus Homo had
diverged by about 2.3 to 2.4 million years ago in Africa.[2][3] Scientists have estimated that humans
branched off from their common ancestor with chimpanzees about 5–7 million years ago. Several species and
subspecies of Homo evolved and are now extinct. These include Homo erectus, which inhabited Asia, and
Homo sapiens neanderthalensis, which inhabited Europe. Archaic Homo sapiens evolved between 400,000
and 250,000 years ago.
The dominant view among scientists concerning the origin of anatomically modern humans is the
"Out of Africa" or recent African origin hypothesis,[4][5][6][7] which argues that Homo sapiens arose in Africa
and migrated out of the continent around 50,000 to 100,000 years ago, replacing populations of Homo
erectus in Asia and Homo neanderthalensis in Europe. Scientists supporting the alternative multiregional
hypothesis argue that Homo sapiens evolved as geographically separate but interbreeding populations
stemming from a worldwide migration of Homo erectus out of Africa nearly 2.5 million years ago.
Contents
[hide]
• 1 History of ideas
• 2 Before Homo
• 2.1 Evolution of the great apes
• 2.2 Divergence of the human lineage from other
Great Apes
• 3 Genus Homo
• 3.1 Habilis
• 3.2 Rudolfensis and Georgicus
• 3.3 Ergaster and Erectus
• 3.4 Cepranensis and Antecessor
• 3.5 Heidelbergensis
• 3.6 Rhodesiensis, and the Gawis cranium
• 3.7 Neanderthalensis
• 3.8 Sapiens
• 3.9 Floresiensis
• 3.10 Denisova hominin
• 3.11 Comparative table of Homo species
• 4 Use of tools
• 4.1 Stone tools
[edit] History of ideas
The word homo, the name of the biological genus to which humans belong, is Latin for "human". It
was chosen originally by Carolus Linnaeus in his classification system. The word "human" is from the Latin
humanus, the adjectival form of homo. The Latin "homo" derives from the Indo-European root, dhghem, or
"earth".[8]
Carolus Linnaeus and other scientists of his time also considered the great apes to be the closest
relatives of humans due to morphological and anatomical similarities. The possibility of linking humans with
earlier apes by descent only became clear after 1859 with the publication of Charles Darwin's On the Origin
of Species. This argued for the idea of the evolution of new species from earlier ones. Darwin's book did not
address the question of human evolution, saying only that "Light will be thrown on the origin of man and his
history".
 Fossil Hominid Evolution Display at The Museum of Osteology, Oklahoma City, USA.
The first debates about the nature of human evolution arose between Thomas Huxley and Richard
Owen. Huxley argued for human evolution from apes by illustrating many of the similarities and differences
between humans and apes and did so particularly in his 1863 book Evidence as to Man's Place in Nature.
However, many of Darwin's early supporters (such as Alfred Russel Wallace and Charles Lyell) did not agree
that the origin of the mental capacities and the moral sensibilities of humans could be explained by natural
selection. Darwin applied the theory of evolution and sexual selection to humans when he published The
Descent of Man in 1871.[9]
 A major problem was the lack of fossil intermediaries. It was only in the 1920s that such fossils were
discovered in Africa. In 1925, Raymond Dart described Australopithecus africanus. The type specimen was
the Taung Child, an Australopithecine infant discovered in a cave. The child's remains were a remarkably
well-preserved tiny skull and an endocranial cast of the individual's brain. Although the brain was small
(410 cm³), its shape was rounded, unlike that of chimpanzees and gorillas, and more like a modern human
brain. Also, the specimen showed short canine teeth, and the position of the foramen magnum was evidence
of bipedal locomotion. All of these traits convinced Dart that the Taung baby was a bipedal human ancestor,
a transitional form between apes and humans.
The classification of humans and their relatives has changed considerably over time. The gracile
Australopithecines are now thought to be ancestors of the genus Homo, the group to which modern humans
belong. Both Australopithecines and Homo sapiens are part of the tribe Hominini. Recent data suggests
Australopithecines were a diverse group and that A. africanus may not be a direct ancestor of modern
humans. Reclassification of Australopithecines that originally were split into either gracile or robust varieties
has put the latter into a family of its own, Paranthropus. Taxonomists place humans, Australopithecines and
related species in the same family as other great apes, in the Hominidae.

Hominin species distributed through time edit

 Note: 1e+06 years = 1 × 106 years = 1 million years ago = 1 Ma

[edit] Before Homo

For evolutionary history before primates, see evolution of mammals and evolutionary history of life.

[edit] Evolution of the great apes

Plesiadapis.
The evolutionary history of the primates can be traced back 65 million years, as one of the oldest of
all surviving placental mammal groups. The oldest known primate-like mammal species, the Plesiadapis,
come from North America, but they were widespread in Eurasia and Africa during the tropical conditions of
the Paleocene and Eocene.
 Notharctus.
With the beginning of modern climates, marked by the formation of the first Antarctic ice in the early
Oligocene around 30 million years ago. A primate from this time was Notharctus. Fossil evidence found in
Germany in the 1980s was determined to be about 16.5 million years old, some 1.5 million years older than
similar species from East Africa and challenging the original theory regarding human ancestry originating on
the African continent.
David Begun[10] says that these primates flourished in Eurasia and that the lineage leading to the
African apes and humans— including Dryopithecus—migrated south from Europe or Western Asia into Africa.
The surviving tropical population, which is seen most completely in the upper Eocene and lowermost
Oligocene fossil beds of the Fayum depression southwest of Cairo, gave rise to all living primates—lemurs of
Madagascar, lorises of Southeast Asia, galagos or "bush babies" of Africa, and the anthropoids; platyrrhines
or New World monkeys, and catarrhines or Old World monkeys and the great apes and humans.
 The earliest known catarrhine is Kamoyapithecus from uppermost Oligocene at Eragaleit in the
northern Kenya Rift Valley, dated to 24 million years ago. Its ancestry is generally thought to be species
related to Aegyptopithecus, Propliopithecus, and Parapithecus from the Fayum, at around 35 million years
ago.[citation needed] In 2010, Saadanius was described as a close relative of the last common ancestor of
the crown catarrhines, and tentatively dated to 29–28 million years ago, helping to fill an 11-million-year gap
in the fossil record.[11]

Reconstructed tailless Proconsul skeleton.

In the early Miocene, about 22 million years ago, the many kinds of arboreally adapted primitive
catarrhines from East Africa suggest a long history of prior diversification. Fossils at 20 million years ago
include fragments attributed to Victoriapithecus, the earliest Old World Monkey. Among the genera thought to
be in the ape lineage leading up to 13 million years ago are Proconsul, Rangwapithecus, Dendropithecus,
Limnopithecus, Nacholapithecus, Equatorius, Nyanzapithecus, Afropithecus, Heliopithecus, and
Kenyapithecus, all from East Africa. The presence of other generalized non-cercopithecids of middle
Miocene age from sites far distant—Otavipithecus from cave deposits in Namibia, and Pierolapithecus and
Dryopithecus from France, Spain and Austria—is evidence of a wide diversity of forms across Africa and the
Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle
Miocene. The youngest of the Miocene hominoids, Oreopithecus, is from 9 million year old coal beds in Italy.
Molecular evidence indicates that the lineage of gibbons (family Hylobatidae) became distinct from
Great Apes between 18 and 12 million years ago, and that of orangutans (subfamily Ponginae) became
distinct from the other Great Apes at about 12 million years; there are no fossils that clearly document the
ancestry of gibbons, which may have originated in a so-far-unknown South East Asian hominoid population,
but fossil proto-orangutans may be represented by Ramapithecus from India and Griphopithecus from
Turkey, dated to around 10 million years ago.

[edit] Divergence of the human lineage from other Great Apes

Species close to the last common ancestor of gorillas, chimpanzees and humans may be
represented by Nakalipithecus fossils found in Kenya and Ouranopithecus found in Greece. Molecular
evidence suggests that between 8 and 4 million years ago, first the gorillas, and then the chimpanzees
(genus Pan) split off from the line leading to the humans; human DNA is approximately 98.4% identical to
that of chimpanzees when comparing single nucleotide polymorphisms (see Human evolutionary genetics).
The fossil record of gorillas and chimpanzees is quite limited. Both poor preservation (rain forest soils tend to
be acidic and dissolve bone) and sampling bias probably contribute to this problem.
Other hominines likely adapted to the drier environments outside the equatorial belt, along with
antelopes, hyenas, dogs, pigs, elephants, and horses. The equatorial belt contracted after about 8 million
years ago. Fossils of these hominans - the species in the human lineage following divergence from the
chimpanzees - are relatively well known. The earliest are Sahelanthropus tchadensis (7 Ma) and Orrorin
tugenensis (6 Ma), followed by:
• Ardipithecus (5.5–4.4 Ma), with species Ar. kadabba and Ar. ramidus;
• Australopithecus (4–1.8 Ma), with species Au. anamensis, Au. afarensis, Au. africanus, Au.
bahrelghazali, Au. garhi, and Au. sediba;
• Kenyanthropus (3–2.7 Ma), with species Kenyanthropus platyops;
• Paranthropus (3–1.2 Ma), with species P. aethiopicus, P. boisei, and P. robustus;
• Homo (2 Ma–present), with species Homo habilis, Homo rudolfensis, Homo ergaster, Homo
georgicus, Homo antecessor, Homo cepranensis, Homo erectus, Homo heidelbergensis, Homo
rhodesiensis, Homo neanderthalensis, Homo sapiens idaltu, Archaic Homo sapiens, Homo
floresiensis.

[edit] Genus Homo

Homo sapiens is the only extant species of its genus, Homo. While some other, extinct, Homo
species might have been ancestors of Homo sapiens, many were likely our "cousins", having speciated away
from our ancestral line.[12] There is not yet a consensus as to which of these groups should count as
separate species and which as subspecies. In some cases this is due to the dearth of fossils, in other cases it
is due to the slight differences used to classify species in the Homo genus. The Sahara pump theory
(describing an occasionally passable "wet" Sahara Desert) provides an explanation of the early variation in
the genus Homo.
 Based on archaeological and paleontological evidence, it has been possible to infer, to some extent,
the ancient dietary practices of various Homo species and to study the role of diet in physical and behavioral
evolution within Homo.[13][14][15][16][17]

[edit] Habilis
Homo habilis lived from about 2.4 to 1.4 Ma. Homo habilis, the first species of the genus Homo,
evolved in South and East Africa in the late Pliocene or early Pleistocene, 2.5–2 Ma, when it diverged from
the Australopithecines. Homo habilis had smaller molars and larger brains than the Australopithecines, and
made tools from stone and perhaps animal bones. One of the first known hominids, it was nicknamed 'handy
man' by its discoverer, Louis Leakey due to its association with stone tools. Some scientists have proposed
moving this species out of Homo and into Australopithecus due to the morphology of its skeleton being more
adapted to living on trees rather than to moving on two legs like Homo sapiens.[18]

[edit] Rudolfensis and Georgicus

These are proposed species names for fossils from about 1.9–1.6 Ma, the relation of which with
Homo habilis is not yet clear.
• Homo rudolfensis refers to a single, incomplete skull from Kenya. Scientists have suggested
that this was another Homo habilis, but this has not been confirmed.[19]
• Homo georgicus, from Georgia, may be an intermediate form between Homo habilis and
Homo erectus,[20] or a sub-species of Homo erectus.[21]
[edit] Ergaster and Erectus
 One current view of the temporal and geographical distribution of hominid populations.[22] Other
interpretations differ mainly in the taxonomy and geographical distribution of hominid species.
The first fossils of Homo erectus were discovered by Dutch physician Eugene Dubois in 1891 on the
Indonesian island of Java. He originally gave the material the name Pithecanthropus erectus based on its
morphology that he considered to be intermediate between that of humans and apes.[23] Homo erectus (H
erectus) lived from about 1.8 Ma to about 70,000 years ago (which would indicate that they were probably
wiped out by the Toba catastrophe; however, Homo erectus soloensis and Homo floresiensis survived it).
Often the early phase, from 1.8 to 1.25 Ma, is considered to be a separate species, Homo ergaster, or it is
seen as a subspecies of Homo erectus, Homo erectus ergaster. In the early Pleistocene, 1.5–1 Ma, in Africa,
Asia, and Europe, some populations of Homo habilis are thought to have evolved larger brains and made
more elaborate stone tools; these differences and others are sufficient for anthropologists to classify them as
a new species, Homo erectus. In addition Homo erectus was the first human ancestor to walk truly upright.
[24] This was made possible by the evolution of locking knees and a different location of the foramen
magnum (the hole in the skull where the spine enters). They may have used fire to cook their meat.
See also: Control of fire by early humans
A famous example of Homo erectus is Peking Man; others were found in Asia (notably in Indonesia),
Africa, and Europe. Many paleoanthropologists now use the term Homo ergaster for the non-Asian forms of
this group, and reserve Homo erectus only for those fossils that are found in Asia and meet certain skeletal
and dental requirements which differ slightly from H. ergaster.
[edit] Cepranensis and Antecessor
These are proposed as species that may be intermediate between H. erectus and H.
heidelbergensis.
• H. antecessor is known from fossils from Spain and England that are dated 1.2 Ma–500 ka.
[25][26]
• H. cepranensis refers to a single skull cap from Italy, estimated to be about 800,000 years
old.[27]

[edit] Heidelbergensis
H. heidelbergensis (Heidelberg Man) lived from about 800,000 to about 300,000 years ago. Also
proposed as Homo sapiens heidelbergensis or Homo sapiens paleohungaricus.[28]

[edit] Rhodesiensis, and the Gawis cranium

• H. rhodesiensis, estimated to be 300,000–125,000 years old. Most current experts believe
Rhodesian Man to be within the group of Homo heidelbergensis, though other designations such as
Archaic Homo sapiens and Homo sapiens rhodesiensis have also been proposed.
• In February 2006 a fossil, the Gawis cranium, was found which might possibly be a species
intermediate between H. erectus and H. sapiens or one of many evolutionary dead ends. The skull
from Gawis, Ethiopia, is believed to be 500,000–250,000 years old. Only summary details are known,
and no peer reviewed studies have been released by the finding team. Gawis man's facial features
suggest its being either an intermediate species or an example of a "Bodo man" female.[29]
 [edit] Neanderthalensis

Le Ferrassie Neanderthal skull (cast)

H. neanderthalensis lived from 400,000[30] years ago. Also proposed as Homo sapiens
neanderthalensis: there is ongoing debate over whether the "Neanderthal Man" was a separate species,
Homo neanderthalensis, or a subspecies of H. sapiens.[31] While the debate remains unsettled, evidence
from sequencing mitochondrial DNA indicates that no significant gene flow occurred between H.
neanderthalensis and H. sapiens, and, therefore, the two were separate species that shared a common
ancestor about 660,000 years ago.[32][33] In 1997, Mark Stoneking stated: "These results [based on
mitochondrial DNA extracted from Neanderthal bone] indicate that Neanderthals did not contribute
mitochondrial DNA to modern humans… Neanderthals are not our ancestors." Subsequent investigation of a
second source of Neanderthal DNA supported these findings.[34] However, the 2010 sequencing of the
Neanderthal genome indicates that Neanderthals did indeed interbreed with H. sapiens circa 75,000 BC
(after H. sapiens moved out from Africa, but before they separated into Europe, the Middle East, and Asia).
[35] Nearly all modern humans have 1% to 4% of their DNA derived from Neanderthal DNA.[35] (To
appreciate how big of a percentage this is, consider that humans and chimps only differ in 1.5% of their
DNA.)[citation needed] This 1-4% bit of DNA is only present in non-african humans.[35] However, supporters
of the multiregional hypothesis point to recent studies indicating non-African nuclear DNA heritage dating to
one Ma,[36] although the reliability of these studies has been questioned.[37] Competition from Homo
sapiens probably contributed to Neanderthal extinction.[38][39] They could have coexisted in Europe for as
long as 10,000 years.[40]

[edit] Sapiens
Main article: Early Homo sapiens
H. sapiens (the adjective sapiens is Latin for "wise" or "intelligent") have lived from about 250,000
years ago to the present. Between 400,000 years ago and the second interglacial period in the Middle
Pleistocene, around 250,000 years ago, the trend in skull expansion and the elaboration of stone tool
technologies developed, providing evidence for a transition from H. erectus to H. sapiens. The direct
evidence suggests there was a migration of H. erectus out of Africa, then a further speciation of H. sapiens
from H. erectus in Africa. A subsequent migration within and out of Africa eventually replaced the earlier
dispersed H. erectus. This migration and origin theory is usually referred to as the recent single origin or Out
of Africa theory. Current evidence does not preclude some multiregional evolution or some admixture of the
migrant H. sapiens with existing Homo populations. This is a hotly debated area of paleoanthropology.
Current research has established that humans are genetically highly homogenous; that is, the DNA
of individuals is more alike than usual for most species, which may have resulted from their relatively recent
evolution or the possibility of a population bottleneck resulting from cataclysmic natural events such as the
Toba catastrophe.[41][42][43] Distinctive genetic characteristics have arisen, however, primarily as the result
of small groups of people moving into new environmental circumstances. These adapted traits are a very
small component of the Homo sapiens genome, but include various characteristics such as skin color and
nose form, in addition to internal characteristics such as the ability to breathe more efficiently in high
altitudes.
H. sapiens idaltu, from Ethiopia, is a possible extinct sub-species who lived from about 160,000
years ago.

[edit] Floresiensis
Main article: Homo floresiensis
H. floresiensis, which lived from approximately 100,000 to 12,000 before present, has been
nicknamed hobbit for its small size, possibly a result of insular dwarfism.[44] H. floresiensis is intriguing both
for its size and its age, being a concrete example of a recent species of the genus Homo that exhibits derived
traits not shared with modern humans. In other words, H. floresiensis share a common ancestor with modern
humans, but split from the modern human lineage and followed a distinct evolutionary path. The main find
was a skeleton believed to be a woman of about 30 years of age. Found in 2003 it has been dated to
approximately 18,000 years old. The living woman was estimated to be one meter in height, with a brain
volume of just 380 cm3 (considered small for a chimpanzee and less than a third of the H. sapiens average of
1400 cm3).
However, there is an ongoing debate over whether H. floresiensis is indeed a separate species.[45]
Some scientists presently believe that H. floresiensis was a modern H. sapiens suffering from pathological
dwarfism.[46] This hypothesis is supported in part, because some modern humans who live on Flores, the
island where the skeleton was found, are pygmies. This coupled with pathological dwarfism, it is argued,
could indeed create a hobbit-like human. The other major attack on H. floresiensis is that it was found with
tools only associated with H. sapiens.[46]
The hypothesis of pathological dwarfism, however, fails to explain additional anatomical features that
are unlike those of modern humans (diseased or not) but much like those of ancient members of our genus.
Aside from cranial features, these features include the form of bones in the wrist, forearm, shoulder, knees,
and feet.

[edit] Denisova hominin

Main article: Denisova hominin
In 2008, archeologists working at the site of Denisova Cave in the Altai Mountains of Siberia
uncovered a small bone fragment from the fifth finger of a juvenile hominin, dubbed the "X-woman" (referring
to the maternal descent of mitochondrial DNA[47], or the Denisova hominin. Artifacts, including a bracelet,
excavated in the cave at the same level were carbon dated to around 40,000 BP. As DNA had survived in the
fossil fragment due to the cool climate of the Denisova Cave, a team of scientists from the Max Planck
Institute for Evolutionary Anthropology in Leipzig, Germany sequenced mtDNA extracted from the fragment.
[48]
The analysis indicated that modern humans, Neanderthals, and the Denisova hominin last shared a
common ancestor around 1 million years ago.[49] Modern humans are known to have overlapped with
Neanderthals in Europe for more than 10,000 years, and the discovery raises the possibility that
Neanderthals, modern humans and the Denisovan hominin may have co-existed together.
 The DNA analysis further indicated that this new hominin species was the result of an early migration
out of Africa, distinct from the later out-of-Africa migrations associated with Neanderthals and modern
humans, but also distinct from the earlier African exodus of Homo erectus.[49] Professor Chris Stringer,
human origins researcher at London's Natural History Museum and one of the leading proponents of the
recent single-origin hypothesis, remarked: "This new DNA work provides an entirely new way of looking at
the still poorly understood evolution of humans in central and eastern Asia." Pääbo noted that the existence
of this distant branch creates a much more complex picture of humankind during the Late Pleistocene.[47]

[edit] Comparative table of Homo species

view • talk • edit
Comparative table of Homo species
Lived when Lived Adult Adult Crani
Species
(Ma) where height mass capacity (cm³)

1.75 m 90 kg
H. antecessor 1.2 – 0.8 Spain 1,000
(5.7 ft) (200 lb)

H. cepranensis 0.9 – 0.8? Italy 1,000

Africa,
850
Eurasia (Java, 1.8 m 60 kg
H. erectus 1.5 – 0.2 (early) – 1,100
China, India, (5.9 ft) (130 lb)
(late)
Caucasus)
 Eastern
1.9 m
H. ergaster 1.9 – 1.4 and Southern 700–85
(6.2 ft)
Africa

1.0 m 25 kg
H. floresiensis 0.10? – 0.012 Indonesia 400
(3.3 ft) (55 lb)

South 1.0 m
H. gautengensis >2 – 0.6
Africa (3.3 ft)

H. georgicus 1.8 Georgia 600

1.0–1.5 33–55 kg
H. habilis 2.3 – 1.4 Africa 510–66
m (3.3–4.9 ft) (73–120 lb)

Europe, 1.8 m 60 kg 1,100–

H. heidelbergensis 0.6 – 0.35
Africa, China (5.9 ft) (130 lb) 1,400

55–70 kg
Europe, 1.6 m 1,200–
H. neanderthalensis 0.35 – 0.03 (120–150 lb)
Western Asia (5.2 ft) 1,900
(heavily built)

H. rhodesiensis 0.3 – 0.12 Zambia 1,300

H. rudolfensis 1.9 Kenya

 H. sapiens idaltu 0.16 – 0.15 Ethiopia 1,450

H. sapiens sapiens 1.4–1.9 50–100 1,000–

0.2 – present Worldwide
(modern humans) m (4.6–6.2 ft) kg (110–220 lb) 1,850

[edit] Use of tools

"A sharp rock", an Oldowan pebble tool, the most basic of human stone tools
Fire, one of the greatest human discoveries

An Acheulean hand axe, the pinnacle of Homo erectus stone working

Venus of Willendorf, an example of Paleolithic art
 See also: Hunting hypothesis
Using tools has been interpreted as a sign of intelligence, and it has been theorized that tool use may
have stimulated certain aspects of human evolution—most notably the continued expansion of the human
brain. Paleontology has yet to explain the expansion of this organ over millions of years despite being
extremely demanding in terms of energy consumption. The brain of a modern human consumes about 20
watts (400 kilocalories per day), which is one fifth of the energy consumption of a human body. Increased tool
use would allow hunting for energy-rich meat products, and would enable processing more energy-rich plant
products. Researchers have suggested that early hominids were thus under evolutionary pressure to
increase their capacity to create and use tools.[50]
Precisely when early humans started to use tools is difficult to determine, because the more primitive
these tools are (for example, sharp-edged stones) the more difficult it is to decide whether they are natural
objects or human artifacts. There is some evidence that the australopithecines (4 Ma) may have used broken
bones as tools, but this is debated.
It should be noted that many species make and use tools, but it is the human species that dominates
the areas of making and using more complex tools. The oldest known tools are the "Oldowan stone tools"
from Ethiopia. It was discovered that these tools are from 2.5 to 2.6 million years old, which predates the
earliest known "Homo" species. There is no known evidence that any "Homo" specimens appeared by 2.5
Ma. A Homo fossil was found near some Oldowan tools, and its age was noted at 2.3 million years old,
suggesting that maybe the Homo species did indeed create and use these tools. It is surely possible, but not
solid evidence. Bernard Wood noted that "Paranthropus" coexisted with the early Homo species in the area
of the "Oldowan Industrial Complex" over roughly the same span of time. Although there is no direct evidence
that points to Paranthropus as the tool makers, their anatomy lends to indirect evidence of their capabilities in
this area. Most paleoanthropologists agree that the early "Homo" species were indeed responsible for most
of the Oldowan tools found. They argue that when most of the Oldowan tools were found in association with
human fossils, Homo was always present, but Paranthropus was not.[51]
In 1994, Randall Susman used the anatomy of opposable thumbs as the basis for his argument that
both the Homo and Paranthropus species were toolmakers. He compared bones and muscles of human and
chimpanzee thumbs, finding that humans have 3 muscles that chimps lack. Humans also have thicker
metacarpals with broader heads, making the human hand more successful at precision grasping than the
chimpanzee hand. Susman defended that modern anatomy of the human thumb is an evolutionary response
to the requirements associated with making and handling tools and that both species were indeed
toolmakers.[51]

[edit] Stone tools

Stone tools are first attested around 2.6 Ma, when H. habilis in Eastern Africa used so-called pebble
tools, choppers made out of round pebbles that had been split by simple strikes.[52] This marks the
beginning of the Paleolithic, or Old Stone Age; its end is taken to be the end of the last Ice Age, around
10,000 years ago. The Paleolithic is subdivided into the Lower Paleolithic (Early Stone Age, ending around
350,000–300,000 years ago), the Middle Paleolithic (Middle Stone Age, until 50,000–30,000 years ago), and
the Upper Paleolithic.
The period from 700,000–300,000 years ago is also known as the Acheulean, when H. ergaster (or
erectus) made large stone hand-axes out of flint and quartzite, at first quite rough (Early Acheulian), later
"retouched" by additional, more subtle strikes at the sides of the flakes. After 350,000 BP (Before Present)
the more refined so-called Levallois technique was developed. It consisted of a series of consecutive strikes,
by which scrapers, slicers ("racloirs"), needles, and flattened needles were made.[52] Finally, after about
50,000 BP, ever more refined and specialized flint tools were made by the Neanderthals and the immigrant
Cro-Magnons (knives, blades, skimmers). In this period they also started to make tools out of bone.

[edit] Modern humans and the "Great Leap Forward" debate

See also: Behavioral modernity
Until about 50,000–40,000 years ago the use of stone tools seems to have progressed stepwise.
Each phase (H. habilis, H. ergaster, H. neanderthalensis) started at a higher level than the previous one, but
once that phase started further development was slow. These Homo species were culturally conservative,
but after 50,000 BC modern human culture started to change at a much greater speed. Jared Diamond,
author of The Third Chimpanzee, and some anthropologists characterize this as a "Great Leap Forward".
Modern humans started burying their dead, making clothing out of hides, developing sophisticated
hunting techniques (such as using trapping pits or driving animals off cliffs), and engaging in cave painting.
[53] As human culture advanced, different populations of humans introduced novelty to existing technologies:
artifacts such as fish hooks, buttons and bone needles show signs of variation among different populations of
humans, something that had not been seen in human cultures prior to 50,000 BP. Typically, H.
neanderthalensis populations do not vary in their technologies.
Among concrete examples of Modern human behavior, anthropologists include specialization of
tools, use of jewellery and images (such as cave drawings), organization of living space, rituals (for example,
burials with grave gifts), specialized hunting techniques, exploration of less hospitable geographical areas,
and barter trade networks. Debate continues as to whether a "revolution" led to modern humans ("the big
bang of human consciousness"), or whether the evolution was more gradual.[54]
[edit] Models of human evolution
Today, all humans belong to one, undivided by species barrier, population of Homo sapiens sapiens.
However, according to the "Out of Africa" model this is not the first species of hominids: the first species of
genus Homo, Homo habilis, evolved in East Africa at least 2 Ma, and members of this species populated
different parts of Africa in a relatively short time. Homo erectus evolved more than 1.8 Ma, and by 1.5 Ma had
spread throughout the Old World.
Anthropologists have been divided as to whether current human population evolved as one
interconnected population (as postulated by the Multiregional Evolution hypothesis), or evolved only in East
Africa, speciated, and then migrating out of Africa and replaced human populations in Eurasia (called the
"Out of Africa" Model or the "Complete Replacement" Model).

[edit] Multiregional model

Main article: Multiregional hypothesis
Multiregional evolution, a model to account for the pattern of human evolution , was proposed by
Milford H. Wolpoff[55] in 1988.[56] Multiregional evolution holds that human evolution from the beginning of
the Pleistocene 2.5 million years BP to the present day has been within a single, continuous human species,
evolving worldwide to modern Homo sapiens.
According to the multiregional hypothesis, fossil and genomic data are evidence for worldwide
human evolution and contradict the recent speciation postulated by the Recent African origin hypothesis. The
fossil evidence was insufficient for Richard Leakey to resolve this debate.[57] Studies of haplogroups in Y-
chromosomal DNA and mitochondrial DNA have largely supported a recent African origin.[58] Evidence from
autosomal DNA also supports the Recent African origin. However the presence of archaic admixture in
modern humans remains a possibility and has been suggested by some studies.[59]

[edit] Out of Africa

See also: Recent single origin hypothesis and Early human migrations
According to the Out of Africa model, developed by Chris Stringer and Peter Andrews, modern H.
sapiens evolved in Africa 200,000 years ago. Homo sapiens began migrating from Africa between 70,000 –
50,000 years ago and eventually replaced existing hominid species in Europe and Asia.[60][61] Out of Africa
has gained support from research using mitochondrial DNA (mtDNA). After analysing genealogy trees
constructed using 133 types of mtDNA, researchers concluded that all were descended from a woman from
Africa, dubbed Mitochondrial Eve. Out of Africa is also supported by the fact that mitochondrial genetic
diversity is highest among African populations.[62]
There are differing theories on whether there was a single exodus or several. A multiple dispersal
model involves the Southern Dispersal theory,[63] which has gained support in recent years from genetic,
linguistic and archaeological evidence. In this theory, there was a coastal dispersal of modern humans from
the Horn of Africa around 70,000 years ago. This group helped to populate Southeast Asia and Oceania,
explaining the discovery of early human sites in these areas much earlier than those in the Levant. A second
wave of humans dispersed across the Sinai peninsula into Asia, resulting in the bulk of human population for
Eurasia. This second group possessed a more sophisticated tool technology and was less dependent on
coastal food sources than the original group. Much of the evidence for the first group's expansion would have
been destroyed by the rising sea levels at the end of the Holocene era.[63] The multiple dispersal model is
contradicted by studies indicating that the populations of Eurasia and the populations of Southeast Asia and
Oceania are all descended from the same mitochondrial DNA lineages, which support a single migration out
of Africa that gave rise to all non-African populations.[64]
The broad study of African genetic diversity headed by Dr. Sarah Tishkoff found the San people to
express the greatest genetic diversity among the 113 distinct populations sampled, making them one of 14
"ancestral population clusters". The research also located the origin of modern human migration in south-
western Africa, near the coastal border of Namibia and Angola.[65]
According to the Toba catastrophe theory to which some anthropologists and archeologists
subscribe, the supereruption of Lake Toba on Sumatra island in Indonesia roughly 70,000 years ago had
global consequences,[66] killing most humans then alive and creating a population bottleneck that affected
the genetic inheritance of all humans today.[67]

[edit] Recent and current human evolution

See also: Transhuman
Natural selection is being observed in contemporary human populations, with recent findings
demonstrating the population which is at risk of the severe debilitating disease kuru has significant over-
representation of an immune variant of the prion protein gene G127V versus non-immune alleles. Scientists
postulate one of the reasons for the rapid selection of this genetic variant is the lethality of the disease in non-
immune persons.[68][69] Other reported evolutionary trends in other populations include a lengthening of the
reproductive period, reduction in cholesterol levels, blood glucose and blood pressure.[70]
In their 2009 book The 10,000 Year Explosion, Gregory Cochran and Henry Harpending argue that
human evolution has accelerated since and as a result of the development of agriculture and civilisation
since some 50,000 years ago, and that there are consequently substantial genetic differences between
different current human populations.

[edit] Genetics
Main articles: Human evolutionary genetics and Human genetic variation
Human evolutionary genetics studies how one human genome differs from the other, the evolutionary
past that gave rise to it, and its current effects. Differences between genomes have anthropological, medical
and forensic implications and applications. Genetic data can provide important insight into human evolution.

[edit] Notable human evolution researchers

See also: :Category:Human evolution theorists
• Robert Broom, a Scottish physician and palaeontologist whose work on South Africa led to
the discovery and description of the Paranthropus genus of hominins, and of "Mrs. Ples"
• James Burnett, Lord Monboddo, a British judge most famous today as a founder of modern
comparative historical linguistics
• Raymond Dart, an Australian anatomist and palaeoanthropologist, whose work at Taung, in
South Africa, led to the discovery of Australopithecus africanus
• Charles Darwin, a British naturalist who documented considerable evidence that species
originate through evolutionary change
• Richard Dawkins, a British ethologist, evolutionary biologist who has promoted a gene-
centered view of evolution
 • J. B. S. Haldane, a British geneticist and evolutionary biologist
• William D. Hamilton, a British Evolutionary Biologist who expounded a rigorous genetic basis
for kin selection, and on the evolution of HIV and other human diseases.
• Sir Alister Hardy, a British zoologist, who first hypothesised the aquatic ape theory of human
evolution
• Henry McHenry, an American anthropologist who specializes in studies of human evolution,
the origins of bipedality, and paleoanthropology
• Jeffrey Laitman, an American anatomist and physical anthropologist whose work has
explored the evolution of the vocal tract and speech
• Louis Leakey, an African archaeologist and naturalist whose work was important in
establishing human evolutionary development in Africa
• Mary Leakey, a British archaeologist and anthropologist whose discoveries in Africa include
the Laetoli footprints
• Richard Leakey, an African paleontologist and archaeologist, son of Louis and Mary Leakey
• Svante Pääbo, a Swedish biologist specializing in evolutionary genetics
• David Pilbeam, a paleoanthropologist, researcher and writer on a range of topics involving
human and primate evolution.
• Jeffrey H. Schwartz, an American physical anthropologist and professor of biological
anthropology
• Chris Stringer, anthropologist, leading proponent of the recent single origin hypothesis
• Alan Templeton, geneticist and statistician, proponent of the multiregional hypothesis
• Philip V. Tobias, a South African palaeoanthropologist is one of the world's leading
authorities on the evolution of humankind
 • Erik Trinkaus, a prominent American paleoanthropologist and expert on Neanderthal biology
and human evolution
• Alfred Russel Wallace, a British naturalist, sometimes called the "father of biogeography",
who independently from Charles Darwin proposed the principles of evolution of animal species
• Milford H. Wolpoff, an American paleoanthropologist who is the leading proponent of the
multiregional evolution hypothesis.

[edit] Species list

This list is in chronological order across the page by genus.
• Sahelanthropus • Australopithecus • Homo
• Sahela • Australo • Homo
nthropus pithecus habilis
tchadensis anamensis • Homo
• Orrorin • Australo rudolfensis
• Orrorin pithecus • Homo
tugenensis afarensis ergaster
• Ardipithecus • Australo • Homo
• Ardipith pithecus georgicus
ecus kadabba bahrelghazali • Homo
• Ardipith • Australo erectus
ecus ramidus pithecus • Homo
africanus cepranensis
 • Australo • Homo
pithecus garhi antecessor
• Australo • Homo
pithecus sediba heidelbergensis
• Paranthropus • Homo
• Paranthr rhodesiensis
opus • Homo
aethiopicus neanderthalensis
• Paranthr • Homo
opus boisei sapiens idaltu
• Paranthr • Homo
opus robustus sapiens (Cro-
• Kenyanthropus Magnon)
• Kenyant • Homo
hropus platyops sapiens sapiens
• Homo
floresiensis

[edit] See also

• List of human evolution
fossils
Evolutionary biology portal
• Timeline of human
 evolution
• Archaeogenetics
• Dual inheritance theory
• Dysgenics
• Evolutionary anthropology
• Evolutionary medicine
• Evolutionary neuroscience
• Evolution of human
intelligence
• Evolution of morality
• Evolutionary origin of
religions
• Evolutionary psychology
• History of Earth
• Hominid intelligence
• Human behavioral ecology
• Human skeletal changes due to
bipedalism
• Human vestigiality
• Ida (fossil)
• Pan prior
• Physical anthropology
• Sahara pump theory
• Sexual selection in human evolution
• Sociocultural evolution

[edit] References
[edit] Notes
1. ^ Heng HH (May 2009). "The genome-centric concept: resynthesis of evolutionary theory".
Bioessays 31 (5): 512–25. doi:10.1002/bies.200800182. PMID 19334004.
 2. ^ Stringer, C.B. (1994). "Evolution of early humans". in Steve Jones, Robert Martin & David
Pilbeam (eds.). The Cambridge Encyclopedia of Human Evolution . Cambridge: Cambridge University
Press. p. 242. ISBN 0-521-32370-3. Also ISBN 0-521-46786-1 (paperback)
3. ^ McHenry, H.M (2009). "Human Evolution". in Michael Ruse & Joseph Travis. Evolution:
The First Four Billion Years. Cambridge, Massachusetts: The Belknap Press of Harvard University
Press. p. 265. ISBN 978-0-674-03175-3.
4. ^ "Out of Africa Revisited - 308 (5724): 921g - Science". Sciencemag.org. 2005-05-13.
doi:10.1126/science.308.5724.921g.
http://www.sciencemag.org/cgi/content/summary/sci;308/5724/921g. Retrieved 2009-11-23.
5. ^ Nature (2003-06-12). "Access : Human evolution: Out of Ethiopia". Nature.
http://www.nature.com/nature/journal/v423/n6941/full/423692a.html. Retrieved 2009-11-23.
6. ^ "Origins of Modern Humans: Multiregional or Out of Africa?". ActionBioscience.
http://www.actionbioscience.org/evolution/johanson.html. Retrieved 2009-11-23.
7. ^ "Modern Humans - Single Origin (Out of Africa) vs Multiregional". Asa3.org.
http://www.asa3.org/ASA/education/origins/migration.htm. Retrieved 2009-11-23.
8. ^ "dhghem" ([dead link]). The American Heritage Dictionary of the English Language (4th
ed.). Houghton Mifflin Company. 2000. http://www.bartelby.org/61/roots/IE104.html.
9. ^ Darwin, Charles (1871. This edition published 1981, with Introduction by John Tyler Bonner
& Robert M. May). The Descent of Man, and Selection in Relation to Sex . Princeton, New Jersey:
Princeton University Press. ISBN 0-691-02369-7..
10.^ Kordos L, Begun DR (2001). "Primates from Rudabánya: allocation of specimens to
individuals, sex and age categories". J. Hum. Evol. 40 (1): 17–39. doi:10.1006/jhev.2000.0437.
PMID 11139358.
 11.^ Zalmout, I.S.; Sanders, W.J.; MacLatchy, L.M.; Gunnell, G.F.; Al-Mufarreh, Y.A.; Ali, M.A.;
Nasser, A.-A.H.; Al-Masari, A.M. et al. (2010). "New Oligocene primate from Saudi Arabia and the
divergence of apes and Old World Monkeys". Nature 466 (7304): 360–364. doi:10.1038/nature09094.
PMID 20631798.
12.^ Strait DS, Grine FE, Moniz MA (1997). "A reappraisal of early hominid phylogeny". J. Hum.
Evol. 32 (1): 17–82. doi:10.1006/jhev.1996.0097. PMID 9034954.
13.^ Walker, Alan (2006). "Early Hominin Diets: Overview and Historical Perspectives". in
Ungar, Peter S.. Evolution of the Human Diet: The Known, the Unknown, and the Unknowable . US:
Oxford University Press. pp. 3–10. ISBN 0195183460. http://books.google.com/?
id=6mxZ1hNBHgkC&pg=PA357&lpg=PA357&dq=
%22The+Calcaneus+of+Australopithecus+afarensis+and+its+implications+for+the+Evolution+of+Bip
edality%22&q=%22The%20Calcaneus%20of%20Australopithecus%20afarensis%20and%20its
%20implications%20for%20the%20Evolution%20of%20Bipedality%22. (scroll up to view chapter 1
& part of chapter 2, which is a serendipitous result from another search. Subsequent attempts get a
targeted search result gave returns without chapter 1).
14.^ Ungar, Peter S. & Teaford, Mark F. (2002). Human Diet: Its Origin and Evolution. Westport,
CT: Bergin & Garvey. p. 206. ISBN 0897897366.
15.^ Bogin, Barry (1997). "The evolution of human nutrition". in Romanucci-Ross, Lola;
Moerman, Daniel E.; & Tancredi, Laurence R.. The Anthropology of Medicine: From Culture to
Method (3 ed.). South Hadley, Mass.: Bergen and Garvey. pp. 96–142. ISBN 0897895169.
http://web.archive.org/web/20031203003838/http://citd.scar.utoronto.ca/ANTAO1/Projects/Bogin.htm
l.
16.^ Barnicot NA (2005, April/June). "Human nutrition: evolutionary perspectives". Integr Physiol
Behav Sci 40 (2): 114–17. doi:10.1007/BF02734246. PMID 17393680.
 17.^ Leonard WR, Snodgrass JJ, Robertson ML (2007). "Effects of brain evolution on human
nutrition and metabolism" (PDF). Annu Rev Nutr. 27: 311–27.
doi:10.1146/annurev.nutr.27.061406.093659. PMID 17439362.
http://www.pinniped.net/LeonardARN.pdf. Retrieved 2008-12-29.
18.^ Wood, B. & Collard, M. (1999) The changing face of Genus Homo. Evol. Anth. 8(6) 195-207
19.^ Wood B (1999). "'Homo rudolfensis' Alexeev, 1986-fact or phantom?". J. Hum. Evol. 36 (1):
115–8. doi:10.1006/jhev.1998.0246. PMID 9924136.
20.^ Gabounia L. de Lumley M. Vekua A. Lordkipanidze D. de Lumley H. (2002). "Discovery of a
new hominid at Dmanisi (Transcaucasia, Georgia)". Comptes Rendus Palevol, 1 (4): 243–53.
doi:10.1016/S1631-0683(02)00032-5.
21.^ Lordkipanidze D, Vekua A, Ferring R, et al. (2006). "A fourth hominin skull from Dmanisi,
Georgia". The anatomical record. Part A, Discoveries in molecular, cellular, and evolutionary biology
288 (11): 1146–57. doi:10.1002/ar.a.20379. PMID 17031841.
22.^ Genetic Analysis of Lice Supports Direct Contact between Modern and Archaic Humans
Reed DL, Smith VS, Hammond SL, Rogers AR, Clayton DH PLoS Biology Vol. 2, No. 11, e340
doi:10.1371/journal.pbio.0020340 http://biology.plosjournals.org/perlserv/?
request=slideshow&type=figure&doi=10.1371/journal.pbio.0020340&id=15540
23.^ Turner W (1895). "On M. Dubois' Description of Remains recently found in Java, named by
him Pithecanthropus erectus: With Remarks on so-called Transitional Forms between Apes and
Man". Journal of anatomy and physiology 29 (Pt 3): 424–45. PMID 17232143.
24.^ Spoor F, Wood B, Zonneveld F (1994). "Implications of early hominid labyrinthine
morphology for evolution of human bipedal locomotion". Nature 369 (6482): 645–8.
doi:10.1038/369645a0. PMID 8208290.
 25.^ Bermúdez de Castro JM, Arsuaga JL, Carbonell E, Rosas A, Martínez I, Mosquera M
(1997). "A hominid from the lower Pleistocene of Atapuerca, Spain: possible ancestor to Neandertals
and modern humans". Science 276 (5317): 1392–5. doi:10.1126/science.276.5317.1392.
PMID 9162001.
26.^ Carbonell, Eudald; José M. Bermúdez de Castro et al. (2008-03-27). "The first hominin of
Europe". Nature 452 (7186): 465–469. doi:10.1038/nature06815. PMID 18368116.
http://www.nature.com/nature/journal/v452/n7186/full/nature06815.html. Retrieved 2008-03-26.
27.^ Manzi G, Mallegni F, Ascenzi A (2001). "A cranium for the earliest Europeans: phylogenetic
position of the hominid from Ceprano, Italy". Proc. Natl. Acad. Sci. U.S.A. 98 (17): 10011–6.
doi:10.1073/pnas.151259998. PMID 11504953.
28.^ Czarnetzki, A (2003). "Palaeopathological and variant conditions of the Homo
heidelbergensis type specimen (Mauer, Germany)". Journal of Human Evolution 44: 479.
doi:10.1016/S0047-2484(03)00029-0.
29.^ Indiana University (March 27, 2006). "Scientists discover hominid cranium in Ethiopia".
Press release. http://newsinfo.iu.edu/news/page/normal/3142.html. Retrieved 2006-11-26.
30.^ Herrera, K. J.; Somarelli, J. A.; Lowery, R. K.; Herrera, R. J. (2009). "To what extent did
Neanderthals and modern humans interact?". Biological Reviews 84 (2): 245–257.
doi:10.1111/j.1469-185X.2008.00071.x. PMID 19391204. edit
31.^ Harvati K (2003). "The Neanderthal taxonomic position: models of intra- and inter-specific
craniofacial variation". J. Hum. Evol. 44 (1): 107–32. doi:10.1016/S0047-2484(02)00208-7.
PMID 12604307.
32.^ Krings M, Stone A, Schmitz RW, Krainitzki H, Stoneking M, Pääbo S (1997). "Neandertal
DNA sequences and the origin of modern humans". Cell 90 (1): 19–30. doi:10.1016/S0092-
8674(00)80310-4. PMID 9230299.
 33.^ Green RE, et al. (2008). "A Complete Neandertal Mitochondrial Genome Sequence
Determined by High-Throughput Sequencing". Cell 134 (3): 416–426. doi:10.1016/j.cell.2008.06.021.
PMID 18692465.
34.^ Serre D, Langaney A, Chech M, et al. (2004). "No evidence of Neandertal mtDNA
contribution to early modern humans". PLoS Biol. 2 (3): E57. doi:10.1371/journal.pbio.0020057.
PMID 15024415.
35.^ a b c Jennifer Viegas (2010-05-06). "Neanderthals, humans interbred, DNA proves".
Discovery News. http://news.discovery.com/human/neanderthal-human-interbreed-dna.html.
Retrieved 2010-08-17.
36.^ Gutiérrez G, Sánchez D, Marín A (2002). "A reanalysis of the ancient mitochondrial DNA
sequences recovered from Neandertal bones". Mol. Biol. Evol. 19 (8): 1359–66. PMID 12140248.
37.^ Hebsgaard MB, Wiuf C, Gilbert MT, Glenner H, Willerslev E (2007). "Evaluating
Neanderthal genetics and phylogeny". J. Mol. Evol. 64 (1): 50–60. doi:10.1007/s00239-006-0017-y.
PMID 17146600.
38.^ Diamond, Jared (1992). The Third Chimpanzee: The Evolution and Future of the Human
Animal. Harper Perennial. ISBN 0060984031.
39.^ How Neanderthals met a grisly fate: devoured by humans. The Observer. May 17, 2009.
40.^ "DNA identifies new ancient human dubbed 'X-woman'". BBC News. March 25, 2010.
41.^ Supervolcanoes, BBC2, 3 February 2000
42.^ Stanley H. Ambrose (1998). "Late Pleistocene human population bottlenecks, volcanic
winter, and differentiation of modern humans". Journal of Human Evolution 34 (6): 623–651.
doi:10.1006/jhev.1998.0219. PMID 9650103.
43.^ Ambrose, Stanley H. (2005). "Volcanic Winter, and Differentiation of Modern Humans".
Bradshaw Foundation. http://www.bradshawfoundation.com/evolution/. Retrieved 2006-04-08.
 44.^ Brown P, Sutikna T, Morwood MJ, et al. (2004). "A new small-bodied hominin from the Late
Pleistocene of Flores, Indonesia". Nature 431 (7012): 1055–61. doi:10.1038/nature02999.
PMID 15514638.
45.^ Argue D, Donlon D, Groves C, Wright R (2006). "Homo floresiensis: microcephalic,
pygmoid, Australopithecus, or Homo?". J. Hum. Evol. 51 (4): 360–74.
doi:10.1016/j.jhevol.2006.04.013. PMID 16919706.
46.^ a b Martin RD, Maclarnon AM, Phillips JL, Dobyns WB (2006). "Flores hominid: new
species or microcephalic dwarf?". The anatomical record. Part A, Discoveries in molecular, cellular,
and evolutionary biology 288 (11): 1123–45. doi:10.1002/ar.a.20389. PMID 17031806.
47.^ a b Sample, Ian (March 24, 2010). "New species of human ancestor found in Siberia". The
Guardian. http://www.guardian.co.uk/science/2010/mar/24/new-human-species-siberia
48.^ Krause, Johannes; Fu, Qiaomei; Good, Jeffrey M.; Viola, Bence; Shunkov, Michael V.;
Derevianko, Anatoli P.; Pääbo, Svante (2010). "The complete mitochondrial DNA genome of an
unknown hominin from southern Siberia". Nature 464 (7290): 894–897. doi:10.1038/nature08976.
PMID 20336068
49.^ a b Katsnelson, Alla (March 24, 2010). "New hominin found via mtDNA". The Scientist.
http://www.the-scientist.com/blog/display/57254/#ixzz0j820ioz1
50.^ Gibbons, Ann (1998). "Solving the Brain's Energy Crisis". Science 280 (5368): 1345–47.
doi:10.1126/science.280.5368.1345. PMID 9634409.
51.^ a b Freeman, Scott; Jon C. Herron. Evolutionary Analysis (4th ed.)., Pearson Education,
Inc. (2007). ISBN 0-13-227584-8 pages 786-788
 52.^ a b Plummer T (2004). "Flaked stones and old bones: Biological and cultural evolution at
the dawn of technology". Am. J. Phys. Anthropol. Suppl 39: 118–64. doi:10.1002/ajpa.20157.
PMID 15605391.
53.^ Ambrose SH (2001). "Paleolithic technology and human evolution". Science 291 (5509):
1748–53. doi:10.1126/science.1059487. PMID 11249821.
54.^ Mcbrearty S, Brooks AS (2000). "The revolution that wasn't: a new interpretation of the
origin of modern human behavior". J. Hum. Evol. 39 (5): 453–563. doi:10.1006/jhev.2000.0435.
PMID 11102266.
55.^ Wolpoff, MH; Hawks J, Caspari R (2000). "Multiregional, not multiple origins". Am J Phys
Anthropol 112 (1): 129–36. doi:10.1002/(SICI)1096-8644(200005)112:1<129::AID-AJPA11>3.0.CO;2-
K. PMID 10766948. http://www3.interscience.wiley.com/journal/71008905/abstract.
56.^ Wolpoff, MH; JN Spuhler, FH Smith, J Radovcic, G Pope, DW Frayer, R Eckhardt, and G
Clark (1988). "Modern Human Origins". Science 241 (4867): 772–4. doi:10.1126/science.3136545.
PMID 3136545. http://www.sciencemag.org/cgi/pdf_extract/241/4867/772.
57.^ Leakey, Richard (1994). The Origin of Humankind. Science Masters Series. New York, NY:
Basic Books. pp. 87–89. ISBN 0465053130.
58.^ Jorde LB, Bamshad M, Rogers AR (February 1998). "Using mitochondrial and nuclear DNA
markers to reconstruct human evolution". Bioessays 20 (2): 126–36. doi:10.1002/(SICI)1521-
1878(199802)20:2<126::AID-BIES5>3.0.CO;2-R. PMID 9631658.
59.^ Wall, J. D.; Lohmueller, K. E.; Plagnol, V. (2009). "Detecting Ancient Admixture and
Estimating Demographic Parameters in Multiple Human Populations". Molecular Biology and
Evolution 26 (8): 1823. doi:10.1093/molbev/msp096. PMID 19420049.
60.^ Modern Humans Came Out of Africa, "Definitive" Study Says
 61.^ Stringer CB, Andrews P (March 1988). "Genetic and fossil evidence for the origin of
modern humans". Science 239 (4845): 1263–8. doi:10.1126/science.3125610. PMID 3125610.
62.^ Cann RL, Stoneking M, Wilson AC (1987). "Mitochondrial DNA and human evolution".
Nature 325 (6099): 31–6. doi:10.1038/325031a0. PMID 3025745.
http://artsci.wustl.edu/~landc/html/cann/.
63.^ a b Searching for traces of the Southern Dispersal, by Dr. Marta Mirazón Lahr, et al.
64.^ Macaulay, V.; Hill, C; Achilli, A; Rengo, C; Clarke, D; Meehan, W; Blackburn, J; Semino, O
et al. (2005). "Single, Rapid Coastal Settlement of Asia Revealed by Analysis of Complete
Mitochondrial Genomes". Science 308 (5724): 1034. doi:10.1126/science.1109792. PMID 15890885.
http://www.sciencemag.org/cgi/content/abstract/308/5724/1034.
65.^ BBC World News "Africa's genetic secrets unlocked", 1 May 2009; the results were
published in the online edition of the journal Science.
66.^ "The new batch - 150,000 years ago". BBC - Science & Nature - The evolution of man.
67.^ "When humans faced extinction". BBC. 2003-06-09.
http://news.bbc.co.uk/2/hi/science/nature/2975862.stm. Retrieved 2007-01-05.
68.^ Medical Research Council (UK) ((November 21, 2009)). "Brain Disease 'Resistance Gene'
Evolves in Papua New Guinea Community; Could Offer Insights Into CJD". Science Daily (online)
(Science News). http://www.sciencedaily.com/releases/2009/11/091120091959.htm. Retrieved 2009-
11-22.
69.^ Mead, S.; Whitfield, J.; Poulter, M.; Shah, P.; Uphill, J.; Campbell, T.; Al-Dujaily, H.;
Hummerich, H. et al. (2009). "A Novel Protective Prion Protein Variant that Colocalizes with Kuru
Exposure.". The New England journal of medicine 361 (21): 2056–2065.
doi:10.1056/NEJMoa0809716. PMID 19923577. edit
 70.^ Byars, S. G.; Ewbank, D.; Govindaraju, D. R.; Stearns, S. C. (2009). "Evolution in Health
and Medicine Sackler Colloquium: Natural selection in a contemporary human population".
Proceedings of the National Academy of Sciences 107: 1787. doi:10.1073/pnas.0906199106. edit

[edit] Further reading

Main article: Bibliography of evolution and human behavior
• Hill, Andrew; Ward, Steven (1988). "Origin of the hominidae: The record of african large
hominoid evolution between 14 my and 4 my". Yearbook of Physical Anthropology 31 (59): 49–83.
doi:10.1002/ajpa.1330310505
• Alexander, R. D. (1990). "How Did Humans Evolve? Reflections on the Uniquely Unique
Species". University of Michigan Museum of Zoology Special Publication (University of Michigan
Museum of Zoology) (1): 1–38. http://insects.ummz.lsa.umich.edu/pdfs/Alexander1990.pdf.
• Flinn, M. V., Geary, D. C., & Ward, C. V. (2005). Ecological dominance, social competition,
and coalitionary arms races: Why humans evolved extraordinary intelligence. Evolution and Human
Behavior, 26, 10-46. Full text.PDF (345 KB)
• edited by Steve Jones, Robert Martin, and David Pilbeam ; foreword by Richard Dawkins.
(1994). Jones, S., Martin, R., & Pilbeam, D.. ed. The Cambridge Encyclopedia of Human Evolution .
Cambridge: Cambridge University Press. ISBN 0-521-32370-3. Also ISBN 0-521-46786-1
• Wolfgang Enard et al. (2002-08-22). "Molecular evolution of FOXP2, a gene involved in
speech and language". Nature 418: 870.
• DNA Shows Neandertals Were Not Our Ancestors
 • J. W. IJdo, A. Baldini, D. C. Ward, S. T. Reeders, R. A. Wells (October 1991). "Origin of
human chromosome 2: An ancestral telomere-telomere fusion" (PDF). Genetics 88: 9051–9055.
http://www.pnas.org/cgi/reprint/88/20/9051.pdf. —two ancestral ape chromosomes fused to give rise
to human chromosome 2.
• Ovchinnikov, et al.; Götherström, Anders; Romanova, Galina P.; Kharitonov, Vitaliy M.;
Lidén, Kerstin; Goodwin, William (2000). "Molecular analysis of Neanderthal DNA from the Northern
Caucasus". Nature 404 (6777): 490. doi:10.1038/35006625. PMID 10761915.
• Heizmann, Elmar P J, Begun, David R (2001). "The oldest Eurasian hominoid". Journal of
Human Evolution 41 (5): 463. doi:10.1006/jhev.2001.0495. PMID 11681862.
• BBC: Finds test human origins theory. 2007-08-08 Homo habilis and Homo erectus are sister
species that overlapped in time.

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 W000

Histoire évolutive des homininés

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

L'histoire évolutive de la lignée humaine retrace les étapes de l'évolution de la famille des homininés
au cours des temps géologiques. Elle ne se limite donc pas à l'étude du genre Homo mais inclut plus
généralement tous les membres de la famille des homininés (y compris donc les australopithèques qui ne
sont pas, à proprement parler, nos ancêtres). Son étude revient surtout à décrire l'histoire de la lignée
humaine. Ce travail scientifique correspond à ce qu'on appelle la paléoanthropologie.
Sommaire
[masquer]
• 1 Du début de la Vie au premier bipède
• 1.1 De l'apparition de la vie à la sortie des eaux
• 1.2 La radiation évolutive des mammifères
• 2 La séparation de la lignée du chimpanzé et de la lignée
humaine
• 2.1 Le plus ancien primate bipède
• 3 Évolution de la lignée humaine
• 3.1 Une évolution plus « buissonnante » que
linéaire
• 3.2 Évolution des hominines
• 3.3 L'émergence du genre Homo
• 3.4 Les origines de l'homme moderne
• 3.5 Fossiles
• 4 Caractéristiques de la lignée humaine
• 4.1 Bipédie
• 4.2 L'évolution crânienne
• 4.3 Activités culturelles
• 4.4 Évolution technique et sociale
• 5 Histoire de la théorie de l'évolution humaine
Du début de la Vie au premier bipède [modifier]

Les bactéries: des « cousines » très éloignées

Un des premiers tétrapodes : ichthyostega

De l'apparition de la vie à la sortie des eaux [modifier]

Il y a 3,5 milliards d'années, la vie apparaît sur Terre. Suivent des formes simples, probablement
assez proches de certaines bactéries actuelles. Ensuite la vie a pris certaines formes plus complexes, toutes
aquatiques, dont les expressions les plus complexes ressemblaient aux poissons osseux actuels. C'est de
cette branche (ostéichtyens) que proviendraient les premiers vertébrés terrestres, les plantes et les
arthropodes étant déjà installées hors de l'eau.
Les premiers animaux à quatre pattes connus ou tétrapodes sont des amphibiens tel Ichthyostega.
Le rayonnement d'espèces issues de cette souche a été important, les « reptiles » de la classification
traditionnelle en sont une variante. Une lignée relativement primitive de reptiles, les reptiles mammaliens, est
à l'origine des mammifères.

La radiation évolutive des mammifères [modifier]

Articles détaillés : Histoire évolutive des mammifères et Histoire évolutive des primates.
Les mammifères subissent une radiation évolutive dès le début du cénozoïque: de nombreuses
espèces différentes apparaissent et occupent des milieux variés : les airs, les milieux aquatiques… Des
mammifères arboricoles apparaissent, les primates. Parmi eux, un grand nombre sont frugivores et c'est
d'ailleurs parallèlement à l'acquisition de ce régime frugivore que devient possible la vision en couleur, pour
le repérage des fruits mûrs [1]. Parmi eux, les singes (simiens) puis les grands singes, se seraient diversifiés,
ces derniers n'ayant pas de queue préhensile.
La classification phylogénétique permet aujourd'hui de positionner de façon précise la lignée
humaine par rapport aux huit genres non-éteints de singes qui forment la super-famille des Hominoïdes :
• le regroupement des diverses espèces d'hommes (genre Homo, à 46 chromosomes) avec
les diverses espèces d'australopithèques forme la tribu des Hominines,
• le regroupement de la tribu des Hominines avec celle des Panines (genre Pan, à 48
chromosomes) forme la sous-famille des Homininés,
 • le regroupement de la sous-famille des Homininés avec celle des Gorillinés (genre Gorilla, à
48 chromosomes) forme la famille des Hominidés,
• le regroupement de la famille des Hominidés avec celles des Pongidés (famille maintenant
limitée au seul genre Pongo, à 48 chromosomes, comprenant l'espèce orang outan) [2] et des
Hylobatidés (quatre genres : le genre Hylobates, à 44 chromosomes, qui comprend les espèces de
gibbons, le genre Hoolock, à 38 chromosomes, le genre Nomascus, à 52 chromosomes, et le genre
Symphalangus, à 50 chromosomes) forme la super-famille des Hominoïdes ;
À un degré plus élevé dans la phylogénie :
• le regroupement de la super-famille des Hominoïdes avec celle des Cercopithécoïdes forme
le groupe zoologique des Catarhiniens (singes du Vieux-Monde),
• le regroupement des Catarhiniens avec les Platyrhiniens (singes du Nouveau-Monde) forme
l'infra-ordre des Simiiformes (singes),
• ensuite, le regroupement des Simiiformes avec les Tarsiformes forme le sous-ordre des
Haplorhiniens,
• le regroupement du sous-ordre des Haplorhiniens avec celui des Strepsirhiniens (qui
comprend les infra-ordres des Lorisiformes et des Lémuriformes) forme l'ordre des Primates,
On remonte alors à la classe des Mammifères ; on peut ensuite poursuivre jusqu'au sous-
embranchement des Vertébrés, à l'embranchement des Cordés, au sous-règne des Eumétazoaires, au
règne animal, enfin à l'empire eucaryote.
Le regroupement de l'empire eucaryote avec les empires eubacteria et archaea constitue la matière
vivante terrestre. Ces empires résultent de la première bifurcation survenue lors de l'évolution de la matière
vivante sur Terre, mais les frontières précises de ceux-ci ne font pas encore l'unanimité parmi les
biologistes ; la succession de bifurcations qui mènent de l'empire eucaryote au règne animal est
actuellement inconnue.

La séparation de la lignée du chimpanzé et de la lignée humaine [modifier]

Les chimpanzés sont les êtres les plus proches des humains, mais pas nos ancêtres
Les différentes séparations entre les lignées ayant mené aux différentes espèces de singes actuels,
et au genre Homo, se sont produites de manière successive. La séparation la plus récente entre la lignée
humaine et celle d'une autre espèce de singes a été celle qui s'est produite avec les Panines (lignée des
chimpanzés). Selon David Reich de la Harvard Medical School à Boston, qui a comparé l'ADN des
chromosomes X humain et de chimpanzé, cette séparation s'est faite il y a moins de 6,3 millions d'années[3].
Toutefois, ces travaux indiquent également que cette séparation a été progressive, car la comparaison des
séquences des chromosomes X de l'Homo sapiens et du chimpanzé montre des similitudes qui semblent
refléter une période de "ré-hybridation" entre des Hominines et des Panines. Une hybridation significative
entre au moins une espèce de chimpanzé d'une part, des espèces d'australopithèques et probablement des
espèces d'Hommes d'autre part, conduisant à des échanges de gènes entre les deux tribus, a dû exister
pendant peut-être quatre millions d'années selon les auteurs de ces travaux.
Article détaillé : Histoire évolutive des primates#comparaison génétique entre humains et
chimpanzés.

Le plus ancien primate bipède [modifier]

Reconstitution de Toumaï
Au Miocène, entre -10 et -6 Ma, il y a séparation de la lignée humaine et de la lignée des
chimpanzés, espèce la plus proche de nous.
 Le plus ancien fossile de primate bipède, daté de - 7 Ma, a été découvert au Tchad en 2001. Il s'agit
de l'espèce baptisée Sahelanthropus tchadensis, aussi connue sous le nom de Toumaï. Seul le crâne a pu
être étudié. Celui-ci présente, pour certains paléoanthropologues, les caractéristiques d'une station droite, et
donc d'une marche sur deux jambes. La bipédie est reconnue par la majorité des auteurs comme le premier
caractère humain à apparaître dans la lignée humaine. Cette apparition serait l'événement fondateur ayant
permis par la suite les évolutions qui ont mené aux différentes espèces d'hominines dont l'Homme fait partie.
Cette évolution récente, depuis notre ancêtre commun le plus récent avec les chimpanzés, est
relativement bien documentée grâce aux fossiles, bien que des lacunes importantes subsistent.

Évolution de la lignée humaine [modifier]

 Évolution de la lignée humaine par rapport à celle des chimpanzés
La lignée humaine regroupe l'espèce humaine et toutes autres espèces fossiles, ayant une parenté
plus proche avec les humains, qu'avec les chimpanzés, espèce actuelle la plus proche de nous. Un fossile
ne peut être classé dans la lignée humaine que s'il présente au moins un des caractères dérivés propres à
l'espèce humaine actuelle.

Une évolution plus « buissonnante » que linéaire [modifier]

Depuis les années quatre-vingt, les découvertes de gisements de fossiles se sont multipliées, et
avec elles, le nombre d'espèces ou de sous-espèces du genre Homo.
Du même coup, l'histoire évolutionnaire de l'homme est passée d'un arbre linéaire à un arbre à
plusieurs branches, et des espèces que nous pensions être nos ancêtres il y a encore peu sont brusquement
devenues nos défunts cousins.
Cet article se propose donc de présenter l'état des théories actuellement admises, ainsi que
quelques théories alternatives.

Évolution des hominines [modifier]

Jusqu'au début des années 1980, l'état de la recherche permettait de représenter l'évolution des
hommes au Pliocène et au Pléistocène comme suit :
Vue linéaire ancienne de l'évolution des hominines
Les connaissances de l'époque permettaient de penser que l'arbre évolutif des genres Australopithecus et
Homo était linéaire et que les espèces se succédaient dans un processus continu et régulier, chaque espèce
étant l'ancêtre de l'autre. Cette hypothèse a connu son apogée dans les années 1960-1970, époque de forte
influence de la Théorie synthétique de l'évolution (TSE) dans les différentes disciplines de la
paléoanthropologie. Certains chercheurs défendaient même avec force la théorie de l'espèce unique : à une
époque donnée ne pouvait exister qu'une seule espèce d'hominidé. L'arbre évolutif de l'homme était alors
perçu comme « un gros tronc avec très peu de branches ».
Cette théorie simpliste est parfois encore enseignée de nos jours bien qu'elle n'intègre pas les
découvertes de ces dernières années. On sait par exemple que vers 2 millions d'années vivaient dans les
mêmes régions d'Afrique de l'Est des Paranthropus, des Homo rudolfensis et des Homo habilis.

L'émergence du genre Homo [modifier]

Les plus anciens représentants du genre Homo, les Homo habilis et les Homo rudolfensis, semblent
apparaître en Afrique vers 2,5 Ma, alors que des changements climatiques importants ont lieu : la formation
de la calotte glaciaire arctique entraîne une sécheresse en Afrique, ce qui provoque le recul des forêts.
Le genre Homo se définit par une capacité crânienne plus forte (supérieure à 600 cm³) que celle des
australopithèques, une boîte crânienne plus arrondie, une réduction de l'appareil masticateur et de la face,
de petites canines, et une bipédie quasi exclusive.
 Homo habilis était encore adapté à la vie arboricole. Les empreintes visibles sur les os de la boîte
crânienne prouvent qu'il existait déjà une asymétrie entre les cerveaux droit et gauche, ce qui suggère une
plus grande capacité à fabriquer et à utiliser les outils. Il est probablement l'auteur de certains des premiers
galets taillés (industrie oldowayenne).
Homo rudolfensis était plus corpulent et possédait un gros cerveau et des mâchoires plus puissantes
(à mettre en relation avec un régime alimentaire probablement plus carnivore que Homo habilis). Bien qu'il
soit peu connu, on pense que sa bipédie devait être plus évoluée que celle d' Homo habilis.
Homo habilis et Homo rudolfensis disparaissent vers 1,6 Ma.
Contemporain des derniers Homo habilis et Paranthropus, l’Homo ergaster possède des
caractéristiques qui le rapprochent beaucoup de l'homme moderne : taille plus importante, bipédie exclusive,
forte capacité crânienne (supérieure à 800 cm³), boîte crânienne bien arrondie et qui domine la face, face
réduite, etc. Ses outils commencent à être plus sophistiqués : bifaces, hachereaux, etc. (industrie
acheuléenne).
Sa bonne adaptation à la marche bipède et à la course lui permet de parcourir de grandes distances
et il va progressivement occuper une partie de l'ancien monde (Asie et Europe), dès 1,8 Ma, probablement
en suivant ses proies au gré des changements climatiques. Ses nouveaux outils lui permettent d'adopter un
régime alimentaire contenant beaucoup plus de viande (c'est un vrai chasseur). Enfin, il peut communiquer
en pratiquant un langage articulé (probablement déjà esquissé chez Homo habilis).
Les Homo ergaster installés en Asie sont probablement à l'origine des Homo erectus, au squelette
très robuste, mais à la capacité crânienne élevée et à la face réduite. Ces derniers produisent une industrie
lithique souvent proche de l'Oldowayen mais parfois clairement acheuléenne.
 De nos jours, les nombreux fossiles découverts sur tous les continents ont complètement transformé
notre arbre évolutif en un « buisson » très fourni. Plusieurs modèles de filiation peuvent être proposés à
partir des différentes espèces d'hominidés qui se sont succédé :
Succession des différentes espèces d'hominines dans le temps:

Échelle en milliers d'années

Les âges des espèces marquées * sont estimées. - Les traits verticaux symbolisent les possibles lignées (hypothèse
"splitter", dite de l'origine unique)
(1) ou Homo sapiens archaïque ancien
Sources : [1] - [2] - [3] - [4]
v·d·m

Ce tableau ne propose que des filiations pour les représentants du genre Homo (traits verticaux). Il
prend en compte les hypothèses suivantes :
• Homo rudolfensis serait une espèce à part entière et non une sous-espèce de Homo habilis.
• Homo antecessor serait l'ancêtre commun de Homo heidelbergensis et de Homo
rhodesiensis. Ses dates d'apparition et d'extinction sont pour le moment indicatives car cette espèce
n'a été définie qu'à partir d'un seul gisement et n'est pas reconnue par l'ensemble des
paléoanthropologues.
 • Homo floresiensis descendrait directement de Homo erectus, et ses ancêtres seraient
arrivés sur l'île de Flores il y a environ 800 000 ans, mais n'auraient constitué une nouvelle espèce à
part entière que bien plus tard.
• Les hommes de Néandertal appartiendraient à l'espèce Homo neanderthalensis, distincte de
Homo sapiens.
De plus, il existe deux tendances chez les paléoanthropologues. Certains sont partisans de
regrouper les spécimens fossiles au sein du plus petit nombre d'espèces et d'autres préfèrent classer ces
individus parmi le plus grand nombre d'espèces fossiles.

Les origines de l'homme moderne [modifier]

Article détaillé : Origine africaine de l'homme moderne.
Les Homo ergaster d'Afrique s'installent en Europe. Descendant des premiers hommes qui ont
conquis l'Europe (Homo heidelbergensis) et le Moyen-Orient, et qui se sont probablement trouvé isolés lors
de certaines périodes de glaciation, Homo neanderthalensis (Homme de Néandertal) a vécu de 200 000 à 30
000 ans. Son corps présente des adaptations au froid. C'est probablement lui qui adopte les premiers rites
funéraires.
On trouve des représentants d'une lignée prénéandertalienne entre 1 Ma et 120 000 ans, mais les
origines véritables sont loin d'être éclaircies (il y aurait peut-être eu continuité en Asie, mais remplacement
en Europe…).
D'autres populations d’Homo ergaster venant d'Afrique et du Proche-Orient sont à l'origine des
"Proto-Cro-Magnon", puis des Cro-Magnon (Homo sapiens). [réf. nécessaire] Vers 40 000 ans, Homo
neanderthalensis et Homo sapiens (Homme de Cro-Magnon) se côtoient en Europe. Ce dernier constitue le
premier représentant de notre espèce. Sa morphologie longiligne traduit des origines très probablement
africaines[4].
Les plus vieux ossements attribués à Homo sapiens ont en effet été découverts en Afrique.
Aujourd'hui, les paléontologues donnent à Homo sapiens un âge d'environ 200 000 ans puisque les plus
vieux ossements retrouvés sont deux crânes datés de -195 000 ans, et appelés Omo 1 et Omo 2; viennent
ensuite ceux de l'Homme d'Herto encore appelé Homo sapiens idaltu, datés d'environ -154 000 ans.

Fossiles [modifier]
Datation Date de l
Illustration Nom Espèce
en Ma découverte

DPC 2803 plus de Aegyptopithecus

31 Ma zeuxis
Aegyptopithecus zeuxis
image sur
Skulls Unlimited

Proconsul
KNM RU 7290 18 Ma 1948
africanus
Proconsul africanus
image sur
Skulls Unlimited

16-18 Afropithecus
KNM WK 16999 1986
Ma turkanensis
16-18 Turkanapithecus
KNM WK 16950 1986
Ma kalakolensis
 Oreopithecus
IGF 11778 8 Ma 1872
bambolii
Lufengpithecus
IVPP PA644 8 Ma 1978
lufengensis
Sivapithecus indicus
Sivapithecus
image sur GSP 15000 8 Ma 1979
indicus
Skulls Unlimited

Laetoli footprint
image sur empreintes de
3.7 Ma 1976
Modern Human Laetoli
Origins

LH 4
image sur 3.6 - 3.8 Australopithecus
LH 4 1974
Modern Human Ma afarensis
Origins

KNM-WT 40000
image sur Kenyanthropus
KNM WT 40000 3.5 Ma 1999
Modern Human platyops
Origins
 « Little Foot » Australopithecus
3.3 Ma 1994
STW 573 ?
Australopithecus
DIK-1 3.3 Ma 2000
afarensis
AL 200-1
image sur 3 - 3.2 Australopithecus
AL 200-1 1975
Modern Human Ma afarensis
Origins

AL 200-1
image sur 3 - 3.2 Australopithecus novembre
AL 129-1
Modern Human ma afarensis 1973
Origins

Australopithecus
K 12 1995
bahrelghazali
 « Lucy » Australopithecus 30
3.2 Ma
AL 288-1 afarensis novembre 1974

AL444-2
image sur Australopithecus
AL 444-2 3 Ma 1991
Modern Human afarensis
Origins
 « Mrs Ples » 2.6 - 2.8 Australopithecus
1947
STS 5 Ma africanus

STS 5

STS 14
2.6-2.8 Australopithecus
image sur Modern STS 14 1947
Ma africanus
Human Origins

STS 52
Australopithecus
image sur Modern STS 52
africanus
Human Origins

STS 71
Australopithecus
image sur Modern STS 71 2.5 Ma 1947
africanus
Human Origins
 Australopithecus
Taung 1 2.5 Ma 1924
africanus

KNM WT 17000 Paranthropus

2.5 Ma 1985
("Black Skull") aethiopicus

KNM WT 17000
image sur Modern
Human Origins

TM 1517
Paranthropus
image sur Modern TM 1517 2 Ma 1938
robustus
Human Origins
 KNM ER 1813
image sur Modern KNM ER 1813 1.9 Ma Homo habilis 1973
Human Origins

KNM ER 1470 1.9 Ma Homo rudolfensis 1972

KNM ER 1470

Paranthropus
SK 48 1.8 Ma 1948
robustus
SK 48
image sur Modern
Human Origins
 OH 24
image sur Modern OH 24 1.8 Ma Homo habilis 1968
Human Origins

OH 8
image sur Modern OH 8 1.8 Ma Homo habilis 1960
Human Origins

Paranthropus
OH 5 1.8 Ma 1959
boisei

OH 5
image sur Modern
Human Origins

D 2700 1.8 Ma Homo georgicus 2001

OH 7 OH 7 1.75 Ma Homo habilis 1960

image sur Modern
Human Origins

KNM ER 3733
image sur Modern KNM ER 3733 1.75 Ma Homo ergaster 1975
Human Origins

100px KNM ER 1805 1.74 ma Homo habilis

Homo habilis
STW 53
sensu lato
image sur 1.5 - 2
StW 53 ou 1976
Modern Human Ma
non classé
Origins
voir Grine et al. 1996

Homo habilis
SK 847
sensu lato
image sur 1.5 - 2
SK 847 ou 1949
Modern Human Ma
non classé
Origins
voir Grine et al. 1996

DNH 7
image sur 1.5 - 2 Paranthropus
DNH 7 1994
Modern Human Ma robustus
Origins
 Paranthropus
100px SK 46 1.5-2 Ma 1949
robustus
OH 13 1.7 Ma Homo habilis 1963

KNM ER 406
image sur Paranthropus
KNM ER 406 1.7 Ma 1969
Modern Human boisei
Origins

KNM ER 732
image sur Paranthropus
KNM ER 732 1.7 Ma 1970
Modern Human boisei
Origins

KNM ER 23000

Paranthropus
100px KNM WT 17400 1.7 Ma
boisei
KNM WT 15000
Homo erectus
image sur
KNM WT 15000 1.6 Ma ou 1984
Modern Human
Homo ergaster
Origins
 1.4 - 1.6
100px KNM ER 3883 Homo erectus
Ma

Paranthropus
100px Peninj Mandible 1.5 Ma 1964
boisei
Chellean OH 9
Image sur
OH 9 1.5 Ma Homo erectus 1960
Modern Human
Origins

KNM ER 992
Image sur
KNM ER 992 1.5 ma Homo ergaster 1971
Modern Human
Origins

Paranthropus
KGA 10-525 1.4 Ma 1993
boisei
moins
Sangiran 4 Homo erectus 1939
de 1 Ma

Sangiran 2 Sangiran 2 0.7-1.6 Homo erectus 1937

image sur Ma
Modern Human
Origins

Homo erectus ou
Trinil 2 .7 - 1 ma 1891
Pithecanthropus-1

700 000
Ternifine 2-3 Homo erectus 1954
a

Sangiran 17
image sur 700 000
Sangiran 17 Homo erectus 1969
Modern Human a
Origins

Hexian PA830
image sur 400-500
Hexian Homo erectus 1980
Modern Human 000
Origins

230-460 années
Homme de Pékin Homo erectus
000 1930

Zhoukoudian XII
 Bodo Homo
image sur heidelbergensis
600 000 1976
Modern Human ou
Origins Homo erectus
500 Homo
Mauer 1 1907
0000 heidelbergensis
Ndutu

Arago 21 Homo heidelbergensis

400k 1971
homme de Tautavel ou
Homo erectus

Arago XXI

Steinheim
image sur Homo
Steinheim Skull 350 000 1933
Modern Human heidelbergensis ?
Origins
 Petralona 1
image sur 250 -
Petralona 1 Homo erectus ? 1960
Modern Human 500 000 a
Origins

250 000 Homo sapiens ? 1935

Ngandong 7 250 000 Homo sapiens ? 1931

Broken Hill 1 Broken Hill 1 125k - Homo

image sur (crâne de Kabwe 300 000 rhodesiensis
1921
Modern Human ou or
Origins « homme de Rhodésie ») Homo heidelbergensis
Dali Homo
image sur heidelbergensis
Dali 209 000 1978
Modern Human ou
Origins Homo sapiens ?
LH 18 Homo sapiens 1976

Omo 1 Omo 1 190 000 Homo sapiens 1967-197

Image sur a
Modern Human
Origins

Omo II
Image sur
Omo 2 ? Homo sapiens ?
Modern Human
Origins

Herto skull
Homo sapiens 1997 -
images sur Homme de Herto 160 000
idaltu 2003
ABC Science

Jebel Irhoud 3
Machoire Jebel Irhoud 1, 2, 3 160 000
Homo sapiens vers 1991
Image sur et 4 a
PhysOrg.com

Tabun 1
Image at Homo
Tabun C1 120k 1967-197
Modern Human neanderthalensis
Origins

Klasies
Images sur 125 -75 années
Homo sapiens
Modern Human 000 a 1960
Origins
 Krapina C
Images sur 100 000 Homo
1899
Modern Human a neanderthalensis
Origins

Skhul V
images sur 90-100
Skhul V Homo sapiens 1933
Modern Human 000 a
Origins

Qafzeh IX/
Image at 90-100 Homo sapiens ou
Qafzeh IX 1933
Modern Human 000 a neanderthalensis
Origins

Skhul IX Homo sapiens ?

Qafzeh VI
Image sur 90-100
Qafzeh VI Homo sapiens 1933
Modern Human 000 a
Origins
 Homo 17
La Ferrassie 1 70 000 a
neanderthalensis septembre 1909

La Ferrassie 1

40-55 Homo
La Quina 5
000 a neanderthalensis
40-55 Homo
La Quina 18
000 neanderthalensis
Mount Circeo 1
Image sur 40-60 Homo
Monte Circeo 1 1939
Modern Human 000 a neanderthalensis
Origins

Saccopastore 1
Image at Homo
Saccopastore 1 60 000 a
Modern Human neanderthalensis
Origins
 Gibraltar 1
Image sur Homo
Gibraltar 1
Modern Human neanderthalensis
Origins

Shanidar 1
Image sur 60 - 80 Homo
Shanidar 1 1961
Modern Human 000 neanderthalensis
Origins

Amud 1
Image sur Homo
Amud 1
Modern Human neanderthalensis
Origins

Amud 7
Image sur Homo
Amud 7
Modern Human neanderthalensis
Origins

La Chapelle-aux- 60 000 a Homo 3 août 190

Saints neanderthalensis
La Chapelle-aux-Saints

Le Moustier
Image sur Homo
Le Moustier 45 000 a 1909
Modern Human neanderthalensis
Origins

Homo
Neanderthal 1 40 000 a 1856
neanderthalensis

Neanderthal 1

Qafzeh VI
92 - 115
images at années
Jebel Qafzeh 6 000 a ou 30 - 55 Homo sapiens
Modern Human 1930
000 a
Origins
 27 - 53
100px NG 6 Homo erectus 1931
000 a

crâne de Hofmeyr 36 000 a Homo sapiens 1952

Cro-Magnon 1 30 000 a Homo sapiens 1868

Cro-Magnon 1

30 - 35
Homo sapiens 1909
000

Predmosti 3 Predmost 3 26 000 a Homo sapiens 1894

images sur
Modern Human
Origins

Homo
floresiensis
ou
18 000 a Homo sapiens 2003
ou
Homo erectus
ou ?

Minatogawa 1
images sur 16 - 18
Minatogawa 1 Homo sapiens 1970
Modern Human 000 a
Origins

Tandou 15 000 a Homo sapiens 1967

8 - 20
Wadi Kubbaniya Homo sapiens 1982
000 a

Kow Swamp 1
images sur 9 - 13
Kow Swamp 1 Homo sapiens 1968
Modern Human 000
Origins
 8 - 12 années
Afalou 13 Homo sapiens
000 a 1920

6 - 12
Wadi Halfa 25 Homo sapiens 1963
000 a

Wadjak 1
images sur 10 - 12
Wadjak 1 Homo sapiens 1888
Modern Human 000 a
Origins

Tepexpan 11k Homo sapiens 1947

Cerro Sota 2 11 000 a Homo sapiens 1936

4.9 -
SDM 16704 Homo sapiens 1929
11.8 000 a

6 - 9 000
Lo 4b Homo sapiens 1965-197
a

3.5 000
Kerma 27 Homo sapiens 1913-191
a

« Ötzi » 3 300 a Homo sapiens 1991

 1 500 - 3
Five Knolls 18 Homo sapiens 1925-192
500

moins
Humboldt Sink Homo sapiens
de 1500 ans

300 -
SCDG K102 Homo sapiens 1982
400 ans

Caractéristiques de la lignée humaine [modifier]

Ces caractères peuvent être classés en trois catégories :

Bipédie [modifier]
• Quatre courbures au niveau de la colonne vertébrale ;
• Un trou occipital avancé à la base du crâne, ce qui est en relation avec une locomotion
bipède ;
• Bassin large et évasé qui permet une meilleure stabilité durant la marche ;
• Fémurs obliques par rapport à la verticale, ce qui permet aux pieds d'être à l'aplomb du
centre de gravité ;
• Un pouce parallèle aux autres orteils et l'existence d'une voûte plantaire ;
• Des membres antérieurs raccourcis.
L'évolution crânienne [modifier]

Les caractères dérivés propres au crâne humain

• Grande capacité crânienne (entre 500 et 1600 cm³) ;
• Arrondissement du crâne ;
• Face aplatie (pas de prognathisme) ;
• Absence de bourrelet sus-orbitaire.

Activités culturelles [modifier]

• Production d'outils et maîtrise de la taille de la pierre, réalisation d'un outillage diversifié
(bifaces, grattoirs...) ;
• Outils complexes, réutilisables, transmissibles avec industrialisation donc apprentissage et
spécialisation, répartition des tâches ; [réf. nécessaire]
• Maîtrise du feu (Homo heidelbergensis ou Homo erectus) ;
 1850
• Rites funéraires (à partir de
Homo neanderthalensis) ;
• Art (peintures, sculptures...) (à
partir d'Homo sapiens) ;
• Langage articulé à base de
phonèmes avec transferts de
conceptualisation. [réf. nécessaire]

Évolution technique et 1900

sociale [modifier]
Articles détaillés : Préhistoire,
Paléolithique, Mésolithique, Néolithique, art
préhistorique et industrie lithique.

Histoire de la théorie de
l'évolution humaine [modifier] 1950
L'origine de l'homme est un concept
fondamental de la métaphysique ; elle a
longtemps été expliquée par des mythes du récit
originel plus ou moins liés à la religion. La théorie
admise en Occident et au Moyen-Orient pendant
plusieurs siècles est issue de la mythologie

2002
juive : l'homme et la femme descendraient du couple formé par Adam et Ève. Le préadamisme n'est devenu
majoritaire qu'après le début du XVIIIe siècle et l'idée de Giordano Bruno qui voit en l'homme le cousin d'un
singe ne devient majoritaire parmi les scientifiques qu'avec l'affirmation des théories évolutives de Charles
Darwin. Ces idées sont toutes violemment condamnées par les églises chrétiennes occidentales. Le fossile
de l'homme de Néandertal découvert en 1856 appuie l'hypothèse évolutive, même si l'idée même qu'une
espèce d'homme distincte de la nôtre ait existé par le passé (et ait disparu) fut particulièrement difficile à
admettre. Elle ne fut proposée par Charles Darwin qu'en 1871. L'homme de Java est découvert en 1890. La
théorie de l'évolution a été popularisée par Darwin dans L'Origine des espèces ; le principal argument des
contradicteurs de cette théorie est le manque de fossiles intermédiaires. Ceci a permis à des faussaires de
berner le monde scientifique, comme avec l'affaire de l'Homme de Piltdown, composé de morceaux de
squelettes d'homme et d'orang-outan.
La définition du genre Homo reste floue et varie selon les critères retenus ; le principal critère reste le
volume de la boîte crânienne.

Schéma caricatural et scientifiquement inexact de la « marche au progrès » et de la « transition

homme-singe »
 La découverte de Lucy en 1974 a permis d'appuyer la théorie d'une origine est-africaine de la lignée
humaine.
En 2004, la découverte du Piérolapithèque en Catalogne a fourni de nouveaux éléments sur la
question du dernier ancêtre commun. Cette espèce date d'environ 13 millions d'années avant le présent.

Notes et références [modifier]

1. ↑ Les chiens ou les ruminants ont une perception des couleurs bien moindre que celle des
primates.
2. ↑ Le groupement de ces deux familles, qui possèdent cinq chromosomes {6, 19, 21, 22, X}
pratiquement identiques, étant parfois appelé Hominoïdés
3. ↑ publiés dans la revue Nature N° 441, p. 1103 s. ;(en)Nature,17 mai 2006 [archive]
4. ↑ Source : "The 10,000 Year Explosion" - Gregory Cochran. [archive]

Voir aussi [modifier]

Articles connexes [modifier]
• Bipédie
• A.A.T. (Aquatic Ape Theory) ou « théorie du primate aquatique »
• Théorie de l'évolution
 Bibliographie [modifier]
• Les Origines de l'homme, Pascal Picq (2005, dernière édition)
• L'Aventure humaine, des molécules à la culture, Boyd & silk, 2004, éditions de boeck .

[Dérouler]
v·d·m
Liste d'espèces d'hominines
Premiers Sahelanthropus tchadensis (Toumaï) • Orrorin tugenensis (Millenium
hominidés Ancestor)

Ardipithèques A. kadabba • A. ramidus

A. afarensis (Lucy, Selam, Kadanuumuu) • A. africanus • A. anamensis •
Australopithèques
A. bahrelghazali (Abel) • A. garhi
et Paranthropes
P. aethiopicus • P. boisei • P. robustus
Kenyanthrope Kenyanthropus platyops

H. antecessor • H. habilis • H. rudolfensis • H. rhodesiensis •

H. georgicus • H. cepranensis • H. erectus • H. ergaster •
Homo
H. heidelbergensis (Homme de Tautavel) • H. neanderthalensis •
H. floresiensis • H. sapiens (Homme de Cro-Magnon)
 évolution des hominines

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives
 Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de la Préhistoire
 • Portail de l’origine et de l’évolution du vivant
• Portail de l’histoire de la zoologie et de la botanique

• Portail de l’anthropologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_hominin
%C3%A9s ».

Catégories : Paléoanthropologie | Anthropologie physique et biologique | Néogène | Histoire

évolutive | [+]
 W000

Evolution of Influenza
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Influenza (Flu)
 Types
Avian (A/H5N1 subtype) · Canine
Equine · Swine (A/H1N1 subtype)
Vaccines
2009 pandemic (Pandemrix)
ACAM-FLU-A · Fluzone · Influvac
Live attenuated (FluMist) · Optaflu
Treatment
Amantadine · Arbidol · Laninamivir
Oseltamivir · Peramivir · Rimantadine
Vitamin D · Zanamivir
Pandemics
 2009 · 1968–1969 Hong Kong · 1918
Outbreaks
2008 West Bengal
2007 Bernard Matthews H5N1
2007 Australian equine
2006 H5N1 India · 1976 swine flu
See also
Flu season · Influenza evolution
Influenza research
Influenza-like illness
v•d•e
To comply with Wikipedia's guidelines, the introduction of this article may need to be
rewritten. Please discuss this issue on the talk page and read the layout guide to make sure
the section will be inclusive of all essential details. (April 2010)
The virus causing influenza is one of the best known pathogens found in various species. In
particular, the virus is found in birds as well as mammals including horses, pigs, and humans.[1] The
phylogeny, or the evolutionary history of a particular species, is an important component when analyzing the
evolution of influenza. Phylogenetic trees are graphical models of the relationships between various species.
They can be used to trace the virus back to particular species and show how organisms that look so different
may be so closely related.[1]
Contents
[hide]
• 1 Mechanisms of evolution
• 1.1 Reassortment
• 1.2 Drift
• 2 Transmission
• 2.1 Species and barriers
• 2.2 Geographic differences
• 3 Phylogenetic analysis
• 4 Future impact
• 5 References

[edit] Mechanisms of evolution

Two common mechanisms by which viruses evolve are reassortment and genetic drift.[2]

[edit] Reassortment
Reassortment allows new viruses to evolve under both natural conditions and in artificial cultures.[2]
In fact, the 1957 evolution of the H2N2 virus is thought to be a result of reassortment.[2] In this case, human
H1N1 strains and avian influenza A genes were mixed.[2] Infecting tissue cultures can demonstrate how
pathogenic qualities can evolve for a particular species even though the reassorted virus may be
nonpathogenic for another species.[2] A prime example of evolution under natural conditions is the
reassortment of two avian influenza strains that were discovered in dead seals back in 1979.[2]

[edit] Drift
New viruses can also emerge by drift. Drift can refer to genetic drift or antigenic drift.[2] Mutation and
selection for the most advantageous variation of the virus takes place during this form of evolution.[2]
Antigenic mutants can evolve quickly due to the high mutation rate in viruses.[2] This evolution occurs under
the pressure of antibodies or immune system responses.[2]

[edit] Transmission
[edit] Species and barriers
The transmission, or how the influenza virus is passed from one species to another, varies. There are
barriers that prevent the flow of the virus between some species ranging from high to low transmission. For
example, there is no direct pathway between humans and birds.[2] Pigs however, serve as an open pathway.
There is a limited barrier for them to spread the virus.[2] Therefore, pigs act as a donator of the virus
relatively easily.

[edit] Geographic differences

Phylogenetic maps are a graphical representation of the geographic relationships among species.
They indicate that the human influenza virus is minimally impacted by geographic differences.[1] However,
both swine and avian influenza does appear to be geographically dependent.[1] All three groups (avian,
swine, and human) show chronological differences. The human influenza virus is retained in humans only,
meaning it does not spread to other species.[1] Some lineages and sublineages of the virus emerge and may
be more prevalent in certain locations. For instance, many human influenza outbreaks begin in Southeast
Asia.[2]

[edit] Phylogenetic analysis

Phylogenetic analysis can help determine past viruses and their patterns as well as determining a
common ancestor of the virus. Past studies reveal that an avian virus spread to pigs and then to humans
approximately 100 years ago.[2] This resulted in human lineages further evolving and becoming more
prominent and stable.[2]
Analysis can also feature relationships between species. The 1918 Spanish influenza virus
demonstrates this. Even though the hemagglutinin (HA) gene was closer in relation to avian strains than
mammalian ones, it was, in fact, mammalian.[3] The gene may have been adapting in humans even prior to
1918.[3] Breaking down the phylogenetic history of the influenza virus shows that there is a common
ancestor that reaches back before the 1918 outbreak that links the current human virus to the swine virus.[4]
The ancestor was derived from an avian host.[2]

[edit] Future impact

Looking at the past phylogenetic relationships of the influenza virus can help lead to information
regarding treatment resistance, selecting vaccine strains, and future influenza strains.
 In current years, there has been a huge increase in the amount of resistance to certain drugs,
including the antiviral compound adamantine.[5] In fact, its resistance has recently climbed from 2 percent to
nearly 90 percent.[5] These records of built up resistance infer that drugs, such as adamantine, will not be
useful against the influenza virus in the future.
Various lineages may continue their presence and reassort indicating the importance of a complete-
genome approach to determine new influenza strains and future epidemics.[6][7] In terms of vaccine strain
selection, antigenic clades evolve by reassortment, not by antigenic drift.[6] This was shown in the 2003-
2004 influenza outbreak.[6]
Phylogenetic trees can help determine what codons in the HA gene of the influenza A virus have
changed in past outbreaks.[8] The more mutations there are in a virus strain, the more likely that strain is to
be a generator of a new lineage in future influenza seasons.[8]

[edit] References
1. ^ a b c d e Liu, S; Kang, J; Chen, J; Tai, D; Jiang, W; Hou, G; Chen; Li, J et al. (2009).
"Panorama phylogenetic diversity and distribution of type A influenza virus". Plos One 4: 1–20.
http://www.plosone.org/article/info:doi%2F10.1371%2Fjournal.pone.0005022.
2. ^ a b c d e f g h i j k l m n o p Scholtissek, C (1995). "Molecular evolution of influenza viruses".
Virus Genes 11: 209–215. http://springerlink.metapress.com/content/v5747t6n11622627/fulltext.pdf.
3. ^ a b Reid, A; Fanning, T; Hultin, J; Taubenberger, J (1999). "Origin and evolution of the 1918
‘‘Spanish’' influenza virus hemagglutinin gene". Proceedings of the National Academy of Sciences
USA 96: 1651–1656. http://www.pnas.org/content/96/4/1651.full.pdf.
 4. ^ Gorman, O; Donis, R; Kawaoka, Y; Webster, R (1990). "Evolution of influenza A virus PB2
genes: implications for evolution of the ribonucleoprotein complex and origin of human influenza A
virus". Journal of Virology 64: 4893–4902. http://jvi.asm.org/cgi/content/abstract/64/10/4893.
5. ^ a b Simonsen, L; Viboud, C; Grenfell, B; Dushoff, J; Jennings, L; Smit, M; Macken; Hata, M
et al. (2007). "The genesis and spread of reassortment human influenza A/H3N2 viruses conferring
adamantane resistance". Molecular Biology and Evolution 24: 24.
http://mbe.oxfordjournals.org/cgi/content/full/24/8/1811#SEC4.
6. ^ a b c Holmes, E; Ghedin, E; Miller, N; Taylor, J; Bao, Y; St George, K; Grenfell; Salzberg, S
et al. (2005). "Whole-genome analysis of human influenza A virus reveals multiple persistent
lineages and reassortment among recent H3N2 viruses". Plos Biology 3: 1579–1589.
http://www.plosbiology.org/article/info:doi/10.1371/journal.pbio.0030300.
7. ^ Vana, G; Westover, K (2008). "Origin of the 1918 Spanish influenza virus: A comparative
genomic analysis". Molecular Phylogenetics and Evolution 3: 1100–1110.
8. ^ a b Fitch, W; Bush, R; Bender, C; Subbarao, K; Cox, N year=2000. "Predicting the evolution
of human influenza A". Journal of Heredity 91: 183–185.
http://jhered.oxfordjournals.org/cgi/reprint/91/3/183.
 W000

Evolution of insects
From Wikipedia, the free encyclopedia

Jump to: navigation, search

 Evolution has produced astonishing variety in insects. Pictured are some of the possible shapes of
antennae.
The relationships of insects to other animal groups remain unclear. Although more traditionally
grouped with millipedes and centipedes, evidence has emerged favouring closer evolutionary ties with the
crustaceans. In the Pancrustacea theory, insects, together with among others Malacostraca, make up a
monophyletic group (sharing a common ancestor).

Contents
[hide]
• 1 Early evidence
• 1.1 Odonata
• 2 Origin of insect flight
• 2.1 Theories
• 3 Life cycle
• 3.1 Mayflies
• 3.2 Distant ancestors
• 4 See also
• 5 References
• 6 External links
[edit] Early evidence
The oldest definitive insect fossil is the Devonian Rhyniognatha hirsti, estimated at 396-407 million
years old.[1] This species already possessed dicondylic mandibles, a feature associated with winged insects,
suggesting that wings may already have evolved at this time. Thus, the first insects probably appeared
earlier, in the Silurian period.[1]
The subclass Apterygota (wingless insects) is now considered artificial as the silverfish (order
Thysanura) are more closely related to Pterygota (winged insects) than to bristletails (order Archaeognatha).
For instance, just like flying insects, Thysanura have so-called dicondylic mandibles, while Archaeognatha
have monocondylic mandibles. The reason for their resemblance is not due to a particularly close
relationship, but rather because they both have kept a primitive and original anatomy in a much higher
degree than the winged insects. The most primitive order of flying insects, the mayflies (Ephemeroptera), are
also those who are most morphologically and physiologically similar to these wingless insects. Some mayfly
nymphs resemble aquatic thysanurans.
Modern Archaeognatha and Thysanura still have rudimentary appendages on their abdomen called
styli, while more primitive and extinct insects known as Monura had much more developed abdominal
appendages, as seen here. The abdominal and thoracic segments in the earliest terrestrial ancestor of the
insects would have been more similar to each other than they are today, and the head had well developed
compound eyes and long antennae. Their body size is not known yet. As the most primitive group today,
Archaeognatha, is most abundant near the coasts, it could mean that this was the kind of habitat where the
insect ancestors became terrestrial. But this specialization to coastal niches could also have a secondary
origin, just as could their jumping locomotion, as it is the crawling Thysanura who are considered to be most
original (plesiomorphic). By looking at how primitive cheliceratan book gills (still seen in horseshoe crabs)
evolved into book lungs in primitive spiders and finally into tracheae in more advanced spiders (most of them
still have a pair of book lungs intact as well), it is possible the trachea of insects was formed in a similar way,
modifying gills at the base of their appendages.
So far there is nothing that suggests the insects were a particularly successful group of animals
before they got their wings.

[edit] Odonata
The Odonata (dragonflies) are also a good candidate as the oldest living member of the Pterygota.
Mayflies are morphologically and physiologically more primitive, but the derived and advanced
characteristics of dragonflies could have evolved independently in their own direction for a long time. It
seems that orders with aquatic nymphs or larvae become evolutionarily conservative once they had adapted
to water. If mayflies made it to the water first, this could partly explain why they are more primitive than
dragonflies, even if dragonflies have an older origin.
Similarly, stoneflies are the most primitive of the Neoptera, but they were not necessarily the first
order to branch off. This also makes it less likely that an aquatic ancestor would have the evolutionary
potential to give rise to all the different forms and species of insects that we know today.
Dragonfly nymphs have a unique labial "mask" used for catching prey, and the imago has a unique
way of copulating, using a secondary male sex organ on the second abdominal segment. It looks like
abdominal appendages modified for sperm transfer and direct insemination have occurred at least twice in
insect evolution, once in Odonata and once in the other flying insects. If these two different methods are the
original ways of copulating for each group, it is a strong indication that it is the dragonflies who are the oldest,
not the mayflies. There is still not agreement about this. Another scenario is that abdominal appendages
adapted for direct insemination has evolved three times in insects; once Odonata, once in mayflies and once
in the Neoptera, both mayflies and Neoptera choosing the same solution. If so, it is still possible that mayflies
are the oldest order among the flying insects. The power of flight is assumed to have evolved only once,
suggesting sperm transfer in the earliest flying insects still was done indirectly.
One possible scenario on how direct insemination evolved in insects is seen in scorpions. The male
deposits a spermatophore on the ground, locks its claws with the female's claws and then guides her over his
packet of sperm, making sure it comes in contact with her genital opening.
When the early (male) insects laid their spermatophores on the ground, it seems likely that some of
them used the clasping organs at the end of their body to drag the female over the package. The ancestors of
Odonata evolved the habit of grabbing the female behind her head, as they still do today. This action, rather
than not grasping the female at all, would have increased the male's chances of spreading its genes. The
chances would be further increased if they first attached their spermatophore safely on their own abdomen
before they placed their abdominal claspers behind the female's head; the male would then not let the female
go before her abdomen had made direct contact with his sperm storage, allowing the transfer of all sperm.
This also meant increased freedom in searching for a female mate because the males could now
transport the packet of sperm elsewhere if the first female slipped away. This ability would eliminate the need
to either wait for another female at the site of the deposited sperm packet or to produce a new packet,
wasting energy. Other advantages include the possibility of mating in other, safer places than flat ground,
such as in trees or bushes.
If the ancestors of the other flying insects evolved the same habit of clasping the female and
dragging her over their spermathophore, but posterior instead of anterior like the Odonata does, their genitals
would come very close to each others. And from there on, it would be a very short step to modify the vestigial
appendages near the male genital opening to transfer the sperm directly into the female. The same
appendages the male Odonata use to transfer their sperm to their secondary sexual organs at the front of
their abdomen.
All insects with an aquatic nymphal or larval stage seem to have adapted to water secondarily from
terrestrial ancestors. Of the most primitive insects with no wings at all, Archaeognatha and Thysanura, all
members live their entire life cycle in terrestrial environments. As mentioned previously, Archaeognatha were
the first to split off from the branch that led to the winged insects (Pterygota), and then the Thysanura
branched off. This indicates that these three groups (Archaeognatha, Thysanura and Pterygota) have a
common terrestrial ancestor, which probably resembled a primitive model of Apterygota, was an
opportunistic generalist and laid spermatophores on the ground instead of copulating, like Thysanura still do
today. If it had feeding habits similar to the majority of apterygotes of today, it lived mostly as a decomposer.
One should expect that a gill breathing arthropod would modify its gills to breathe air if it were
adapting to terrestrial environments, and not evolve new respiration organs from bottom up next to the
original and still functioning ones.
Then comes the fact that insect (larva and nymph) gills are actually a part of a modified, closed
trachea system specially adapted for water, called tracheal gills. The arthropod trachea can only arise in an
atmosphere and as a consequence of the adaptations of living on land. This too indicates that insects are
descended from a terrestrial ancestor.
And finally when looking at the three most primitive insects with aquatic nymphs (called naiads:
Ephemeroptera, Odonata and Plecoptera), each order has its own kind of tracheal gills that are so different
from one another that they must have separate origins. This would be expected if they evolved from land-
dwelling species.
 This means that one of the most interesting parts of insect evolution is what happened between the
Thysanura-Pterygota split and the first flight.

[edit] Origin of insect flight

The origin of insect flight remains obscure, since the earliest winged insects currently known appear
to have been capable fliers. Some extinct insects (e.g. the Palaeodictyoptera) had an additional pair of
winglets attached to the first segment of the thorax, for a total of three pairs.
The wings themselves are thought by many[who?] to be highly modified (tracheal) gills. And there is
no doubt that the tracheal gills of the mayfly nymph in many species look like wings. By comparing a well
developed pair of gill blades in the naiads and a reduced pair of hind wings on the adults, it is not hard to
imagine that the mayfly gills (tergaliae) and insect wings have a common origin, and newer research also
supports this.[citation needed] The tergaliae are not found in any other order of insects, and they have
evolved in different directions with time. In some nymphs/naiads the most anterior pair has become
sclerotized and works as a gill cover for the rest of the gills. Others can form a large sucker, be used for
swimming or modified into other shapes. But it doesn't have to mean that these structures were originally
gills. It could also mean that the tergaliae evolved from the same structures which gave rise to the wings, and
that flying insects evolved from a wingless terrestrial species with pairs of plates on its body segments: three
on the thorax and nine on the abdomen (mayfly nymphs with nine pairs of tergaliae on the abdomen exist,
but so far no living or extinct insects with plates on the last two segments have been found). If these were
primary gills, it would be a mystery why they should have waited so long to be modified when we see the
different modifications in modern mayfly nymphs.
[edit] Theories
When the first forests arose on earth, new niches for terrestrial animals were created. Spore-feeders
and others who depended on plants and/or the animals living around them would have to adapt too to make
use of them. In a world with no flying animals, it would probably just be a matter of time before some
arthropods who were living in the trees evolved paired structures with muscle attachments from their
exoskeleton and used them for gliding, one pair on each segment. Further evolution in this direction would
give bigger gliding structures on their thorax and gradually smaller ones on their abdomen. Their bodies
would have become stiffer while thysanurans, which didn't evolve flight, kept their flexible abdomen.
Mayfly nymphs must have adapted to water while they still had the "gliders" on their abdomen intact.
So far there is no concrete evidence to support this theory either, but it is one that offers an explanation for
the problems of why presumably aquatic animals evolved in the direction they did.
Leaping and arboreal insects seems like a good explanation for this evolutionary process for several
reasons. Because early winged insects were lacking the sophisticated wing folding mechanism of neopterous
insects, they must have lived in the open and not been able to hide or search for food under leaves, in
cracks, under rocks and other such confined spaced. In these old forests there weren't many open places
where insects with huge structures on their back could have lived without experiencing huge disadvantages.
If insects got their wings on land and not in water, which clearly seems to be the case, the tree canopies
would be the most obvious place where such gliding structures could have emerged, in a time when the air
was a new territory. The question is if the plates used for gliding evolved from "scratch" or by modifying
already existing anatomical details. The thorax in Thysanura and Archaeognatha are known to have some
structures connected to their trachea which share similarities to the wings of primitive insects. This suggests
the origin of both the wings and the spiracles are related.
 Gliding requires universal body modifications, as seen in present-day vertebrates such as some
rodents and marsupials, which have grown wide, flat expansions of skin for this purpose. The flying dragons
(genus Draco) of Indonesia has modified its ribs into gliders, and even some snakes can glide through the air
by spreading their ribs. The main difference is that while vertebrates have an inner skeleton, primitive insects
had a flexible and adaptive exoskeleton.
It is clear that there would have been some animals living in the trees, as animals are always taking
advantage of all available niches, both for feeding and protection. At the time, the reproductive organs were
by far the most nutritious part of the plant, and these early plants show signs of arthropod consumption and
adaptations to protect themselves, for example by placing their reproductive organs as high up as possible.
But there will always be some species who will be able to cope with that by following the their food source up
the trees.
Knowing that insects were terrestrial at that time and that some arthropods (like primitive insects)
were living in the tree crowns, it seems less likely that they would have developed their wings down on the
ground or in the water.
In a three dimensional environment such as trees, the ability to glide would increase the insects'
chances to survive a fall, as well as saving energy. This trait has repeated itself in modern wingless species
such as the gliding ants who are living an arboreal life. When the gliding ability first had originated, gliding
and leaping behavior would be a logical next step, which would eventually be reflected in their anatomical
design.
The need to navigate through vegetation and to land safely would mean good muscle control over
the proto-wings, and further improvements would eventually lead to true (but primitive) wings.
While the thorax got the wings, a long abdomen could have served as a stabilizer in flight.
 It is also worth remembering that some of the earliest flying insects were large predators. This isn't
surprising since there weren't any yet any other predators hunting in the air: it was therefore a totally new
ecological niche. Some of the prey were without a doubt other insects, as insects with proto-wings would
have radiated into other species even before the wings were fully evolved. From this point onwards, the arms
race could continue: the same predator/prey co-evolution which has existed as long as there have been
predators and prey on earth; both the hunters and the hunted were in need of improving and extending their
flight skills even further to keep up with the other.
Insects that had evolved their proto-wings in a world without flying predators could afford to be
exposed openly without risk, but this changed when carnivorous flying insects evolved. It is unknown when
they first evolved, but once these predators had emerged they put a strong selection pressure on their victims
and themselves. Those of the prey who came up with a good solution about how to fold their wings over their
backs in a way that made it possible for them to live in narrow spaces would not only be able to hide from
flying predators (and terrestrial predators if they were on the ground) but also to exploit a wide variety of
niches that were closed to those who couldn't fold their wings in this way. And today the neopterous insects
(those that can fold their wings back over the abdomen) are by far the most dominant group of insects.
The water-skimming theory suggests that skimming on the water surface is the origin of insect flight.
[2] This theory is based on the fact that the first fossil insects, the Devonian Rhyniognatha hirsti, is thought to
have possessed wings, even though the insects closest evolutionary ties are with crustaceans, which are
aquatic.
[edit] Life cycle
[edit] Mayflies
Another primitive trait of the mayflies are the subimago; no other insects have this winged yet
sexually immature stage. A few specialized species have females with no subimago, but retain the subimago
stage for males.
The reasons the subimago still exists in this order could be that there hasn't been enough selection
pressure to get rid of it; it also seems specially adapted to do the transition from water to air.
The male geniatalia are not fully functional at this point. One reason for this could be that the
modification of the abdominal appendages into male copulation organs emerged later than the evolution of
flight. This is indicated by the fact that dragonflies have a different copulation organ than other insects.
As we know, in mayflies the nymphs and the adults are specialized for two different ways of living; in
the water and in the air. The only stage (instar) between these two is the subimago. In more primitive fossil
forms, the preadult individuals had not just one instar but numerous ones (while the modern subimago do not
eat, older and more primitive species with a subimagos were probably feeding in this phase of life too as the
lines between the instars were much more diffuse and gradual than today). Adult form was reached several
moults before maturity. They probably didn't have more instars after becoming fully mature. This way of
maturing is how Apterygota do it, which moult even when mature, but not winged insects.
Modern mayflies have eliminated all the instars between imago and nymph, except the single instar
called subimago, which is still not (at least not in the males) fully sexually mature. The other flying insects
with incomplete metamorphosis (Exopterygota) have gone a little further and completed the trend; here all
the immature structures of the animal from the last nymphal stage are completed at once in a single final
moult. The more advanced insects with larvae and complete metamorphosis (Endopterygota) have gone
even further. An interesting theory here is that the pupal stage is actually a strongly modified and extended
stage of subimago, but so far it is nothing more than a theory. Interestingly enough there are some insects
within the Exopterygota, thrips and whiteflies (Aleyrodidae), who have evolved pupae-like stages too.

[edit] Distant ancestors

The distant ancestor of flying insects, a species with primitive proto-wings, had a more or less
ametabolous life cycle and instars of basically the same type as thysanurans with no defined nymphal,
subimago or adult stages as the individual became older. Individuals developed gradually as they were
growing and moulting, but there were probably no big changes in between instars.
Modern mayfly nymphs do not acquire gills until after their first moult. Before this stage they are so
small that there is no need for gills to extract oxygen from the water. This could be a trait from the common
ancestor all flyers evolved from. An early terrestrial insect would have no need for paired outgrowths from the
body before it started to live in the trees (or in the water, for that matter), so it would not have any.
This would also affect the way their offspring looked like in the early instars, resembling earlier
ametabolous generations even after they had started to adapt to a new way of living, in a habitat where they
actually could have some good use for flaps along their body. Since they matured in the same way as
thysanurans with plenty of moultings as they were growing and very little difference between the adults and
much younger individuals (unlike modern insects, who are hemimetabolous or holometabolous), there
probably wasn't much room for adapting into different niches depending on age and stage. Also, it would
have been difficult for an animal already adapted to a niche to make a switch to a new niche later in life
based on age or size differences alone when these differences were not significant.
 So they had to specialize and focus their whole existence on improving a single lifestyle in a
particular niche. The older the species and the single individuals became, the more would they differ from
their original form as they adapted to their new lifestyle better than the generations before. The final body
design was no longer achieved while still inside the egg, but continued to develop for most of the life, causing
a bigger difference between the youngest and oldest individuals. Assuming that mature individuals most
likely mastered their new element better than did the nymphs who had the same lifestyle, it would appear to
be an advantage if the immatures reached adult shape and form as soon as possible. This may explain why
they evolved fewer but more intense instars and a stronger focus on the adult body, and the differences
between the adults and the first instars were greater, instead of just gradually growing bigger as earlier
generations had done. This evolutionary trend explains how they went from ametabolous to hemimetabolous
insects.
Reaching maturity and a fully grown body became only a part of the development process, gradually
also a new anatomy and new abilities only possible in the later stages of life, were included. The anatomy
they were born and grew up with had limitations the adults who had learned to fly didn't have. If they couldn't
live their early life the way adults did, immature individuals had to adapt to the best way of living and
surviving despite their limitations till the moment came when they could leave them behind. This would be a
starting point in the evolution where imago and nymphs started to live in different niches, some more clearly
defined than others. Also, a final anatomy, size and maturity reached at once with a single final nymphal
stage meant less waste of time and energy, and also made a more complex adult body structure. These
strategies obviously became very successful with time.
Late Carboniferous and Early Permian insect orders include both several current very long-lived
groups (mayflies, (Ephemeroptera), dragonflies (Odonata), cockroaches (Blattodea), and Orthoptera
(grasshoppers and their relatives)) and a number of Paleozoic forms. During this era, some giant dragonfly-
like forms – e.g. Meganeura and Meganeuropsis (Order Protodonata) and Mazothairos (Order
Palaeodictyoptera) – reached wingspans of 55 to 70 cm (22 to 28 in), making them far larger than any living
insect. Also their nymphs must have had a very impressive size. This gigantism may have been due to higher
atmospheric oxygen levels (up to 80% above modern levels during the Carboniferous) that allowed increased
respiratory efficiency relative to today. The lack of flying vertebrates could have been another factor.
Most extant orders of insects developed during the Permian era that began around 270 million years
ago. Many of the early groups became extinct during the Permian-Triassic extinction event, the largest mass
extinction in the history of the Earth, around 252 million years ago.
The remarkably successful Hymenopterans appeared in the Cretaceous but achieved their diversity
more recently, in the Cenozoic. A number of highly successful insect groups — especially the Hymenoptera
and Lepidoptera (butterflies), as well as many types of Diptera (flies) and Coleoptera (beetles) — evolved in
conjunction with flowering plants, a powerful illustration of co-evolution.
Many modern insect genera developed during the Cenozoic; insects from this period on are often
found preserved in amber, often in perfect condition. Such specimens are easily compared with modern
species. The study of fossilized insects is called paleoentomology.

[edit] See also

• Butterfly evolution
• Spider evolution
[edit] References
1. ^ a b [1] Michael S. Engel and David A. Grimaldi. (2004). New light shed on the oldest insect.
Nature 427: 627-630 (12 February 2004) | doi:10.1038/nature02291
2. ^ Marden, J.H. and M.G. Kramer. 1994. Surface-skimming stoneflies: a possible intermediate
stage for insect flight evolution. Science 266, 427-430.
• Grimaldi, David and Michael S. Engel (2005-05-16). Evolution of the Insects. Cambridge
University Press. ISBN 0-521-82149-5. — an up to date review of the evolutionary history of the
insects
• Rasnitsyn, A.P. and Quicke, D.L.J. (2002). History of Insects. Kluwer Academic Publishers.
ISBN 1-4020-0026-X. — detail coverage of various aspects of the evolutionary history of the insects
[2]

[edit] External links

• What arthropod brains say about arthropod phylogeny
• Ecological history of the terrestrial insects
• Geographical history of the insects
• The Primitive Characters of Extant Mayflies (Ephemeroptera)
• The insect abdomen and terminalia
• Morphology of Ephemeroptera
• International Palaeoentomological Society
 [hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_insects"

Categories: Evolution by taxon | Entomology

 W000

Lemur evolutionary history

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 Mouse lemurs, the smallest primates in the world, evolved in isolation along with other lemurs on the
island of Madagascar.
The evolutionary history of lemurs occurred in isolation from other primates on the island of
Madagascar for at least 40 million years. Lemurs are prosimian primates belonging to the suborder
Strepsirrhini, which branched off from other primates less than 63 mya (million years ago). They share some
traits with the most basal primates, and thus are often confused as being ancestral to modern monkeys,
apes, and humans. Instead, they merely resemble ancestral primates.
Lemurs are thought to have evolved during the Eocene or earlier, sharing a closest common
ancestor with lorisiforms. Fossils from Africa and tests of nuclear DNA suggest that lemurs made their way to
Madagascar between 40 and 52 mya. Molecular tests offer an alternative date range of 62 to 65 mya. An
ancestral lemur population is thought to have inadvertently rafted to the island on a floating mat of vegetation,
although hypotheses for land bridges and island hopping have also been proposed. The timing and number
of hypothesized colonizations has traditionally hinged on the phylogenetic affinities of the Aye-aye, the most
basal member of the lemur clade.
Having undergone their own independent evolution on Madagascar, lemurs have diversified to fill
many niches normally filled by other types of mammals. They include the smallest primates in the world, and
once included some of the largest. Since the arrival of humans approximately 2,000 years ago, they are now
restricted to 10% of the island, or approximately 60,000 square kilometres (23,000 sq mi), and many are
facing extinction. For this reason, researchers have been trying to identify and assess every species. Over
the last 10 to 20 years, there has been a steep increase in the number of recognized lemur species and
subspecies, both through the discovery of new species and the elevation of existing subspecies to full
species status. Currently there are approximately 100 or more recognized species or subspecies of living
lemur, which are divided into five families and 15 genera. If the extinct subfossil lemurs are included, an
additional three families, eight genera, and 17 species would be included. The recent rise in species numbers
is due to both improved genetic analysis and a push in conservation to encourage the protection of isolated
and distinct lemur populations. Not everyone in the scientific community supports these taxonomic changes,
with some preferring instead an estimate of 50 living species.
Contents
[hide]
• 1 Evolutionary history
• 1.1 Colonization of Madagascar
• 1.2 Diversification
• 2 Distribution and diversity
• 3 Taxonomic and phylogenetic classification
• 3.1 Suprageneric classification
• 3.2 Genus-level classification
• 3.3 Species-level classification
• 4 Notes
• 5 References

[edit] Evolutionary history

Lemurs are prosimian primates belonging to the suborder Strepsirrhini. Like other strepsirrhine
primates, such as lorises, pottos, and galagos, they share ancestral traits with early primates. In this regard,
lemurs are popularly confused with ancestral primates; however, lemurs did not give rise to monkeys and
apes, but evolved independently on Madagascar.[1]
 Fossil evidence for the evolution of the toothcomb, a trait shared by lemurs with their closest
relatives, the lorises, weighs heavily on the evolutionary history of strepsirrhines and the lemur colonization
of Madagascar.
Primates first evolved sometime between the Middle Cretaceous and the early Paleocene periods on
either the supercontinent of Laurasia or in Africa.[2] According to molecular clock studies, the last common
ancestor of all primates dates to around 79.6 mya,[3] although the earliest known fossil primates are only 54–
55 mya old.[4] The closest relatives of primates are the extinct plesiadapiforms, the modern colugos
(commonly and inaccurately named "flying lemurs"), and treeshrews.[3] Some of the earliest known true
primates are represented by the fossil groups Omomyidae, Eosimiidae, and Adapiformes.
 The relationship between known fossil primate families remains unclear. A conservative estimate for
the divergence of haplorrhines (tarsiers, monkeys, apes, and humans) and strepsirrhines is 58 to 63 mya,[5]
and a consensus is emerging that places tarsiers close to omomyids, while eosimids gave rise to the simians
(non-tarsier haplorrhines) and the adapiforms gave rise to modern strepsirrhines, including lemurs.[6] In
2009, a highly publicized and scientifically criticized publication proclaimed that a 47 million-year-old
adapiform fossil, Darwinius masillae, demonstrated both adapiform and simian traits, making it a transitional
form between the prosimian and simian lineages.[7] Media sources inaccurately dubbed the fossil as a
"missing link" between lemurs and humans.[8]
Lemurs are currently thought to have evolved during the Eocene (37 to 55 mya),[5][9] although
molecular tests suggest the Paleocene (65 to 56 mya) or later.[9] Until recently, they were thought to have
descended directly from the diverse group of adapiforms due to several shared postcranial traits,[10] as well
as long snouts and small brains. Although adapiforms also had lemur-like auditory bullae, a prosimian
characteristic,[11] they had smaller brains and longer snouts than lemurs.[12] There are also several other
morphological differences. Most noticeably, adapiforms lack a key derived trait, the strepsirrhine toothcomb,
and possibly the toilet-claw, found not only in strepsirrhines but also in tarsiers. Unlike lemurs, adapiforms
exhibited a fused mandibular symphysis (a characteristic of simians) and also possessed four premolars,
instead of three or two.[13]
Comparative studies of the cytochrome b gene, which are frequently used to determine phylogenetic
relationships among mammals—particularly within families and genera[14]—have been used to show that
lemurs share common ancestry with lorisiforms.[13][15] This conclusion is also corroborated by the shared
strepsirhine toothcomb, an unusual trait that is unlikely to have evolved twice.[16] If adapiforms were the
ancestors of the living strepsirrhines, then the first strepsirrhines would have to predate the early Eocene, a
view supported by molecular phylogenetic studies,[13] which show that lemurs split from lorises
approximately 62 to 65 mya.[17] The molecular studies also show that lemuriforms diversified before the
modern lorisiforms.[13] Using only nuclear genes, another study dated the split between lemurs and lorises
at 60 mya, lemur diversification at 50 mya, and the lemur colonization of Madagascar somewhere between
these two approximate dates.[18]
The fossil record tells a different story. Although it can only show the earliest possible date for the
appearance of a taxonomic group, other concerns have arisen about these vastly earlier divergence dates
predicted independently of the fossil record. First, palaeontologists have expressed concerns that if primates
have been around for significantly more than 65 million years, then the first one-third of the primate fossil
record is missing. Another problem is that some of the these molecular dates have overestimated the
divergence of other mammalian orders, such as Rodentia, suggesting primate divergence might also be
overestimated. Currently the oldest known strepsirrhine, Djebelemur, dates from the early Eocene in northern
Africa and lacks a fully differentiated toothcomb. Based on fossils and other genetic tests, a more
conservative estimate dates the divergence between lemurs and lorises to around 50 to 55 mya.[5]
To complicate the ancestry puzzle, no terrestrial Eocene or Paleocene fossils have been found on
Madagascar,[19][20] and the fossil record from both Africa and Asia around this time is not much better.[13]
Fossil sites in Madagascar are restricted to only five windows in time, which omit most of the Cenozoic, from
65 mya to ~26,000 years ago. What little fossil-bearing rock exists from this vast span of time is dominated
by marine strata along the west coast.[21] The oldest lemur fossils on Madagascar are actually subfossils
dating to the Late Pleistocene.[10]

[edit] Colonization of Madagascar

Once part of the supercontinent Gondwana, Madagascar broke away from eastern Africa, the likely
source of the ancestral lemur population, about 160 mya and then from Antarctica between 80 and 130 mya.
Initially, the island drifted south from where it split from Africa (around modern Somalia) until it reached its
current position between 80 and 90 mya. Around that time, it split with India, leaving it isolated in the Indian
Ocean and separated from nearby Africa by the Mozambique Channel,[22][23] a deep channel with a
minimum width of approximately 560 kilometers (350 mi).[13] These separation dates and the estimated age
of the primate lineage preclude any possibility that lemurs could have been on the island before the
Madagascar pulled away from Africa.[24] In support of this, mammalian fossils on Madagascar from the
Cretaceous (see Mesozoic mammals of Madagascar) include gondwanatheres and other mammalian groups
that would not have been ancestral to lemurs or the other endemic mammals present on the island today.[13]

A reconstructed map of the Earth during the Early Paleocene, approximately 65 million years ago,
around the time that lemurs evolved and colonized Madagascar
With Madagascar already geographically isolated by the Paleocene and lemur diversification dating
to the same time, an explanation was needed for how lemurs had made it to the island. In the 19th century,
prior to the theory of continental drift, scientists including Philip Sclater, Étienne Geoffroy Saint-Hilaire, and
Ernst Haeckel suggested that Madagascar and India were once part of a southern continent—named Lemuria
by Sclater—that has since disappeared under the Indian Ocean.[25][26] By the early 20th century, oceanic
dispersal emerged as the most popular explanation for how lemurs reached the island.[21][23][27] The idea
first took shape under the anti-plate tectonics movement of the early 1900s, when renowned paleontologist
William Diller Matthew proposed the idea in his influential article "Climate and Evolution" in 1915. In the
article, Matthew could only account for the presence of lemurs in Madagascar by "rafting".[28] Although
unlikely, over long periods of time terrestrial animals can occasionally raft to remote islands on floating mats
of tangled vegetation, which get flushed out to sea from major rivers by floodwaters.[13][28][29] In the 1940s,
American paleontologist George Gaylord Simpson coined the term "sweepstakes dispersal" for such unlikely
events. Today, a rafting event remains the most accepted explanation for the lemur colonization of
Madagascar.[30]
Any extended ocean voyage without fresh water or food would prove difficult for a large, warm-
blooded (or "homeothermic") mammal, but today many small, nocturnal species of lemur exhibit
heterothermy, which allows them to lower their metabolism and become dormant while living off fat reserves.
Such a trait in a small, nocturnal lemur ancestor would have facilitated the ocean voyage and could have
been passed on to its descendants.[29] However, this trait has not been observed in the closely-related
lorisiforms studied to date, and could have evolved on Madagascar in response to the island's harsh
environmental conditions.[13]
Because only five terrestrial orders of mammals have made it to the island, each likely to have
derived from a single colonization,[24] and since these colonizations date to either the early Cenozoic or the
early Miocene, the conditions for oceanic dispersal to Madagascar seem to have been better during two
separate periods in the past.[13] A report published in January 2010 supported this assumption by
demonstrating that both Madagascar and Africa were 1,650 km (1,030 mi) south of their present-day
positions around 60 mya, placing them in a different ocean gyre and reversing the strong current that
presently flows away from Madagascar. The currents were even shown to be stronger than they are today,
shortening the rafting time to approximately 30 days or less, making the crossing much easier for a small
mammal. Over time, as the continental plates drifted northward, the currents gradually changed, and by
20 mya the window for oceanic dispersal had closed.[31]
Since the 1970s, the rafting hypothesis has been called into question by claims that lemur family
Cheirogaleidae might be more closely related to the other Afro-Asian strepsirrhines than to the rest of the
lemurs. This idea was initially based on similarities in behavior and molar morphology, although it gained
support with the 2001 discovery of 30 million-year-old Bugtilemur in Pakistan and the 2003 discovery of
40 million-year-old Karanisia in Egypt. Karanisia is the oldest fossil found that bears a toothcomb, whereas
Bugtilemur was thought to have a toothcomb, but also had even more similar molar morphology to
Cheirogaleus (dwarf lemurs). If these relationships had been correct, the dates of these fossils would have
had implications on the colonization of Madagascar, requiring two separate events. The most parsimonious
explanation, given the genetic evidence and the absence of toothcombed primates in European fossil sites,
[13] is that stem strepsirrhines evolved on the Afro-Arabian landmass, dispersing to Madagascar and more
recently from Africa to Asia.[32] More recently, the structure and general presence of the toothcomb in
Bugtilemur has been questioned, as well as many other dental features, suggesting it is most likely an
adapiform.[5]
An alternative form of oceanic dispersal that had been considered was island hopping, where the
lemur ancestors might have made it to Madagascar in small steps by colonizing exposed seamounts during
times of low sea level.[12][21] However, this is unlikely since the only seamounts found along the Davie
Ridge would have been too small in a such a wide channel. Even though the Comoros Islands between
Africa and Madagascar are significantly larger, they are too young, having been formed by volcanic activity
only around 8 mya.[21] A land bridge between Madagascar and Africa has also been proposed, but a land
bridge would have facilitated the migration of a much greater sampling of Africa's mammalian fauna than is
endemic to the island. Furthermore, deep trenches separate Madagascar from the mainland, and prior to the
Oligocene, sea level was significantly higher than today.[33]
Despite these issues, one variant of the land bridge hypothesis may best explain both how a land
bridge could have formed, and how it other mammalian orders failed to cross it.[5] Geological studies have
shown that following the collision of India and Asia, the Davie Fracture Zone had been pushed up by tectonic
forces, possibly high enough to create a land bridge. Indeed, core samples along the Davie Fracture Zone
suggest that at least parts of the Mozambique Channel were above sea level between 45 and 26 mya,[34] or
possibly as early as 55 mya.[5] Following the Indian-Asian collision, the fault type changed from a strike-slip
fault to a normal fault, and seafloor spreading created compression along the Davie Fracture Zone, causing it
to rise. By the early Miocene, the East African Rift created tension along the fault, causing it to subside
beneath the ocean. Coinciding with this, the divergence dates of nearly all the Malagasy mammalian orders
fall within this window. Old World monkeys, dogs, and cats did not diverge or arrive in Africa until later in the
Miocene.[34] Still, more recent dating of divergence of the Malagasy mammalian clades falls outside of this
land bridge window, and a much greater diversity of mammal groups would be expected on Madagascar had
the land bridge been present during that stretch of time.[18]
The dating of the lemur colonization is controversial for the same reasons as that of strepsirrhine
evolution. Using molecular testing, the colonization has been estimated at 62 to 65 mya based on the split
between the Aye-aye and the rest of the lemurs.[17] On the other hand, the sparse fossil record and
estimates based on nuclear genes support a more conservative estimate of 40 to 52 mya.[5] Once safely
established on Madagascar, with its limited mammalian population, the lemurs were protected from the
increasing competition from evolving arboreal mammalian groups.[19] Monkeys had evolved by the
Oligocene, and their intelligence, aggression, and deceptiveness may have given them the advantage in
exploiting the environment over the diurnal adapiform primates in Africa and Asia, ultimately driving them to
extinction, leaving only the nocturnal lorisiforms.[12][35]

[edit] Diversification

A reconstructed map of the Earth during the Early Oligocene, approximately 35 million years ago, at
a time when lemurs were diversifying.
The ancestral lemur that colonized Madagascar is thought to have been small and nocturnal.[36]
More specifically, it is thought to have had adapiform-like cranial anatomy—particularly the cranial foramina
and the middle ear—comparable to that of lemurids, while being similar to cheirogaleids in dentition and
postcranial anatomy.[5] Lemurs have since diversified greatly, starting with the Aye-aye and its extinct
relations, whose divergence is popularly thought to have occurred shortly after the lemur colonization of
Madagascar.[17] According to molecular studies, there have since been two major episodes of
diversification, from which all other known extant and extinct family lineages emerged. The remaining
families diverged during a 10 to 12 million-year window, between the Late Eocene (42 mya) and into the
Oligocene (30 mya).[17][27] In fact, these dates for this divergence window coincided with the Eocene-
Oligocene extinction event, during which time climate cooling took place and changes in ocean currents
altered weather patterns.[17][5] Outside of Madagascar, these dates also coincide with the divergence of the
lorisiform primates and five major clades of squirrels, all occupying similar niches as lemurs.[17] The dates
do not suggest that increased predation drove family-level divergence since the first carnivores arrived on the
island between 24 and 18 mya.[36]
The second major episode of diversification occurred during the Late Miocene, approximately 8 to
12 mya, and included the true lemurs (Eulemur) and the mouse lemurs (Microcebus).[17][27] In both the true
lemurs and mouse lemurs, their populations were thought to have diverged due to habitat fragmentation
caused when humans arrived on the island roughly 2,000 years ago.[10] Only recently has molecular
research started to show a more distant split in these genera.[37] Most surprising were the mouse lemurs, a
group which is now thought to contain cryptic species, meaning they are indistinguishable from each other
based solely on appearance. In contrast, true lemurs are easier to distinguish and exhibit sexual
dichromatism.[17] Studies in karyology, molecular genetics, and biogeographic patterns have also assisted in
understanding their phylogeny and diversification.[37] Although the divergence times for these two genera
are imprecise, this estimated divergence time would overlap with a change to a wetter climate in
Madagascar, as new weather patterns generated monsoons and likely influenced the plant life.[17][27]
This difference in evolutionary divergence between the two genera may be due to differences in their
activity patterns. True lemurs are often diurnal, allowing sexual partners to distinguish each other as well as
other related species visually. Mouse lemurs, on the other hand, are nocturnal, reducing their ability to use
visual signals for mate selection. Instead, they use olfactory and auditory signaling. For these reasons, true
lemurs may have evolved sexual dichromatism while mouse lemurs evolved to be cryptic species.[17]

[edit] Distribution and diversity

See also: Subfossil lemur
Since their arrival on Madagascar, lemurs have diversified both in behavior and morphology. Their
diversity rivals that of the monkeys and apes found throughout the rest of the world, especially when the
recently extinct subfossil lemurs are considered.[35] Ranging in size from the 30 g (1.1 oz) Madame Berthe's
Mouse Lemur, the world's smallest primate,[38] to the extinct 160–200 kg (350–440 lb) Archaeoindris
fontoynonti,[39] lemurs evolved diverse forms of locomotion, varying levels of social complexity, and unique
adaptations to the local climate. Until recently, they had gone on to fill many niches normally occupied by
monkeys, squirrels, woodpeckers, and large grazing ungulates.[12][19] In addition to the incredible diversity
between lemur families, there has also been great diversification among closely-related lemurs. Yet despite
separation by geographical barriers or by niche differentiation in sympatry, occasionally hybridization can
occur.[35] Lemur diversification has also created generalist species, such as the true lemurs of northern
Madagascar, which are very adaptable, mostly nondescript, and found throughout most of the island's
forests.[10]
 The Diademed Sifaka (Propithecus diadema) is one of the largest of the living lemurs, comparable in
size to the Indri. It lives in the rainforests of Madagascar and eats a varied diet of leaves and fruit.
Most of the 99 living lemur taxa are found only on Madagascar. Two species, the Common Brown
Lemur (Eulemur fulvus) and the Mongoose Lemur (Eulemur mongoz), can also be found on the Comoro
Islands, although it is assumed that both species were introduced to the islands from northwestern
Madagascar by humans within the last few hundred years.[40][41] Molecular studies on Eulemur fulvus
fulvus (from the mainland) and E. f. mayottensis (from the Comoro Islands)[24] and on Comoro and mainland
Mongoose Lemurs have supported this assumption by showing no genetic differences between the two
populations.[41] Because all lemurs, including these two brown lemur species, are only native to the island of
Madagascar, they are considered to be endemic.
Historically, lemurs ranged across the entire island inhabiting a wide variety of habitats, including dry
deciduous forests, lowland forests, spiny thickets, subhumid forests, montane forest, and mangrove. Today,
their collective range is restricted to 10% of the island, or approximately 60,000 km2 (23,000 sq mi).[42] Most
of the remaining forests and lemurs are found along the periphery of the island. The center of the island, the
Hauts Plateaux ("High Plateaus"), was converted by early settlers to rice paddies and grassland through
slash-and-burn agriculture, known locally as tavy. As erosion depleted the soil, the cyclical forest regrowth
and burning ended as the forest gradually failed to return.[43] Today, the level of floral diversity increases
with precipitation, from the dry southern forests to the wetter norther forests to the rainforests along the east
coast. Increased foliage corresponds to increased faunal diversity, including the diversity and complexity of
lemur communities.[10]
Having evolved in Madagascar's challenging environment, replete with poor soils, extreme shifts in
poor, seasonal plant productivity, and devastating climatic events such as extended droughts and annual
cyclones,[9] lemurs have adopted unique combinations of unusual traits to survive, distinguishing them
significantly from other primates. In response to limited, seasonal resources, lemurs may exhibit seasonal fat
storage, hypometabolism (including torpor and hibernation in some cheirogaleids), small group sizes, low
encephalization (relative brain size), cathemerality (activity both day and night), and/or strict breeding
seasons.[9][44] Secondarily, extreme resource limitations and seasonal breeding are thought to have
resulted in three other relatively common lemur traits: female dominance, sexual monomorphism (lack of size
differences between the sexes), and male-male competition for mates involving low levels of agonism
(conflict), such as sperm competition.[45]
The arrival of humans on the island 1,500 to 2,000 years ago has taken a significant toll, not only on
the size of lemur populations, but also on their diversity.[19] Due to habitat destruction and hunting, at least
17 species and 8 genera have gone extinct and the populations of all species have decreased.[39][46] A
couple of species once thought to have gone extinct have since been rediscovered. The Hairy-eared Dwarf
Lemur (Allocebus trichotis) was only known from five museum specimens, most collected in the late 19th
century and one in 1965. It was rediscovered in 1989[47] and has since been identified in five national parks,
although it is very rare within its range.[38] Likewise, the Greater Bamboo Lemur (Prolemur simus) was
thought to be extinct as recently as the late 1970s, but a population was located near Ranomafana National
Park in the late 1980s.[48] Historically, it had a much wider geographic distribution, shown by subfossil
remains, but today it remains one of the world's 25 most endangered primates.[48][49][50][51] One distinctive
morph (possibly a species or subspecies) of sifaka,[N 1] has not been so fortunate, having been extirpated
from all known localities.[54] Unless trends change, extinctions are likely to continue.[55]
 A life restoration of Palaeopropithecus ingens, a giant sloth lemur that went extinct less than a
thousand years ago.
 Until recently, giant species of lemur existed on Madagascar. Now represented only by recent or
subfossil remains, they were modern forms and are counted as part of the rich lemur diversity that has
evolved in isolation. Some of their adaptations were unlike those seen in lemurs today.[19] All 17 extinct
lemurs were larger than the extant forms, some weighing as much as 200 kg (440 lb),[35] and are thought to
have been active during the day.[56] Not only were they unlike the living lemurs in both size and appearance,
they also filled ecological niches that no longer exist or are now left unoccupied.[19] Large parts of
Madagascar, which are now devoid of forests and lemurs, once hosted diverse primate communities that
included more than 20 species covering the full range of lemur sizes.[57]

[edit] Taxonomic and phylogenetic classification

For a more comprehensive list, see List of lemur species
Lemur taxonomy is controversial, and not all experts agree, particularly with the recent increase in
the number of recognized species.[58][59][60] According to Russell Mittermeier, the president of
Conservation International (CI), taxonomist Colin Groves, and others, there are currently 99 recognized
species or subspecies of extant lemur, divided into five families and 15 genera.[61] Conversely, other experts
in the field label this as a possible example of taxonomic inflation,[60] and prefer instead an estimate of at
least 50 species.[58] All sides generally agree that the recently extinct subfossil lemurs should be classified
in three families, eight genera, and 17 species.[39][46]
Closest lemur relations[3]
Euarchonta
Scandentia
 (treeshrews)

Dermoptera
(colugos)

†Plesiadapiforme
s

Primates
Haplorrhini
(tarsiers, monkeys,
apes)
Strepsirrhini
Lorises,
pottos, and
galagos
Lemurs
 Since the first taxonomic classification of lemurs in 1758 by Carl Linnaeus, many changes have been
made to lemur taxonomy. Within the order Primates, treeshrews (order Scandentia) were considered basal,
prosimian primates—close relatives of lemurs—until the 1980s.[62] Colugos, also incorrectly referred to as
"flying lemurs", were once considered lemur-like primates, but were reclassified as close relatives of bats,[63]
and more recently as close relatives of primates within their own order, Dermoptera.[3] Primates, together
with their closest relatives, the treeshrews, colugos, and long-extinct plesiadapiforms, form the taxonomically
unranked Euarchonta clade within the Euarchontoglires. Also, all lorisids originally placed in the genus
Lemur by Carl Linnaeus have since been moved into either their own infraorder (Lorisiformes) or their own
superfamily (Lorisoidea) within Lemuriformes.[64][5]
 For the Malagasy primate fauna, taxonomic nomenclature proliferated during the 1800s, with the aid
of museum systematists, such as Albert Günther and John Edward Gray, as well as naturalists and
explorers, such as Alfred Grandidier and Alphonse Milne-Edwards.[65][66] The taxonomic nomenclature of
lemurs was not sorted out until decades later, when Ernst Schwarz standardized it in 1931.[65][66][67] It was
not until the 1990s that this nomenclature started to see a new wave of taxonomic change.[60]

[edit] Suprageneric classification

Since the 19th century, the classification of lemurs above the genus level has seen many changes.
Early taxonomists proposed a variety of classifications for lemurs, but generally separated indriids from other
lemurs and placed the Aye-aye in a major group of its own; some classified the dwarf and mouse lemurs with
the galagos.[68] In 1915, William King Gregory published a classification[69] that remained generally
accepted over the next decades. He placed all the lemurs together in a "series" Lemuriformes and
recognized three families: Daubentoniidae, Indriidae, and Lemuridae (including the current Cheirogaleidae
and Lepilemuridae).[68] George Gaylord Simpson's influential 1945 classification of mammals placed the
treeshrews and the fossil Anagale (both now classified outside Primates) inside Lemuriformes and classified
the fossil families Plesiadapidae and Adapidae in a superfamily Lemuroidea with most of the lemurs.[70]
Although treeshrews, plesiadapids, and the like are now no longer considered to be closely related to
lemurs, disagreements persist over the classification of lemurs and related groups, resulting in two
competing arrangements of the infraorders and superfamilies within Strepsirrhini. Colin Groves, in the 2005
third edition of Mammal Species of the World, classifies living strepsirrhines under three infraorders and two
superfamilies. This places the Aye-aye within its own infraorder, separate from both lemurs (divided into two
superfamilies) and lorises.[64] Since the publication of Mammal Species of the World, there has been little
support in the academic literature for placing the Aye-aye in its own infraorder, and more recently
Mittermeier, Groves, and other editors have ignored this taxonomic level.[61] An alternative classification
draws the lines for infraorders and superfamilies differently, though using the same general phylogenetic
tree. It classifies all living strepsirrhines under one infraorder, with the lorises and lemurs in separate
superfamilies.[5]
Two alternative lemur classifications at the infraorder and superfamily levels
3 infraorders, 2 superfamilies[64] 1 infraorder, 2 superfamilies[5

• Order Primates • Order Primates

• Suborder Strepsirrhini: non-tarsier • Suborder Strepsirrhini: non-ta
prosimians • Infraorder Lemuriform
• Infraorder Chiromyiformes: Aye- • Superfamily L
aye • Family
• Infraorder Lemuriformes monkey or bab
• Family Cheirogaleidae: • Family
dwarf and mouse lemurs dwarf and mou
• Family • Family
†Archaeolemuridae: monkey or Aye-aye
baboon lemurs • Family
• Family Indriidae: woolly lemurs, sifaka
lemurs, sifakas, and allies • Family
• Family Lemuridae: brown lemurs and all
lemurs and allies • Family
 • Family Lepilemuridae:
sportive lemurs
sportive lemur
• Family †Megaladapidae:
• Family
koala lemurs
koala lemurs
• Family
• Family
†Palaeopropithecidae: sloth
†Palaeopropit
lemurs
• Superfamily L
• Infraorder Lorisiformes: galagos
and lorises
and lorises
• Suborder Haplorrhini: tarsiers,
• Suborder Haplorrhini: tarsiers, monkeys,
and apes

The classification of several lemur taxa has elicited particular debate. Most significantly, the
placement of the Aye-aye has been controversial since its introduction to Western science in 1782, and it has
been a topic of debate up until very recently.[35][72][68] Hinged upon morphological traits and molecular
data, it has had profound implications on scientific theories.[72] Arguing against Darwin's theory of natural
selection, Richard Owen claimed in 1863 that the Aye-aye's distinct characteristics, including its ever-
growing incisors and unique, highly flexible middle finger, are so perfectly adapted for their uses in extractive
foraging that they could not have evolved gradually through natural selection.[72] More recently, the Aye-
aye's placement within the order Primates has posed problems for the rafting hypothesis for the primate
colonization of Madagascar. If this species does not form a monophyletic group with the rest of the lemurs,
then multiple colonization events would have had to occur to explain the present-day distribution of non-
human primates on Madagascar.[35]
 The Aye-aye has traditionally been difficult to classify due to its unique physical traits.
Until Richard Owen published a definitive anatomical study in 1866, early naturalists were uncertain
whether the Aye-aye (genus Daubentonia) was a primate, rodent, or marsupial.[72][73][74] In the late
eighteenth century, for example, the Aye-aye was classified under the squirrel genus Sciurus.[75] By
emphasizing its primate features, such as its postorbital bar, stereoscopic vision, and opposable hallux, over
its rodent-like teeth, Owen demonstrated its affinity with other primates.[72][76] In 1996, Ankel-Simons
demonstrated that the shape and arrangement of the Aye-aye's diminutive deciduous incisors indicate that
this genus has a shared ancestry with the toothcombed primates.[16] However, the placement of the Aye-aye
within the order Primates remained problematic until very recently. The karyotype of the Aye-aye is
noticeably different from that of its closest relatives, the lorises and the rest of the lemurs, with a diploid
chromosome count of 2n=30.[77] Based on its anatomy, researchers have found support for classifying the
genus Daubentonia as a specialized indriid, a monotypic sister group to all strepsirrhines, and an
indeterminate taxon within the order Primates.[15] In 1931, Schwarz labeled the Aye-aye as an offshoot of
Indriidae, claiming that all lemurs were monophyletic, whereas Reginald Innes Pocock had previously placed
the Aye-aye outside of the lemurs.[67] In that same year, Anthony and Coupin classified the Aye-aye under
infraorder Chiromyiformes, a sister group to the other strepsirrhines. Colin Groves upheld this classification in
2005 because he was not entirely convinced the Aye-aye formed a clade with the rest of the Malagasy
lemurs,[78] despite molecular tests that had shown Daubentoniidae was basal to all Lemuriformes.[15][79] In
2008, Russell Mittermeier, Colin Groves, and others ignored addressing higher-level taxonomy by defining
lemurs as monophyletic and containing five living families, including Daubentoniidae.[61]
Another interpretation of the Aye-aye's origins has once again called into question the single origins
of the lemurs. Comparisons have been made between the Aye-aye and a fossil strepsirrhine primate from
Africa, Plesiopithecus. Similarities in the shape of the skull and the morphology of the lower jaw have raised
the question of whether or not this could be an Aye-aye ancestor. However, the placement of an Aye-aye
ancestor in Africa would require multiple colonizations by strepsirrhine primates. Molecular tests may offer
support since they show that the Aye-aye was the first to diverge in the lemur clade and that the other lemur
families did not diverge until much later.[5]
Often classified with the galagos by early students, the cheirogaleids were placed with the other
lemurs from Gregory's 1915 classification until the early 1970s, when several anthropologists proposed that
they are more closely related to lorisiforms, based on morphological data.[68][80] However, relevant genetic
studies nearly unanimously place cheirogaleids within the lemuriform clade and Groves, who had promoted
the cheirogaleid-lorisiform relationship in a 1974 paper, by 2001 himself regarded the idea as refuted.[15][79]
[80]
Lemur phylogeny[27][57][81]
 Strepsirrhini
Lorisiform clade

Galagidae

Lorisidae

Lemur clade
Daubentonia

Lemuridae

†Megaladapidae
 Lepilemuridae

Cheirogaleidae

†Archaeolemurida
e

†Palaeopropithecida
e

Indriidae
 Classifications in the first half of the 20th century divided lemurs into three families—Daubentoniidae,
Indriidae, and Lemuridae, with the latter including the current Cheirogaleidae and Lepilemuridae.[68]
Because of concerns that Lemuridae might not be monophyletic, the family was later split; in 1982, for
example, Tattersall separated the Cheirogaleidae for the dwarf lemurs, mouse lemurs, and relatives and the
Lepilemuridae for the sportive lemurs and bamboo lemurs (including the Greater Bamboo Lemur).[82] This
classification is still used, except that the bamboo lemurs are placed in Lemuridae.[83][61]
From the 1970s to the 1990s, there have been suggestions that the ruffed lemurs might be related to
indriids or a sister group to Lemuridae and Indriidae and that the bamboo lemurs are related to the sportive
lemurs,[84] but neither view is supported by molecular phylogeny.[27] The sportive lemurs and the extinct
koala lemurs (Megaladapidae) both lack upper incisors in the permanent dentition,[73] and in 1981, Groves
placed both together in the family Megaladapidae, which he renamed Lepilemuridae in 2005 because that
older name takes precedence.[85] Genetic research does not support a close relationship between the
sportive and koala lemurs and instead places the koala lemurs as a sister group to Lemuridae; therefore, the
two are now placed in separate families (Lepilemuridae for the sportive lemurs and Megaladapidae for the
koala lemurs).[86][87][81] The sloth lemurs (Palaeopropithecidae) and monkey lemurs (Archaeopithecidae)
were classified as subfamilies within Indriidae as late as 1982,[82] but are now recognized as separate
families.[81]
The relationships among the families of lemurs have been problematic and have yet to be definitively
resolved. Two competing phylogenies exist based on genetic and molecular data. One approach (Horvath,
et al.) looks at a larger number of genes, but among fewer species. This results in Lemuridae being a sister
group to Lepilemuridae, Cheirogaleidae, and Indriidae.[27] The other approach (Orlando, et al.) looks at
fewer genes, but more lemur species. Using this analysis, Lepilemuridae becomes the sister group to
Lemuridae, Cheirogaleidae, and Indriidae.[81] Both phylogenies agree that the Malagasy primates are
monophyletic and that Daubentoniidae (the Aye-aye) is basal to the lemuriform clade, having split off
significantly earlier than the other families.[15][27][81] However, two problems create complications for both
approaches. First, the four most closely related lemur families diverged within a narrow window of
approximately 10 million years, making it much harder to distinguish the splits with molecular evidence.
Second, the divergence occurred approximately 42 mya;[17] such distant splits create a lot of noise for
molecular techniques.

[edit] Genus-level classification

Early classifications of the genera of lemurs differed in a number of ways from current taxonomy. For
example, the fork-marked lemurs were initially placed in the genus Lemur and then in Microcebus with the
mouse lemurs before being placed in their own genus Phaner;[47][88][67][89] and Charles Immanuel Forsyth
Major split the Cheirogaleus medius species group of the dwarf lemurs into a separate genus Opolemur, but
this was not accepted.[89][67] Genus-level taxonomy was largely stabilized by Schwarz in 1931,[67] but a
number of changes have become accepted:
• The Ring-tailed Lemur, ruffed lemurs, and true lemurs were once grouped together in the
genus Lemur due to a host of morphological similarities. For instance, the skeletons of the Ring-
tailed Lemur and the true lemurs are nearly indistinguishable.[50] However, ruffed lemurs were
reassigned to the genus Varecia in 1962,[90] and due to similarities between the Ring-tailed Lemur
and the bamboo lemurs, particularly in regards to molecular evidence and scent glands similarities,
the true lemurs were moved to the genus Eulemur in 1988.[49][50][91] The genus Lemur is now
monotypic, containing only the Ring-tailed Lemur.
• In 2001, Colin Groves concluded that despite similarities, the Greater Bamboo Lemur was
sufficiently distinct from the bamboo lemurs of the genus Hapalemur to merit its own monotypic
genus, Prolemur.[50][92] This follows Schwarz's 1931 opposition to Pocock's decision to separate
Prolemur from Hapalemur.[67]
 • Originally placed in the genus Microcebus (mouse lemurs), the Giant Mouse Lemur was
moved to its own genus, Mirza, in 1985 due to its larger size, morphological differences, dental
characteristics, and behavior.[47][93]
• The Hairy-eared Dwarf Lemur was first placed in the genus Cheirogaleus (dwarf lemurs) in
1875 and was later found to have closer affinities with Microcebus. However, its dentition and
cranium structure were sufficiently distinct to merit elevation to its own genus, Allocebus.[47][94]

[edit] Species-level classification

Over the past two decades, the number of recognized lemur species has more than doubled
according to some experts. In 1994, 32 distinct species were named in the first edition of Conservation
International's field guide, Lemurs of Madagascar, and 68 were described in the 2nd edition, published in
2006.[58][95] In December 2008, Russell Mittermeier, Colin Groves, and other experts co-wrote an article in
the International Journal of Primatology classifying 99 species and subspecies.[61] The number of lemur
species is likely to continue growing in the coming years, as field studies, cytogenetic and molecular genetic
research continues, particularly on cryptic species, such as mouse lemurs, which cannot be distinguished
visually.[58]
 The Sahamalaza Sportive Lemur (Lepilemur sahamalazensis) was identified as a distinct species as
recently as 2006.
This threefold increase in nearly 15 years has not had universal support among taxonomists and
lemur researchers. In many cases, classifications ultimately depends upon which species concept is used.
Due to the critical condition that most Malagasy primate populations are in, taxonomists and conservationists
sometimes favor splitting them into separate species to develop an effective strategy for the conservation of
the full range of lemur diversity.[58][61] Implicitly, this means that full species status will help grant genetically
distinct populations added environmental protection.[58]
The first large wave of new lemur species descriptions came in 2001 when Colin Groves elevated the
Red Ruffed Lemur (Varecia rubra),[50][96] five subspecies of brown lemur (Eulemur albifrons, E. albocollaris,
E. collaris, E. rufus and E. sanfordi),[97] and four subspecies of sifaka (Propithecus coquereli, P. deckenii,
P. edwardsi, and P. perrieri) to full species status.[98] Additional elevations of all remaining subspecies within
the Eulemur and Propithecus genera were made in the years that followed.[49][50][99] These and
subsequent changes in taxonomy were largely due to a shift to the phylogenetic species concept,[100] yet
are not universally endorsed.[60]
By far the most explosive growth in species numbers has been in the genera Microcebus and
Lepilemur. In 2006, 15 new species of Lepilemur were described, with three new species reported in
February,[101] one species in June,[102] and 11 in September.[103] Since then, two additional species have
been described.[61] Genetic and morphological differences seem to suggest that they are cryptic species,
but there is still debate whether these merit full species status or should be regarded as subspecies of
previously identified, "core" species.[60][86]
 In true lemurs and mouse lemurs, both groups were initially divided into a small number of species,
either with no distinguishable subspecies (in the case of mouse lemurs) or with several distinguishable
subspecies (in the case of true lemurs).[10][37] With molecular research suggesting a more distant split in
both genera, these subspecies or undistinguished populations have been promoted to species status.[37]
In the case of mouse lemurs, the rise in species numbers has been only slightly less sudden and
dramatic. Classified as one species by Ernst Schwarz in 1931 (excluding one, Coquerel's Giant Mouse
Lemur, that is no longer classified in Microcebus),[67] the genus was revised to contain two species, the
Gray Mouse Lemur (Microcebus murinus) and the Brown Mouse Lemur (M. rufus), after an extensive field
study in 1972 showed both living in sympatry in southeastern Madagascar.[104] At the time, the Gray Mouse
Lemur was known in the drier parts of the north, west, and south, while the Brown Mouse Lemur inhabited
the humid rainforest regions of the east. However, we now know the species diversity and distribution to be
significantly more complex.[47] Revisions throughout the 1990s and 2000s identified numerous new species
through genetic testing using mitochondrial DNA, demonstrating that the genus is represented by a multitude
of cryptic species.[61][105][106][107] Many, but not all of these defined species have been supported by
nuclear DNA tests.[108]
However, there are still concerns that species are being identified prematurely. Ian Tattersall, an
anthropologist who recognized 42 species of lemur in 1982,[109] has expressed concern that the
geographically organized variety in lemur populations is being recognized with full species status while the
number of subspecies in lemur genera has virtually disappeared. He has argued that taxonomists are
confusing differentiation and speciation, two processes that are often unrelated, while denying the role of
microevolution in evolutionary processes.[60] Still, other researchers who emphasize the framework of the
"general lineage concept of species" contend that lineage divergence or differentiation demarcates the
beginning of a new species.[108]
 New species have been identified due to differences in morphology, karyotypes, cytochrome b
sequences, and other genetic tests, as well as several combinations of these.[60] When nuclear DNA
(nDNA) was tested in conjunction with mitochondrial DNA (mtDNA) in mouse lemurs, a few species, such as
Claire's Mouse Lemur (Microcebus mamiratra) were demonstrated to be indistinguishable from other closely
related species. In such cases, nDNA did not vary, but the mtDNA that had been used to define it as a
species was still distinct. Differences in results between nDNA, which is inherited from both parents, and
mtDNA, which is inherited from the mother, was attributed to female philopatry, where females remain within
or close to the home range into which they were born while males disperse. Since the isolated population
known as Claire's Mouse Lemur has distinct mtDNA, but not nDNA, it is likely to contain a population
descended from a related group of females, but which still disperses and interbreeds with nearby
populations.[108]
Traditionally, karyology has been considered when determining species status. From the lemurs
studied so far, the diploid number of chromosomes in lemurs varies between 2n=20 and 2n=66. In the case
of the true lemurs, the diploid number ranges from 2n=48 to 2n=60 while the individual chromosome sizes
vary considerably.[77]
Sometimes distinctions are made due to very slight differences in pelage coloration. For instance,
three distinctly-colored types of mouse lemur were discovered in a multi-year study in Beza Mahafaly
Reserve in southern Madagascar, but rather than being separate species, DNA tests revealed that they all
belonged to a single species, the Reddish-gray Mouse Lemur (Microcebus griseorufus).[110] For this reason,
further research is needed to confirm or deny the recent species splits. Only through detailed studies of
morphology, ecology, behavior, and genetics can the true number of lemur species be determined.[58]
Lemur species and subspecies count by year and genus
 1982 1994 2005
Tattersall[109] Mittermeier et al.[111] Groves[83] Mitterm

species subspecies species subspecies species subspecies specie

1 0 1 0 1 0 1

1 2 2 0 3 0 4

2 0 2 0 7 0 7

1 0 1 0 1 0 1

– – 5 8 11 2 10

2 3 3 3 4 2 5

1 0 1 0 1 2 1
4 7 1 0 1 0 1

1 6 7 0 8 0 8

2 0 3 0 8 0 12

1 0 1 0 1 0 2

1 0 1 4 4 0 4

– – – – 1 0 1

2 9 3 8 7 4 9

1 2 1 2 1 4 2

20 29 32 25 59 14 68

42 50 67
[edit] Notes
1. ^ Propithecus diadema holomelas was once considered one of five subspecies of Diademed
Sifaka. In 1986, Ian Tattersall subsumed it as a synonym for what is now known as Milne-Edwards'
Sifaka (known then as Propithecus diadema edwardsi). Both subspecies had only slight color
variations and were known to be sympatric with each other in at least one forest.[52] Since it was
extirpated, the taxonomic status of Propithecus diadema holomelas has been questioned, but
nothing definitive has been published.[53]
2. ^ a b In 1931, Allocebus was not recognized as a separate genus, but was lumped under the
genus Cheirogaleus.
3. ^ a b In 1931 and 1982, Eulemur was not recognized as a separate genus, but was lumped
under the genus Lemur.
4. ^ a b The genus Prolemur was not distinguished until 2001. Prior to this, the Greater Bamboo
Lemur was placed in the genus Hapalemur.
5. ^ a b In 1931, Varecia was not recognized as a separate genus, but was lumped under the
genus Lemur.
6. ^ a b In 1931, Mirza was not recognized as a separate genus, but was lumped under
Microcebus.
7. ^ The grand total of species and subspecies for each year will not equal the sum of the
species total and the subspecies total since each subspecies group already counts as one species.
For example, in 2010 there were three subspecies for Varecia variegata recognized: V. v. variegata,
V. v. editorum, and V. v. subcincta. Together, they count as one species, which is already included in
the species total for that year. The same applies that year for Hapalemur. For this reason there are
101 species and subspecies (97 − 2 + 6 = 101), not 103. In 1994, there were seven species with
subspecies listed, so the grand total is: 32 − 7 + 25 = 50

[edit] References
1. ^ Kay, R.F.; Ross, C.; Williams, B.A. (1997). "Anthropoid origins". Science 275 (5301): 797–
804. doi:10.1126/science.275.5301.797.
2. ^ Bloch, J.I.; Silcox, M.T.; Boyer, D.M.; Sargis, E.J.. "New Paleocene skeletons and the
relationship of plesiadapiforms to crown-clade primates". Proceedings of the National Academy of
Sciences 104 (4): 1159–1164. doi:10.1073/pnas.0610579104.
3. ^ a b c d Janečka, J.E.; Miller, W.; Pringle, T.H.; Wiens, F.; Zitzmann, A.; Helgen, K.M.;
Springer, M.S.; Murphy, W.J. (2007). "Molecular and genomic data identify the closest living relative
of Primates". Science 318 (5851): 792–794. doi:10.1126/science.1147555. PMID 17975064.
http://www.bx.psu.edu/miller_lab/dist/flying_lemur.pdf. Retrieved 11 September 2009.
4. ^ Tavaré, S.; Marshall, C.R.; Will, O.; Soligo, C.; Martin, R.D. (2002). "Using the fossil record
to estimate the age of the last common ancestor of extant primates". Nature 416 (6882): 726–729.
doi:10.1038/416726a. PMID 11961552. Lay summary – ScienceDaily (18 April 2002).
5. ^ a b c d e f g h i j k l m Godinot, M. (2006). "Lemuriform origins as viewed from the fossil
record". Folia Primatologica 77: 446–464. doi:10.1159/000095391.
 6. ^ Williams, B.A.; Kay, R.F.; Kirk, E.C.. "New perspectives on anthropoid origins".
Proceedings of the National Academy of Sciences 107 (11): 4797–4804.
doi:10.1073/pnas.0908320107.
7. ^ Franzen, J.L.; Gingerich, P.D.; Habersetzer, J.; Hurum, J.H.; von Koenigswald, W.; Smith,
B.H. (2009). John, Hawks. ed. "Complete Primate Skeleton from the Middle Eocene of Messel in
Germany: Morphology and Paleobiology". PLoS ONE 4 (5): e5723.
doi:10.1371/journal.pone.0005723. http://www.plosone.org/doi/pone.0005723.
8. ^ Handwerk, B. (19 May 2009). ""Missing link" found: New fossil links humans, lemurs?".
National Geographic News. http://news.nationalgeographic.com/news/2009/05/090519-missing-link-
found.html. Retrieved 12 September 2009.
9. ^ a b c d Gould & Sauther 2006, pp. vii–xiii
10.^ a b c d e f Sussman 2003, pp. 149–229
11.^ Ankel-Simons 2007, pp. 392–514
12.^ a b c d Preston-Mafham 1991, pp. 141–188
13.^ a b c d e f g h i j k Tattersall 2006, pp. 3–18
14.^ Castresana, J. (2001). "Cytochrome b phylogeny and the taxonomy of great apes and
mammals". Molecular Biology and Evolution 18 (4): 465–471. PMID 11264397.
http://mbe.oxfordjournals.org/cgi/reprint/18/4/465.
15.^ a b c d e Yoder 2003, pp. 1242–1247
16.^ a b Simons 1997, pp. 142–166
17.^ a b c d e f g h i j k Yoder, A.D.; Yang, Z. (2004). "Divergence dates for Malagasy lemurs
estimated from multiple gene loci: geological and evolutionary context". Molecular Ecology 13 (4):
757–773. doi:10.1046/j.1365-294X.2004.02106.x. PMID 15012754.
http://www.biology.duke.edu/yoderlab/reprints/2004Yoder&Yang_jme.pdf.
18.^ a b Poux, C.; Madsen, O.; Marquard, E.; Vieites, D.R.; De Jong, W.W.; Vences, M. (2005).
"Asynchronous Colonization of Madagascar by the Four Endemic Clades of Primates, Tenrecs,
Carnivores, and Rodents as Inferred from Nuclear Genes". Systematic Biology 54 (5): 719–730.
doi:10.1080/10635150500234534. http://sysbio.oxfordjournals.org/cgi/reprint/54/5/719.pdf.
19.^ a b c d e f Sussman 2003, pp. 107–148
20.^ Samonds, K.E.; Zalmout, I.S.; Irwin, M.T.; Krause, D.W.; Rogers, R.R.; Raharivony, L.L.
(2009). "Eotheroides lambondrano, new middle Eocene seacow (Mammalia, Sirenia) from the
Mahajanga Basin, northwestern Madagascar". Journal of Vertebrate Paleontology 29 (4): 1233–1243.
doi:10.1671/039.029.0417.
21.^ a b c d Krause 2003, pp. 40–47
22.^ Flynn & Wyss 2003, pp. 34–40
23.^ a b Mittermeier et al. 2006, pp. 23–26
24.^ a b c Goodman, Ganzhorn & Rakotondravony 2003, pp. 1159–1186
25.^ Neild, T. (2007). Supercontinent: Ten Billion Years in the Life of Our Planet . Harvard
University Press. pp. 38–39. ISBN 978-0674026599.
26.^ de Camp, L.S. (1954). Lost Continents (1st ed.). p. 52. ISBN 978-0486226682.
27.^ a b c d e f g h Horvath, J.E.; Weisrock, D.W.; Embry, S.L.; Fiorentino, I.; Balhoff, J.P.;
Kappeler, P.; Wray, G.A.; Willard, H.F. et al. (2008). "Development and application of a
phylogenomic toolkit: resolving the evolutionary history of Madagascar's lemurs". Genome Research
18: 490. doi:10.1101/gr.7265208.
http://www.biology.duke.edu/yoderlab/reprints/2008Horvath_etalGR.pdf.
 28.^ a b Matthew, W.D. (1915). "Climate and evolution". Annals of the New York Academy of
Sciences 24: 171–318. doi:10.1111/j.1749-6632.1914.tb55346.x.
29.^ a b Garbutt 2007, pp. 14–15
30.^ Brumfiel, G. (20 January 2010). "Lemurs' wet and wild past". Nature News.
doi:10.1038/news.2010.23. http://www.nature.com/news/2010/100120/full/news.2010.23.html.
Retrieved 29 January 2010.
31.^ Ali, J.R.; Huber, M. (2010). "Mammalian biodiversity on Madagascar controlled by ocean
currents". Nature 463: 653–656. doi:10.1038/nature08706. Lay summary (21 January 2010).
32.^ Seiffert, E.R.; Simons, E.L.; Attia, Y. (2003). "Fossil evidence for an ancient divergence of
lorises and galagos". Nature 422 (6930): 421–424. doi:10.1038/nature01489. PMID 12660781.
33.^ Krause, Hartman & Wells 1997, pp. 3–43
34.^ a b McCall, R.A. (1997). "Implications of recent geological investigations of the Mozambique
Channel for the mammalian colonization of Madagascar". Proceedings of the Royal Society B 264:
663–665.
35.^ a b c d e f Garbutt 2007, pp. 85–86
36.^ a b Curtis 2006, pp. 133–158
37.^ a b c d Johnson 2006, pp. 187–210
38.^ a b Mittermeier et al. 2006, pp. 89–182
39.^ a b c Mittermeier et al. 2006, pp. 37–51
40.^ Harcourt, C.. "Introduction". Lemurs of Madagascar and the Comoros: The IUCN Red Data
Book. pp. 7–13. ISBN 2880329574.
 41.^ a b Pastorini, J.; Thalmann, U.; Martin, R.D. (2003). "A molecular approach to comparative
phylogeography of extant Malagasy lemurs". Proceedings of the National Academy of Sciences 100
(10): 5879–5884. doi:10.1073/pnas.1031673100. http://www.pnas.org/content/100/10/5879.full.pdf.
42.^ Mittermeier et al. 2006, pp. 15–17
43.^ Preston-Mafham 1991, pp. 10–21
44.^ Godfrey, Jungers & Schwartz 2006, pp. 41–64
45.^ Dunham, A.E.; Rudolf, V.H.W. (2009). "Evolution of sexual size monomorphism: the
influence of passive mate guarding". Journal of Evolutionary Biology 22 (7): 1376–1386.
doi:10.1111/j.1420-9101.2009.01768.x. PMID 19486235. Lay summary – ScienceDaily (1 August
2009).
46.^ a b Gommery, D.; Ramanivosoa, B.; Tombomiadana-Raveloson, S.; Randrianantenaina, H.;
Kerloc'h, P. (2009). "A new species of giant subfossil lemur from the North-West of Madagascar
(Palaeopropithecus kelyus, Primates)". Comptes Rendus Palevol 3 (5): 471–480.
doi:10.1016/j.crpv.2009.02.001. Lay summary (27 May 2009).
47.^ a b c d e Garbutt 2007, pp. 86–114
48.^ a b Mutschler & Tan 2003, pp. 1324–1329
49.^ a b c Mittermeier et al. 2006, pp. 209–323
50.^ a b c d e f Garbutt 2007, pp. 137–175
51.^ "The Top 25 most endangered primates 2006–2008". IUCN/SSC Primate Specialist Group.
http://www.primate-sg.org/T25full07.htm. Retrieved 13 January 2010.
52.^ Pastorini, J.; Forstner, M.R.J.; Martin, R.D. (2001). "Phylogenetic history of sifakas
(Propithecus: Lemuriformes) derived from mtDNA sequences". American Journal of Primatology 53:
1–17. doi:10.1002/1098-2345(200101)53:1<1::AID-AJP1>3.0.CO;2-J.
 53.^ Groves, C.P.; Helgen, K.M. (2007). "Craniodental characters in the taxonomy of
Propithecus". International Journal of Primatology (Springer Netherlands) 28 (6): 1363–1383.
doi:10.1007/s10764-007-9226-5.
54.^ Irwin 2006, pp. 305–326
55.^ Burney 2003, pp. 47–51
56.^ Sussman 2003, pp. 257–269
57.^ a b Godfrey & Jungers 2003, pp. 1247–1252
58.^ a b c d e f g Yoder, A.D. (2007). "Lemurs: a Quick Guide". Current Biology 17 (20): 866–868.
http://www.biology.duke.edu/yoderlab/reprints/2007YoderCB.pdf.
59.^ Groeneveld, L.F.; Weisrock, D.W.; Rasoloarison, R.M.; Yoder, A.D.; Kappeler, P.M. (2009).
"Species delimitation in lemurs: multiple genetic loci reveal low levels of species diversity in the
genus Cheirogaleus". BMC Evolutionary Biology 9 (30). doi:10.1186/1471-2148-9-30.
http://www.biomedcentral.com/1471-2148/9/30. Retrieved 14 January 2010.
60.^ a b c d e f g Tattersall, I. (2007). "Madagascar's lemurs: Cryptic diversity or taxonomic
inflation?". Evolutionary Anthropology 16: 12–23. doi:10.1002/evan.20126.
61.^ a b c d e f g h Mittermeier, R.; Ganzhorn, J.; Konstant, W.; Glander, K.; Tattersall, I.; Groves,
C.; Rylands, A.; Hapke, A. et al. (2008). "Lemur diversity in Madagascar". International Journal of
Primatology 29 (6): 1607–1656. doi:10.1007/s10764-008-9317-y.
http://www.aeecl.org/documents/28.pdf.
62.^ Rowe 1996, p. 13
63.^ Rowe 1996, p. 27
 64.^ a b c Groves, C. (2005). "Strepsirrhini". in Wilson, D. E., & Reeder, D. M, eds. Mammal
Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100002.
65.^ a b Jolly & Sussman 2006, pp. 19–40
66.^ a b Mittermeier et al. 2006, pp. 27–36
67.^ a b c d e f g h Schwarz, E. (1931). "A revision of the genera and species of Madagascar
Lemuridae". Proceedings of the Zoological Society of London 1931: 399–428.
68.^ a b c d e Tattersall 1982, pp. 246–247
69.^ Gregory, W.K. (1915). "1. On the relationships of the Eocene lemur Notharctus to the
Adapidae and other primates. 2. On the classification and phylogeny of the Lemuroidea". Bulletin of
the Geological Society of America 26: 419–446. http://biodiversitylibrary.org/page/4544186#505.
70.^ Simpson, G.G. (1945). Tyler, R.. ed. "The principles of classification and the classification
of mammals". Bulletin of the America Museum of Natural History (New York: Order of the Trustees)
85. http://digitallibrary.amnh.org/dspace/bitstream/2246/1104/1/B085a00.pdf.
71.^ Cartmill 2010, pp. 10–30
72.^ a b c d e Sterling & McCreless 2006, pp. 159–184
73.^ a b Ankel-Simons 2007, pp. 224–283
74.^ Garbutt 2007, pp. 205–207
75.^ Elliot, D.G. (1907). A Catalogue of the Collection of Mammals in the Field Columbian
Museum. Chicago: Harvard University. p. 552. http://books.google.com/books?
id=YnQrAAAAYAAJ&pg=PA552&dq=Sciurus+madagascariensis&cd=2#v=onepage&q=Sciurus
%20madagascariensis&f=false.
 76.^ Owen, R. (1868). Anatomy of Vertebrates. Volume III: Mammals. London: Longmans,
Green, and Co.. http://books.google.com/books?
id=S21XPqd2fF8C&printsec=frontcover&dq=1866+Richard+Owen+On+the+anatomy+of+vertebrates
&source=gbs_similarbooks_s&cad=1#v=onepage&q&f=false.
77.^ a b Ankel-Simons 2007, pp. 533–559
78.^ Groves, C. (2005). "Chiromyiformes". in Wilson, D. E., & Reeder, D. M, eds. Mammal
Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100096.
79.^ a b Yoder, A.D.; Vilgalys, R.; Ruvolo, M. (1996). "Molecular evolutionary dynamics of
cytochrome b in strepsirrhine primates: The phylogenetic significance of third-position transversions" .
Molecular Biology and Evolution 13 (10): 1339–1350. ISSN 0737-4038. PMID 8952078.
http://www.biology.duke.edu/yoderlab/reprints/1996YoderVilgalysMBE.pdf.
80.^ a b Groves 2001, p. 55
81.^ a b c d e Orlando, L.; Calvignac, S.; Schnebelen, C.; Douady, C.J.; Godfrey, L.R.; Hänni, C.
(2008). "DNA from extinct giant lemurs links archaeolemurids to extant indriids". BMC Evolutionary
Biology 8 (121). doi:10.1186/1471-2148-8-121. http://www.biomedcentral.com/1471-2148/8/121.
82.^ a b Tattersall 1982, pp. 263
83.^ a b Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the
World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3.
84.^ Groves 2001, pp. 54–55
 85.^ Groves, C. (2005). "Lepilemuridae". in Wilson, D. E., & Reeder, D. M, eds. Mammal
Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100065.
86.^ a b Garbutt 2007, pp. 115–136
87.^ Karanth, P.; Rakotosamimanana, B.; Parsons, T.J.; Yoder, A.D. (2005). "Ancient DNA from
giant extinct lemurs verifies single origin of Malagasy primates". Proceedings of the National
Academy of Sciences 102: 5090–5095. doi:10.1073/pnas.0408354102.
http://www.biology.duke.edu/yoderlab/reprints/2005Karanth_et_alPNAS.pdf.
88.^ Groves, C. (2005). "Phaner". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species of the
World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100027.
89.^ a b Groves 2001, pp. 65–66
90.^ Groves, C. (2005). "Varecia". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100059.
91.^ Groves, C. (2005). "Eulemur". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100034.
92.^ Groves, C. (2005). "Prolemur". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100057.
93.^ Groves, C. (2005). "Mirza". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species of the
World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100025.
 94.^ Groves, C. (2005). "Allocebus". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species of
the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100006.
95.^ a b Mittermeier et al. 2006, pp. 85–88
96.^ Groves, C. (2005). "Varecia rubra". in Wilson, D. E., & Reeder, D. M, eds. Mammal
Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100060.
97.^ Groves, C. (2005). "Eulemur fulvus". in Wilson, D. E., & Reeder, D. M, eds. Mammal
Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100040.
98.^ Groves, C. (2005). "Propithecus". in Wilson, D. E., & Reeder, D. M, eds. Mammal Species
of the World (3rd ed.). Baltimore: Johns Hopkins University Press. ISBN 0-801-88221-4.
http://www.bucknell.edu/msw3/browse.asp?id=12100084.
99.^ Garbutt 2007, pp. 176–204
100.^ Thalmann, U. (December 2006). "Lemurs – Ambassadors for Madagascar". Madagascar
Conservation & Development 1: 4–8. ISSN 1662-2510. http://www.mwc-
info.net/en/services/Journal_PDF%27s/Issue1/Flagshipspecies.pdf.
101.^ Andriaholinirina, N.; Fausser, J.; Roos, C.; Zinner, D.; Thalmann, U.; Rabarivola, C.;
Ravoarimanana, I.; Ganzhorn, J.U. et al. (2006). "Molecular phylogeny and taxonomic revision of the
sportive lemurs (Lepilemur, Primates)". BMC Evolutionary Biology 6: 17. doi:10.1186/1471-2148-6-
17. http://www.biomedcentral.com/content/pdf/1471-2148-6-17.pdf.
102.^ Rabarivola, C.; Zaramody, A.; Fausser, J.; Andriaholinirina, N.; Roos, C.; Zinner, D.;
Marcel, H.; Rumpler, Y. (2006). "Cytogenetic and molecular characteristics of a new species of
sportive lemur from Northern Madagascar". Lemur News 11: 45–49. ISSN 0343-3528.
http://www.primate-sg.org/PDF/LN11.pdf.
103.^ Louis, Jr., E.E.; Engberg, S.E.; Lei, R.; Geng, H.; Sommer, J.A.; Randriamamapionona,
R.; Randriamanana, J.C.; Zaonarivelo, J.R. et al. (2006). "Molecular and morphological analyses of
the sportive lemurs (Family Megaladapidae: Genus Lepilemur) reveals 11 previously unrecognized
species". Texas Tech University Special Publications (49): 1–49. ISSN 0169-0237.
http://www.nsrl.ttu.edu/publications/opapers/specpubs/SP49.pdf.
104.^ Kappeler & Rasoloarison 2003, pp. 1310–1315
105.^ Braune, P.; Schmidt, S.; Zimmermann, E. (2008). "Acoustic divergence in the
communication of cryptic species of nocturnal primates (Microcebus ssp.)". BMC Biology 6 (19).
doi:10.1186/1741-7007-6-19. http://www.biomedcentral.com/content/pdf/1741-7007-6-19.pdf. Lay
summary – ScienceDaily (14 May 2008).
106.^ Olivieri, G.; Zimmermann, E.; Randrianambinina, B.; Rassoloharijaona, S.;
Rakotondravony, D.; Guschanski, K.; Radespiel, U. (2006). "The ever-increasing diversity in mouse
lemurs: three new species in north and northwestern Madagascar". Molecular Phylogenetics and
Evolution 43 (1): 309–327. doi:10.1016/j.ympev.2006.10.026. PMID 17197200.
107.^ Louis, Jr., E.E.; Engberg, S.E.; McGuire, S.M.; McCormick, M.J.; Randriamampionona, R.;
Ranaivoarisoa, J.F.; Bailey, C.A.; Mittermeier, R.A. et al. (2008). "Revision of the mouse lemurs,
Microcebus (Primates, Lemuriformes), of northern and northwestern Madagascar with descriptions of
two new species at Montagne d'Ambre National Park and Antafondro Classified Forest". Primate
Conservation 23: 19–38. doi:10.1896/052.023.0103. http://www.primate-
sg.org/PDF/PC23.new.microcebus.V3.pdf.
 108.^ a b c Weisrock, D.W.; Rasoloarison, R.M.; Fiorentino, I.; Ralison, J.M.; Goodman, S.M.;
Yoder, A.D. (2010). "Delimiting species without nuclear monophyly in Madagascar's mouse lemurs".
PLoS ONE 5 (3). doi:10.1371/journal.pone.0009883.
109.^ a b Tattersall 1982
110.^ Heckman, K.L.; Rasoazanabary, E.; Machlin, E.; Godfrey, L.R.; Yoder, A.D. (2007).
"MHC-associated mating strategies and the importance of overall genetic diversity in an obligate
pair-living primate". BMC Evolutionary Biology 6 (98): 98. doi:10.1186/1471-2148-6-98.
http://www.biology.duke.edu/yoderlab/reprints/2006heckman_etal.BMCEB.pdf. Lay
summary – ScienceDaily (16 November 2006).
111.^ Mittermeier, R.A.; Tattersall, I.; Konstant, W.R.; Meyers, D.M.; Mast, R.B. (1994). Lemurs
of Madagascar. Illustrated by S.D. Nash (1st ed.). Conservation International. pp. 80–82. ISBN 1-
881173-08-9.
112.^ Mittermeier, R.A.; Louis Jr., E.E.; Richardson, M.; Schwitzer, C.; Langrand, O.; Rylands,
A.B.; Hawkins, F.; Ratsimbazafy, J. et al. (in press). Lemurs of Madagascar. Illustrated by S.D. Nash
(3rd ed.). Arlington, VA: Conservation International.
Books cited
• Ankel-Simons, F. (2007). Primate Anatomy (3rd ed.). Academic Press. ISBN 0-12-372576-3.
• Garbutt, N. (2007). Mammals of Madagascar, A Complete Guide. A&C Black Publishers.
ISBN 978-0-300-12550-4.
• Goodman, S.M.; Benstead, J.P., eds (2003). The Natural History of Madagascar. University
of Chicago Press. ISBN 0-226-30306-3.
 Chapter 2 - Geology and Soils
• Flynn, J.J.; Wyss, A.R. (2003). "Mesozoic Terrestrial Vertebrate Faunas: The Early
History of Madagascar's Vertebrate Diversity". pp. 34–40.
• Krause, D.W. (2003). "Late Cretaceous Vertebrates of Madagascar: A Window into
Gondwanan Biogeography at the End of the Age of Dinosaurs". pp. 40–47.
• Burney, D.A. (2003). "Madagascar's Prehistoric Ecosystems". pp. 47–51.
Chapter 13 - Mammals
• Goodman, S.M.; Ganzhorn, J.U.; Rakotondravony, D. (2003). "Introduction to the
Mammals". pp. 1159–1186.
• Yoder, A.D. (2003). "Phylogeny of the Lemurs". pp. 1242–1247.
• Godfrey, L.R.; Jungers, W.L. (2003). "Subfossil Lemurs". pp. 1247–1252.
• Kappeler, P.M.; Rasoloarison, R.M. (2003). "Microcebus, Mouse Lemurs, Tsidy".
pp. 1310–1315.
• Mutschler, T.; Tan, C.L. (2003). "Hapalemur, Bamboo or Gentle Lemur". pp. 1324–
1329.
• Goodman, S.M.; Patterson, B.D., eds (1997). Natural Change and Human Impact in
Madagascar. Smithsonian Institution Press. ISBN 978-1560986829.
 • Krause, D.W.; Hartman; Wells, N.A. (1997). "Chapter 1: Late Cretaceous Vertebrates
from Madagascar: Implications for Biotic Change in Deep Time". pp. 3–43.
• Simons, E.L. (1997). "Chapter 6: Lemurs: Old and New". pp. 142–166.
• Gould, L.; Sauther, M.L., eds (2006). Lemurs: Ecology and Adaptation. Springer. ISBN 978-
0387-34585-7.
• Gould, L.; Sauther, M.L. (2006). "Preface". pp. vii–xiii.
• Tattersall, I. (2006). "Chapter 1: Origin of the Malagasy Strepsirhine Primates". pp. 3–
18.
• Jolly, A.; Sussman, R.W. (2006). "Chapter 2: Notes on the History of Ecological
Studies of Malagasy Lemurs". pp. 19–40.
• Godfrey, L.R.; Jungers, W.L.; Schwartz, G.T. (2006). "Chapter 3: Ecology and
Extinction of Madagascar's Subfossil Lemurs". pp. 41–64.
• Freed, B.Z. (2006). "Chapter 6: Polyspecific Associations of Crowned Lemurs and
Sanford's Lemurs in Madagascar". pp. 111–132.
• Curtis, D.J. (2006). "Chapter 7: Cathemerality in Lemurs". pp. 133–158.
• Sterling, E.J.; McCreless, E.E. (2006). "Chapter 8: Adaptations in the Aye-aye: A
Review". pp. 159–184.
 • Johnson, S.E. (2006). "Chapter 9: Evolutionary Divergence in the Brown Lemur
Species Complex". pp. 187–210.
• Irwin, M.T. (2006). "Chapter 14: Ecologically Enigmatic Lemurs: The Sifakas of the
Eastern Forests (Propithecus candidus, P. diadema, P. edwardsi, P. perrieri, and P.
tattersalli)". pp. 305–326.
• Groves, C.P. (2001). Primate Taxonomy. Washington, D.C.: Smithsonian Institution Press.
ISBN 978-1560988724.
• Mittermeier, R.A.; Konstant, W.R.; Hawkins, F.; Louis, E.E.; Langrand, O.; Ratsimbazafy, J.;
Rasoloarison, R.; Ganzhorn, J.U. et al. (2006). Lemurs of Madagascar. Illustrated by S.D. Nash (2nd
ed.). Conservation International. ISBN 1-881173-88-7.
• Platt, M.; Ghazanfar, A., eds (2010). Primate Neuroethology. Oxford University Press.
ISBN 978-0195-32659-8.
• Cartmill, M. (2010). "Chapter 2: Primate Classification and Diversity". pp. 10–30.
http://books.google.com/books?
hl=en&lr=&id=hv28p1tCnnEC&oi=fnd&pg=PA10&dq=lemuridae+cladogram&ots=f-w-
l07gyT&sig=OpbCHr0_N82RSJ-Db9TGRvwLhao#v=onepage&q&f=false.
• Preston-Mafham, K. (1991). Madagascar: A Natural History. Facts on File. ISBN 978-
0816024032.
• Rowe, N. (1996). The Pictorial Guide to the Living Primates. Pogonias Press. ISBN 978-
0964882515.
 • Sussman, R.W. (2003). Primate Ecology and Social Structure. Pearson Custom Publishing.
ISBN 978-0536743633.
• Tattersall, I. (1982). Primates of Madagascar. Columbia University Press. ISBN 978-
0231047043.

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Categories: Lemurs | Evolution by taxon

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Evolutionary history of life

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Evolution

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Adaptation
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Biology portal · v • d • e
The evolutionary history of life on Earth traces the processes by which living and fossil organisms
evolved. It stretches from the origin of life on Earth, thought to be over 3,500 million years ago, to the present
day. The similarities between all present day organisms indicate the presence of a common ancestor from
which all known species have diverged through the process of evolution.[1]
Microbial mats of coexisting bacteria and archaea were the dominant form of life in the early Archean
and many of the major steps in early evolution are thought to have taken place within them.[2] The evolution
of oxygenic photosynthesis, around 3,500 million years ago, eventually led to the oxygenation of the
atmosphere, beginning around 2,400 million years ago.[3] The earliest evidence of eukaryotes (complex cells
with organelles), dates from 1,850 million years ago,[4][5] and while they may have been present earlier, their
diversification accelerated when they started using oxygen in their metabolism. Later, around 1,700 million
years ago, multicellular organisms began to appear, with differentiated cells performing specialised functions.
[6]
The earliest land plants date back to around 450 million years ago,[7] though evidence suggests that
algal scum formed on the land as early as 1,200 million years ago. Land plants were so successful that they
are thought to have contributed to the late Devonian extinction event.[8] Invertebrate animals appear during
the Vendian period,[9] while vertebrates originated about 525 million years ago during the Cambrian
explosion.[10]
During the Permian period, synapsids, including the ancestors of mammals, dominated the land,[11]
but the Permian–Triassic extinction event 251 million years ago came close to wiping out all complex life.[12]
During the recovery from this catastrophe, archosaurs became the most abundant land vertebrates,
displacing therapsids in the mid-Triassic.[13] One archosaur group, the dinosaurs, dominated the Jurassic
and Cretaceous periods,[14] with the ancestors of mammals surviving only as small insectivores.[15] After
the Cretaceous–Tertiary extinction event 65 million years ago killed off the non-avian dinosaurs[16] mammals
increased rapidly in size and diversity.[17] Such mass extinctions may have accelerated evolution by
providing opportunities for new groups of organisms to diversify.[18]
Fossil evidence indicates that flowering plants appeared and rapidly diversified in the Early
Cretaceous, between 130 million years ago and 90 million years ago, probably helped by coevolution with
pollinating insects. Flowering plants and marine phytoplankton are still the dominant producers of organic
matter. Social insects appeared around the same time as flowering plants. Although they occupy only small
parts of the insect "family tree", they now form over half the total mass of insects. Humans evolved from a
lineage of upright-walking apes whose earliest fossils date from over 6 million years ago. Although early
members of this lineage had chimp-sized brains, there are signs of a steady increase in brain size after about
3 million years ago.
Contents
[hide]
• 1 Earliest history of Earth
• 2 Earliest evidence for life on Earth
• 3 Origins of life on Earth
• 3.1 Life "seeded" from elsewhere
• 3.2 Independent emergence on Earth
• 3.2.1 Replication first: RNA world
• 3.2.2 Metabolism first: Iron-sulfur world
• 3.2.3 Membranes first: Lipid world
• 3.2.4 The clay theory
• 4 Environmental and evolutionary impact of microbial mats
• 5 Diversification of eukaryotes
• 6 Multicellular organisms and sexual reproduction
• 6.1 Multicellularity
• 6.2 Evolution of sexual reproduction
• 6.3 Fossil evidence for multicellularity and sexual
reproduction
• 7 Emergence of animals
• 8 Colonization of land
• 8.1 Evolution of soil
[edit] Earliest history of Earth

History of Earth and its life

-4500 —
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0—
Hadean
Archean
Protero
-zoic
Phanero
-zoic
Eo
Paleo
Meso
Neo
Paleo
Meso
Neo
Paleo
Meso
Ceno

←
Solar system formed

←
Impact formed Moon

←
? Cool surface, oceans, atmosphere

←
Late Heavy Bombardment

←
? Earliest evidence of life

←
Oxygenation of atmosphere
 ←
Earliest multicellular organism[19]

←
Earliest known fungi

←
Earliest known cnidarians

←
? Cambrian explosion

←
Earliest land invertebrates and plants
 ←
Earliest land vertebrates

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Earliest known dinosaur

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Extinction of non-avian dinosaurs
 Scale:
Millions of years
Main article: History of the Earth
The oldest meteorite fragments found on Earth are about 4,540 million years old, this, coupled
primarily with the dating of ancient lead deposits, has put the estimated age of Earth at around that time.[20]
About 40 million years later a planetoid struck the Earth, throwing into orbit the material that formed the
Moon.[21]
Until recently the oldest rocks found on Earth were about 3,800 million years old,[20] leading
scientists to believe for decades that Earth's surface had been molten until then. Accordingly, they named
this part of Earth's history the Hadean eon, whose name means "hellish".[22] However analysis of zircons
formed 4,400 to 4,000 million years ago indicates that Earth's crust solidified about 100 million years after the
planet's formation and that the planet quickly acquired oceans and an atmosphere, which may have been
capable of supporting life.[23]
Evidence from the Moon indicates that from 4,000 to 3,800 million years ago it suffered a Late Heavy
Bombardment by debris that was left over from the formation of the Solar system, and the Earth should have
experienced an even heavier bombardment due to its stronger gravity.[22][24] While there is no direct
evidence of conditions on Earth 4,000 to 3,800 million years ago, there is no reason to think that the Earth
was not also affected by this late heavy bombardment.[25] This event may well have stripped away any
previous atmosphere and oceans; in this case gases and water from comet impacts may have contributed to
their replacement, although volcanic outgassing on Earth would have contributed at least half.[26]
[edit] Earliest evidence for life on Earth
The earliest identified organisms were minute and relatively featureless, their fossils look like small
rods, which are very difficult to tell apart from structures that arise through abiotic physical processes. The
oldest undisputed evidence of life on Earth, interpreted as fossilized bacteria, dates to 3,000 million years
ago.[27] Other finds in rocks dated to about 3,500 million years ago have been interpreted as bacteria,[28]
with geochemical evidence also seeming to show the presence of life 3,800 million years ago.[29] However
these analyses were closely scrutinized, and non-biological processes were found which could produce all of
the "signatures of life" that had been reported.[30][31] While this does not prove that the structures found had
a non-biological origin, they cannot be taken as clear evidence for the presence of life. Currently, the oldest
unchallenged evidence for life is geochemical signatures from rocks deposited 3,400 million years ago,[27]
[32] although these statements have not been thoroughly examined by critics.
[edit] Origins of life on Earth

Evolutionary tree showing the divergence of modern species from their common ancestor in the
center.[33] The three domains are colored, with bacteria blue, archaea green, and eukaryotes red.
Further information: Evidence of common descent, Common descent, and Homology (biology)
 Biologists reason that all living organisms on Earth must share a single last universal ancestor,
because it would be virtually impossible that two or more separate lineages could have independently
developed the many complex biochemical mechanisms common to all living organisms.[34][35] As previously
mentioned the earliest organisms for which fossil evidence is available are bacteria, cells far too complex to
have arisen directly from non-living materials.[36] The lack of fossil or geochemical evidence for earlier
organisms has left plenty of scope for hypotheses, which fall into two main groups: 1) that life arose
spontaneously on Earth or 2) that it was "seeded" from elsewhere in the universe.[37]

[edit] Life "seeded" from elsewhere

Main articles: Panspermia, Life on Mars, Fermi paradox, and Rare Earth hypothesis
The idea that life on Earth was "seeded" from elsewhere in the universe dates back at least to the
fifth century BC.[38] In the twentieth century it was proposed by the physical chemist Svante Arrhenius,[39]
by the astronomers Fred Hoyle and Chandra Wickramasinghe,[40] and by molecular biologist Francis Crick
and chemist Leslie Orgel.[41] There are three main versions of the "seeded from elsewhere" hypothesis:
from elsewhere in our Solar system via fragments knocked into space by a large meteor impact, in which
case the only credible source is Mars;[42] by alien visitors, possibly as a result of accidental contamination by
micro-organisms that they brought with them;[41] and from outside the Solar system but by natural means.
[39][42] Experiments suggest that some micro-organisms can survive the shock of being catapulted into
space and some can survive exposure to radiation for several days, but there is no proof that they can
survive in space for much longer periods.[42] Scientists are divided over the likelihood of life arising
independently on Mars,[43] or on other planets in our galaxy.[42]
[edit] Independent emergence on Earth
Main article: Abiogenesis
Life on earth is based on carbon and water. Carbon provides stable frameworks for complex
chemicals and can be easily extracted from the environment, especially from carbon dioxide. The only other
element with similar chemical properties, silicon, forms much less stable structures and, because most of its
compounds are solids, would be more difficult for organisms to extract. Water is an excellent solvent and has
two other useful properties: the fact that ice floats enables aquatic organisms to survive beneath it in winter;
and its molecules have electrically negative and positive ends, which enables it to form a wider range of
compounds than other solvents can. Other good solvents, such as ammonia, are liquid only at such low
temperatures that chemical reactions may be too slow to sustain life, and lack water's other advantages.[44]
Organisms based on alternative biochemistry may however be possible on other planets.[45]
Research on how life might have emerged unaided from non-living chemicals focuses on three
possible starting points: self-replication, an organism's ability to produce offspring that are very similar to
itself; metabolism, its ability to feed and repair itself; and external cell membranes, which allow food to enter
and waste products to leave, but exclude unwanted substances.[46] Research on abiogenesis still has a long
way to go, since theoretical and empirical approaches are only beginning to make contact with each other.
[47][48]

[edit] Replication first: RNA world

Main articles: Last universal ancestor and RNA world
 Even the simplest members of the three modern domains of life use DNA
to record their "recipes" and a complex array of RNA and protein molecules to
"read" these instructions and use them for growth, maintenance and self-
replication. This system is far too complex to have emerged directly from non-
living materials.[36] The discovery that some RNA molecules can catalyze both
their own replication and the construction of proteins led to the hypothesis of
earlier life-forms based entirely on RNA.[49] These ribozymes could have formed
an RNA world in which there were individuals but no species, as mutations and
horizontal gene transfers would have meant that the offspring in each generation
were quite likely to have different genomes from those that their parents started
with.[50] RNA would later have been replaced by DNA, which is more stable and
therefore can build longer genomes, expanding the range of capabilities a single
organism can have.[50][51][52] Ribozymes remain as the main components of
ribosomes, modern cells' "protein factories".[53]
Although short self-replicating RNA molecules have been artificially
produced in laboratories,[54] doubts have been raised about where natural non-
biological synthesis of RNA is possible.[55] The earliest "ribozymes" may have
been formed of simpler nucleic acids such as PNA, TNA or GNA, which would
have been replaced later by RNA.[56][57]
In 2003 it was proposed that porous metal sulfide precipitates would
assist RNA synthesis at about 100 °C (212 °F) and ocean-bottom pressures near
The replicator in
virtually all known life is
deoxyribonucleic acid.
DNA's structure and
replication systems are
far more complex than
those of the original
replicator.[36]
hydrothermal vents. In this hypothesis lipid membranes would be the last major cell components to appear
and until then the proto-cells would be confined to the pores.[58]

[edit] Metabolism first: Iron-sulfur world

Main article: Iron-sulfur world theory
A series of experiments starting in 1997 showed that early stages in the formation of proteins from
inorganic materials including carbon monoxide and hydrogen sulfide could be achieved by using iron sulfide
and nickel sulfide as catalysts. Most of the steps required temperatures of about 100 °C (212 °F) and
moderate pressures, although one stage required 250 °C (482 °F) and a pressure equivalent to that found
under 7 kilometres (4.3 mi) of rock. Hence it was suggested that self-sustaining synthesis of proteins could
have occurred near hydrothermal vents.[59]

[edit] Membranes first: Lipid world

It has been suggested that double-walled "bubbles" of
lipids like those that form the external membranes of cells may = water-attracting heads of lipid
have been an essential first step.[60] Experiments that
simulated the conditions of the early Earth have reported the = water-repellent tails
formation of lipids, and these can spontaneously form
liposomes, double-walled "bubbles", and then reproduce
themselves. Although they are not intrinsically information-
carriers as nucleic acids are, they would be subject to natural

Cross-section through a liposome.

selection for longevity and reproduction. Nucleic acids such as RNA might then have formed more easily
within the liposomes than they would have outside.[61]

[edit] The clay theory

Main articles: Graham Cairns-Smith#Clay Theory and RNA world
RNA is complex and there are doubts about whether it can be produced non-biologically in the wild.
[55] Some clays, notably montmorillonite, have properties that make them plausible accelerators for the
emergence of an RNA world: they grow by self-replication of their crystalline pattern; they are subject to an
analog of natural selection, as the clay "species" that grows fastest in a particular environment rapidly
becomes dominant; and they can catalyze the formation of RNA molecules.[62] Although this idea has not
become the scientific consensus, it still has active supporters.[63]
Research in 2003 reported that montmorillonite could also accelerate the conversion of fatty acids
into "bubbles", and that the "bubbles" could encapsulate RNA attached to the clay. These "bubbles" can then
grow by absorbing additional lipids and then divide. The formation of the earliest cells may have been aided
by similar processes.[64]
A similar hypothesis presents self-replicating iron-rich clays as the progenitors of nucleotides, lipids
and amino acids.[65]

[edit] Environmental and evolutionary impact of microbial mats

Main articles: Microbial mat and Oxygen catastrophe
 Modern stromatolites in Shark Bay, Western Australia.
Microbial mats are multi-layered, multi-species colonies of bacteria and other organisms that are
generally only a few millimeters thick, but still contain a wide range of chemical environments, each of which
favors a different set of micro-organisms.[66] To some extent each mat forms its own food chain, as the by-
products of each group of micro-organisms generally serve as "food" for adjacent groups.[67]
Stromatolites are stubby pillars built as microbes in mats slowly migrate upwards to avoid being
smothered by sediment deposited on them by water.[66] There has been vigorous debate about the validity
of alleged fossils from before 3,000 million years ago,[68] with critics arguing that so-called stromatolites
could have been formed by non-biological processes.[30] In 2006 another find of stromatolites was reported
from the same part of Australia as previous ones, in rocks dated to 3,500 million years ago.[69]
In modern underwater mats the top layer often consists of photosynthesizing cyanobacteria which
create an oxygen-rich environment, while the bottom layer is oxygen-free and often dominated by hydrogen
sulfide emitted by the organisms living there.[67] It is estimated that the appearance of oxygenic
photosynthesis by bacteria in mats increased biological productivity by a factor of between 100 and 1,000.
The reducing agent used by oxygenic photosynthesis is water, which is much more plentiful than the
geologically-produced reducing agents required by the earlier non-oxygenic photosynthesis.[70] From this
point onwards life itself produced significantly more of the resources it needed than did geochemical
processes.[71] Oxygen is toxic to organisms that are not adapted to it, but greatly increases the metabolic
efficiency of oxygen-adapted organisms.[72][73]
Oxygen became a significant component of Earth's atmosphere about 2,400 million years ago.[74]
Although eukaryotes may have been present much earlier,[75][76] the oxygenation of the atmosphere was a
prerequisite for the evolution of the most complex eukaryotic cells, from which all multicellular organisms are
built.[77] The boundary between oxygen-rich and oxygen-free layers in microbial mats would have moved
upwards when photosynthesis shut down overnight, and then downwards as it resumed on the next day. This
would have created selection pressure for organisms in this intermediate zone to acquire the ability to
tolerate and then to use oxygen, possibly via endosymbiosis, where one organism lives inside another and
both of them benefit from their association.[2]
Cyanobacteria have the most complete biochemical "toolkits" of all the mat-forming organisms.
Hence they are the most self-sufficient of the mat organisms and were well-adapted to strike out on their own
both as floating mats and as the first of the phytoplankton, providing the basis of most marine food chains.[2]
 Eukaryotes
Bikonta

Apusozoa

[edit] Diversification of eukaryotes Archaeplastida (Land plants

Main article: Eukaryote green algae, red algae, and
glaucophytes)
Eukaryotes may have been present long before the
oxygenation of the atmosphere,[75] but most modern eukaryotes
Chromalveolata
require oxygen, which their mitochondria use to fuel the
production of ATP, the internal energy supply of all known cells.
[77] In the 1970s it was proposed and, after much debate, widely
accepted that eukaryotes emerged as a result of a sequence of Rhizaria
endosymbioses between "procaryotes". For example: a
predatory micro-organism invaded a large procaryote, probably
an archaean, but the attack was neutralized, and the attacker Excavata
took up residence and evolved into the first of the mitochondria;
one of these chimeras later tried to swallow a photosynthesizing
cyanobacterium, but the victim survived inside the attacker and
the new combination became the ancestor of plants; and so on.
After each endosymbiosis began, the partners would have Unikonta
eliminated unproductive duplication of genetic functions by re-
arranging their genomes, a process which sometimes involved Amoebozoa
transfer of genes between them.[80][81][82] Another hypothesis
proposes that mitochondria were originally sulfur- or hydrogen- Opisthokonta
metabolising endosymbionts, and became oxygen-consumers
later.[83] On the other hand mitochondria might have been part Metazoa
of eukaryotes' original equipment.[84] (Animals)

Choanozoa

Eumycota
(Fungi)
 There is a debate about when eukaryotes first appeared: the presence of steranes in Australian
shales may indicate that eukaryotes were present 2,700 million years ago;[76] however an analysis in 2008
concluded that these chemicals infiltrated the rocks less than 2,200 million years ago and prove nothing
about the origins of eukaryotes.[85] Fossils of the alga Grypania have been reported in 1,850 million-year-old
rocks (originally dated to 2,100 million years ago but later revised[5]), and indicates that eukaryotes with
organelles had already evolved.[86] A diverse collection of fossil algae were found in rocks dated between
1,500 million years ago and 1,400 million years ago.[87] The earliest known fossils of fungi date from 1,430
million years ago.[88]

[edit] Multicellular organisms and sexual reproduction

[edit] Multicellularity
Main articles: Multicellular organism , Evolution of multicellularity , and Sexual reproduction

A slime mold solves a maze. The mold (yellow) explored and filled the maze (left). When the
researchers placed sugar (red) at two separate points, the mold concentrated most of its mass there and left
only the most efficient connection between the two points (right).[89]
 The simplest definitions of "multicellular", for example "having multiple cells", could include colonial
cyanobacteria like Nostoc. Even a professional biologist's definition such as "having the same genome but
different types of cell" would still include some genera of the green alga Volvox, which have cells that
specialize in reproduction.[90] Multicellularity evolved independently in organisms as diverse as sponges and
other animals, fungi, plants, brown algae, cyanobacteria, slime moulds and myxobacteria.[5][91] For the sake
of brevity this article focuses on the organisms that show the greatest specialization of cells and variety of
cell types, although this approach to the evolution of complexity could be regarded as "rather
anthropocentric".[92]
The initial advantages of multicellularity may have included: increased resistance to predators, many
of which attacked by engulfing; the ability to resist currents by attaching to a firm surface; the ability to reach
upwards to filter-feed or to obtain sunlight for photosynthesis;[93] the ability to create an internal environment
that gives protection against the external one;[92] and even the opportunity for a group of cells to behave
"intelligently" by sharing information.[89] These features would also have provided opportunities for other
organisms to diversify, by creating more varied environments than flat microbial mats could.[93]
Multicellularity with differentiated cells is beneficial to the organism as a whole but disadvantageous
from the point of view of individual cells, most of which lose the opportunity to reproduce themselves. In an
asexual multicellular organism, rogue cells which retain the ability to reproduce may take over and reduce the
organism to a mass of undifferentiated cells. Sexual reproduction eliminates such rogue cells from the next
generation and therefore appears to be a prerequisite for complex multicellularity.[93]
The available evidence indicates that eukaryotes evolved much earlier but remained inconspicuous
until a rapid diversification around 1,000 million years ago. The only respect in which eukaryotes clearly
surpass bacteria and archaea is their capacity for variety of forms, and sexual reproduction enabled
eukaryotes to exploit that advantage by producing organisms with multiple cells that differed in form and
function.[93]

[edit] Evolution of sexual reproduction

Main article: Evolution of sexual reproduction
The defining characteristic of sexual reproduction is recombination, in which each of the offspring
receives 50% of its genetic inheritance from each of the parents.[94] Bacteria also exchange DNA by
bacterial conjugation, the benefits of which include resistance to antibiotics and other toxins, and the ability to
utilize new metabolites.[95] However conjugation is not a means of reproduction, and is not limited to
members of the same species – there are cases where bacteria transfer DNA to plants and animals.[96]
The disadvantages of sexual reproduction are well-known: the genetic reshuffle of recombination
may break up favorable combinations of genes; and since males do not directly increase the number of
offspring in the next generation, an asexual population can out-breed and displace in as little as 50
generations a sexual population that is equal in every other respect.[94] Nevertheless the great majority of
animals, plants, fungi and protists reproduce sexually. There is strong evidence that sexual reproduction
arose early in the history of eukaryotes and that the genes controlling it have changed very little since then.
[97] How sexual reproduction evolved and survived is an unsolved puzzle.[98]
The Red Queen Hypothesis suggests that sexual reproduction provides protection against parasites,
because it is easier for parasites to evolve means of overcoming the defenses of genetically identical clones
than those of sexual species that present moving targets, and there is some experimental evidence for this.
However there is still doubt about whether it would explain the survival of sexual species if multiple similar
clone species were present, as one of the clones may survive the attacks of parasites for long enough to out-
breed the sexual species.[94]
The Mutation Deterministic Hypothesis assumes that each organism has more than one harmful
mutation and the combined effects of these mutations are more harmful than the sum of the harm done by
each individual mutation. If so, sexual recombination of genes will reduce the harm that bad mutations do to
offspring and at the same time eliminate some bad mutations from the gene pool by isolating them in
individuals that perish quickly because they have an above-average number of bad mutations. However the
evidence suggests that the MDH's assumptions are shaky, because many species have on average less
than one harmful mutation per individual and no species that has been investigated shows evidence of
synergy between harmful mutations.[94]

Horodyskia apparently re-arranged itself into fewer but larger main masses as the sediment grew
deeper round its base.[5]
 The random nature of recombination causes the relative abundance of alternative traits to vary from
one generation to another. This genetic drift is insufficient on its own to make sexual reproduction
advantageous, but a combination of genetic drift and natural selection may be sufficient. When chance
produces combinations of good traits, natural selection gives a large advantage to lineages in which these
traits become genetically linked. On the other hand the benefits of good traits are neutralized if they appear
along with bad traits. Sexual recombination gives good traits the opportunities to become linked with other
good traits, and mathematical models suggest this may be more than enough to offset the disadvantages of
sexual reproduction.[98] Other combinations of hypotheses that are inadequate on their own are also being
examined.[94]
The following hypotheses attempt to explain how and why sex evolved:
• It may have enabled organisms to repair genetic damage.[99] The most primitive form of sex
may have been one organism repairing damaged DNA by replicating an undamaged strand from a
similar organism.[100]
• Sexual reproduction may have originated from selfish parasitic genetic elements propagating
themselves by transfer to new hosts.[101]
• It may have evolved from cannibalism, where some of the victim's DNA was incorporated into
the cannibal organism.[100]
• Sexual reproduction may have evolved from ancient haloarchaea through a combination of
jumping genes, and swapping plasmids.[102]
• Or it may have evolved as a form of vaccination in which infected hosts exchanged
weakened symbiotic copies of parasitic DNA as protection against more virulent versions. The
meiosis stage of sexual reproduction may then have evolved as a way of removing the symbiotes.
[103]
 Bacteria also exchange DNA by bacterial conjugation, the benefits of which include resistance to
antibiotics and other toxins, and the ability to utilize new metabolites.[95] However conjugation is not a
means of reproduction and is not limited to members of the same species, and there are cases where
bacteria transfer DNA to plants and animals.[96] Nevertheless it may be an example of the "selfish genetic
element" hypothesis, as it transfers DNA by means of such a "selfish gene", the F-plasmid.[100]

[edit] Fossil evidence for multicellularity and sexual reproduction

The Francevillian Group Fossil, dated to 2,100 million years ago, is the
earliest known fossil organism that is clearly multicellular.[19] They may have
had differentiated cells.[105] Another early multicellular fossil, Qingshania,
[note 1] dated to 1,700 million years ago, appears to consist of virtually
identical cells. The red alga called Bangiomorpha, dated at 1,200 million years
ago, is the earliest known organism which certainly has differentiated,
specialized cells, and is also the oldest known sexually-reproducing organism.
[93] The 1,430 million-year-old fossils interpreted as fungi appear to have been
multicellular with differentiated cells.[88] The "string of beads" organism
Horodyskia, found in rocks dated from 1,500 million years ago to 900 million
years ago, may have been an early metazoan;[5] however it has also been
interpreted as a colonial foraminiferan.[104]
Horodyskia may
[edit] Emergence of animals have been an early
Main articles: Animal, Ediacara biota, Cambrian Explosion, Burgess metazoan,[5] or a colonial
shale type fauna, and Stem group foraminiferan[104]
 Animals are multicellular eukaryotes,[note 2] and are Bilaterians
distinguished from plants, algae, and fungi by lacking cell walls.
[107] All animals are motile,[108] if only at certain life stages. All Deuterostomes
animals except sponges have bodies differentiated into separate (chordates
tissues, including muscles, which move parts of the animal by hemichordates
contracting, and nerve tissue, which transmits and processes echinoderms
signals.[109] Protostomes
The earliest widely-accepted animal fossils are rather Ecdysozoa
modern-looking cnidarians (the group that includes jellyfish, sea (arthropods,
anemones and hydras), possibly from around 580 million years nematodes, tardigrades,
ago, although fossils from the Doushantuo Formation can only etc.)
be dated approximately. Their presence implies that the
cnidarian and bilaterian lineages had already diverged.[110] Lophotrochozoa
(molluscs, annelids,
The Ediacara biota, which flourished for the last brachiopods, etc.)
40 million years before the start of the Cambrian,[111] were the
first animals more than a very few centimeters long. Many were
flat and had a "quilted" appearance, and seemed so strange that
there was a proposal to classify them as a separate kingdom,
Vendozoa.[112] Others, however, been interpreted as early
molluscs (Kimberella[113][114]), echinoderms (Arkarua[115]),
and arthropods (Spriggina,[116] Parvancorina[117]). There is
still debate about the classification of these specimens, mainly
because the diagnostic features which allow taxonomists to
classify more recent organisms, such as similarities to living organisms, are generally absent in the
Ediacarans. However there seems little doubt that Kimberella was at least a triploblastic bilaterian animal, in
other words significantly more complex than cnidarians.[118]
The small shelly fauna are a very mixed collection of fossils found between the Late Ediacaran and
Mid Cambrian periods. The earliest, Cloudina, shows signs of successful defense against predation and may
indicate the start of an evolutionary arms race. Some tiny Early Cambrian shells almost certainly belonged to
molluscs, while the owners of some "armor plates", Halkieria and Microdictyon, were eventually identified
when more complete specimens were found in Cambrian lagerstätten that preserved soft-bodied animals.
[119]

Opabinia made the largest single contribution to modern interest in the Cambrian explosion.[120]
In the 1970s there was already a debate about whether the emergence of the modern phyla was
"explosive" or gradual but hidden by the shortage of Pre-Cambrian animal fossils.[119] A re-analysis of
fossils from the Burgess Shale lagerstätte increased interest in the issue when it revealed animals, such as
Opabinia, which did not fit into any known phylum. At the time these were interpreted as evidence that the
modern phyla had evolved very rapidly in the "Cambrian explosion" and that the Burgess Shale's "weird
wonders" showed that the Early Cambrian was a uniquely experimental period of animal evolution.[121] Later
discoveries of similar animals and the development of new theoretical approaches led to the conclusion that
many of the "weird wonders" were evolutionary "aunts" or "cousins" of modern groups[122] – for example that
Opabinia was a member of the lobopods, a group which includes the ancestors of the arthropods, and that it
may have been closely related to the modern tardigrades.[123] Nevertheless there is still much debate about
whether the Cambrian explosion was really explosive and, if so, how and why it happened and why it
appears unique in the history of animals.[124]

Acanthodians were among the earliest vertebrates with jaws[125]

Most of the animals at the heart of the Cambrian explosion debate are protostomes, one of the two
main groups of complex animals. One deuterostome group, the echinoderms, many of which have hard
calcite "shells", are fairly common from the Early Cambrian small shelly fauna onwards.[119] Other
deuterostome groups are soft-bodied, and most of the significant Cambrian deuterostome fossils come from
the Chengjiang fauna, a lagerstätte in China.[126] The Chengjiang fossils Haikouichthys and Myllokunmingia
appear to be true vertebrates,[127] and Haikouichthys had distinct vertebrae, which may have been slightly
mineralized.[128] Vertebrates with jaws, such as the Acanthodians, first appeared in the Late Ordovician.
[129]
[edit] Colonization of land
Adaptation to life on land is a major challenge: all land organisms need to avoid drying-out and all
those above microscopic size have to resist gravity; respiration and gas exchange systems have to change;
reproductive systems cannot depend on water to carry eggs and sperm towards each other.[130][131]
Although the earliest good evidence of land plants and animals dates back to the Ordovician period
(488 to 444 million years ago), modern land ecosystems only appeared in the late Devonian, about
385 to 359 million years ago.[132]

[edit] Evolution of soil

Before the colonization of land, soil, a combination of mineral particles and decomposed organic
matter, did not exist. Land surfaces would have been either bare rock or unstable sand produced by
weathering. Water and any nutrients in it would have drained away very quickly.[132]

Lichens growing on concrete

 Films of cyanobacteria, which are not plants but use the same photosynthesis mechanisms, have
been found in modern deserts, and only in areas that are unsuitable for vascular plants. This suggests that
microbial mats may have been the first organisms to colonize dry land, possibly in the Precambrian. Mat-
forming cyanobacteria could have gradually evolved resistance to desiccation as they spread from the seas
to tidal zones and then to land.[132] Lichens, which are symbiotic combinations of a fungus (almost always
an ascomycete) and one or more photosynthesizers (green algae or cyanobacteria),[133] are also important
colonizers of lifeless environments,[132] and their ability to break down rocks contributes to soil formation in
situations where plants cannot survive.[133] The earliest known ascomycete fossils date from 423 to 419
million years ago in the Silurian.[132]
Soil formation would have been very slow until the appearance of burrowing animals, which mix the
mineral and organic components of soil and whose feces are a major source of the organic components.
[132] Burrows have been found in Ordovician sediments, and are attributed to annelids ("worms") or
arthropods.[132][134]

[edit] Plants and the Late Devonian wood crisis

Main article: Evolutionary history of plants
Reconstruction of Cooksonia, a vascular plant from the Silurian.
 Fossilized trees from the Mid-Devonian Gilboa fossil forest.
In aquatic algae, almost all cells are capable of photosynthesies and are nearly independent. Life on
land required plants to become internally more complex and specialized: photosynthesis was most efficient
at the top; roots were required in order to extract water from the ground; the parts in between became
supports and transport systems for water and nutrients.[130][135]
Spores of land plants, possibly rather like liverworts, have been found in Mid Ordovician rocks dated
to about 476 million years ago. In Mid Silurian rocks 430 million years ago there are fossils of actual plants
including clubmosses such as Baragwanathia; most were under 10 centimetres (3.9 in) high, and some
appear closely related to vascular plants, the group that includes trees.[135]
By the Late Devonian 370 million years ago, trees such as Archaeopteris were so abundant that they
changed river systems from mostly braided to mostly meandering, because their roots bound the soil firmly.
[136] In fact they caused a "Late Devonian wood crisis",[137] because:
• They removed more carbon dioxide from the atmosphere, reducing the greenhouse effect
and thus causing an ice age in the Carboniferous period.[138] In later ecosystems the carbon dioxide
"locked up" in wood is returned to the atmosphere by decomposition of dead wood. However the
earliest fossil evidence of fungi that can decompose wood also comes from the Late Devonian.[139]
• The increasing depth of plants' roots led to more washing of nutrients into rivers and seas by
rain. This caused algal blooms whose high consumption of oxygen caused anoxic events in deeper
waters, increasing the extinction rate among deep-water animals.[138]
 [edit] Land invertebrates
Animals had to change their feeding and excretory systems, and most land animals developed
internal fertilization of their eggs. The difference in refractive index between water and air required changes
in their eyes. On the other hand in some ways movement and breathing became easier, and the better
transmission of high-frequency sounds in air encouraged the development of hearing.[131]
Some trace fossils from the Cambrian-Ordovician boundary about 490 million years ago are
interpreted as the tracks of large amphibious arthropods on coastal sand dunes, and may have been made
by euthycarcinoids,[140] which are thought to be evolutionary "aunts" of myriapods.[141] Other trace fossils
from the Late Ordovician a little over 445 million years ago probably represent land invertebrates, and there
is clear evidence of numerous arthropods on coasts and alluvial plains shortly before the Silurian-Devonian
boundary, about 415 million years ago, including signs that some arthropods ate plants.[142] Arthropods
were well pre-adapted to colonise land, because their existing jointed exoskeletons provided protection
against desiccation, support against gravity and a means of locomotion that was not dependent on water.
[143]
The fossil record of other major invertebrate groups on land is poor: none at all for non-parasitic
flatworms, nematodes or nemerteans; some parasitic nematodes have been fossilized in amber; annelid
worm fossils are known from the Carboniferous, but they may still have been aquatic animals; the earliest
fossils of gastropods on land date from the Late Carboniferous, and this group may have had to wait until leaf
litter became abundant enough to provide the moist conditions they need.[131]
The earliest confirmed fossils of flying insects date from the Late Carboniferous, but it is thought that
insects developed the ability to fly in the Early Carboniferous or even Late Devonian. This gave them a wider
range of ecological niches for feeding and breeding, and a means of escape from predators and from
unfavorable changes in the environment.[144] About 99% of modern insect species fly or are descendants of
flying species.[145]

[edit] Land vertebrates

Main article: Tetrapod

Acanthostega changed views about the early evolution of tetrapods[146]

"Fish"

Acanthostega
 Ichthyostega

Tulerpeton

Early
amphibians

Anthracosauria

Amniotes
 Family tree of tetrapods[147]
Tetrapods, vertebrates with four limbs, evolved from other rhipidistians over a relatively short
timespan during the Late Devonian, between 370 million years ago and 360 million years ago.[148] From the
1950s to the early 1980s it was thought that tetrapods evolved from fish that had already acquired the ability
to crawl on land, possibly in order to go from a pool that was drying out to one that was deeper. However in
1987 nearly-complete fossils of Acanthostega from about 363 million years ago showed that this Late
Devonian transitional animal had legs and both lungs and gills, but could never have survived on land: its
limbs and its wrist and ankle joints were too weak to bear its weight; its ribs were too short to prevent its
lungs from being squeezed flat by its weight; its fish-like tail fin would have been damaged by dragging on
the ground. The current hypothesis is that Acanthostega, which was about 1 metre (3.3 ft) long, was a wholly
aquatic predator that hunted in shallow water. Its skeleton differed from that of most fish, in ways that
enabled it to raise its head to breathe air while its body remained submerged, including: its jaws show
modifications that would have enabled it to gulp air; the bones at the back of its skull are locked together,
providing strong attachment points for muscles that raised its head; the head is not joined to the shoulder
girdle and it has a distinct neck.[146]
The Devonian proliferation of land plants may help to explain why air-breathing would have been an
advantage: leaves falling into streams and rivers would have encouraged the growth of aquatic vegetation;
this would have attracted grazing invertebrates and small fish that preyed on them; they would have been
attractive prey but the environment was unsuitable for the big marine predatory fish; air-breathing would have
been necessary because these waters would have been short of oxygen, since warm water holds less
dissolved oxygen than cooler marine water and since the decomposition of vegetation would have used
some of the oxygen.[146]
Later discoveries revealed earlier transitional forms between Acanthostega and completely fish-like
animals.[149] Unfortunately there is then a gap of about 30 million years between the fossils of ancestral
tetrapods and Mid Carboniferous fossils of vertebrates that look well-adapted for life on land. Some of these
look like early relatives of modern amphibians, most of which need to keep their skins moist and to lay their
eggs in water, while others are accepted as early relatives of the amniotes, whose water-proof skins and
eggs enable them to live and breed far from water.[147]
[edit] Dinosaurs, birds and mammals
Main articles: Dinosaur evolution, Origin of Birds, and Evolution of mammals
Amniotes
Synapsids
Early synapsids
(extinct)

Pelycosaurs

Extinct pelycosaurs

Therapsids

Extinct therapsids

Mammaliformes

Extinct mammaliformes

Mammals
 Sauropsids

Diapsids
 Squ
(lizards and
snakes)
Archosaurs
Extinct
archosaurs

Crocodilians

Pterosaurs
(extinct)

Dinosaurs
Theropods
Extinct
theropods
Birds

Sauropods
(extinct)

Ornithischians
(extinct)
 Possible family tree of dinosaurs, birds and mammals[151][152]
Amniotes, whose eggs can survive in dry environments, probably evolved in the Late Carboniferous
period, between 330 million years ago and 314 million years ago. The earliest fossils of the two surviving
amniote groups, synapsids and sauropsids, date from around 313 million years ago.[151][152] The synapsid
pelycosaurs and their descendants the therapsids are the most common land vertebrates in the best-known
Permian fossil beds, between 229 million years ago and 251 million years ago. However at the time these
were all in temperate zones at middle latitudes, and there is evidence that hotter, drier environments nearer
the Equator were dominated by sauropsids and amphibians.[153]
The Permian-Triassic extinction wiped out almost all land vertebrates,[154] as well as the great
majority of other life.[155] During the slow recovery from this catastrophe, estimated to be 30M years,[156] a
previously obscure sauropsid group became the most abundant and diverse terrestrial vertebrates: a few
fossils of archosauriformes ("shaped like archosaurs") have been found in Late Permian rocks,[157] but by
the Mid Triassic archosaurs were the dominant land vertebrates. Dinosaurs distinguished themselves from
other archosaurs in the Late Triassic, and became the dominant land vertebrates of the Jurassic and
Cretaceous periods, between 199 million years ago and 65 million years ago.[158]
During the Late Jurassic, birds evolved from small, predatory theropod dinosaurs.[159] The first birds
inherited teeth and long, bony tails from their dinosaur ancestors,[159] but some developed horny, toothless
beaks by the very Late Jurassic[160] and short pygostyle tails by the Early Cretaceous.[161]
While the archosaurs and dinosaurs were becoming more dominant in the Triassic, the mammaliform
successors of the therapsids could only survive as small, mainly nocturnal insectivores. This apparent set-
back may actually have promoted the evolution of mammals, for example nocturnal life may have
accelerated the development of endothermy ("warm-bloodedness") and hair or fur.[162] By 195 million years
ago in the Early Jurassic there were animals that were very nearly mammals.[163] Unfortunately there is a
gap in the fossil record throughout the Mid Jurassic.[164] However fossil teeth discovered in Madagascar
indicate that true mammals existed at least 167 million years ago.[165] After dominating land vertebrate
niches for about 150 million years, the dinosaurs perished 65 million years ago in the Cretaceous–Tertiary
extinction along with many other groups of organisms.[166] Mammals throughout the time of the dinosaurs
had been restricted to a narrow range of taxa, sizes and shapes, but increased rapidly in size and diversity
after the extinction,[167][168] with bats taking to the air within 13 million years,[169] and cetaceans to the sea
within 15 million years.[170]

[edit] Flowering plants

Main articles: Flowering plant and Gymnosperm
 Gymnosperms Gymnosp
Gnetales
(gymnosperm)

Welwitschia Cycads and

The 250,000 to 400,000 species of flowering plants outnumber all other ground plants combined,
(gymnosperm) (gymnosperm)

Bennettitales

Ephedra
(gymnosperm)

Gingk

Bennettitales

Angiosperms Gnetales
(flowering plants) (gymnosperm)

Conifers
(gymnosperm)

One possible family tree of flowering plants.[171]

Another possible family tree.[172]

[edit] Social insects
Main article: Social insects
The social insects are remarkable because the great majority of individuals in each colony are sterile.
This appears contrary to basic concepts of evolution such as natural selection and the selfish gene. In fact
there are very few eusocial insect species: only 15 out of approximately 2,600 living families of insects
contain eusocial species, and it seems that eusociality has evolved independently only 12 times among
arthropods, although some eusocial lineages have diversified into several families. Nevertheless social
insects have been spectacularly successful; for example although ants and termites account for only about
2% of known insect species, they form over 50% of the total mass of insects. Their ability to control a territory
appears to be the foundation of their success.[173]
 These termite mounds have survived a bush fire.
The sacrifice of breeding opportunities by most individuals has long been explained as a
consequence of these species' unusual haplodiploid method of sex determination, which has the paradoxical
consequence that two sterile worker daughters of the same queen share more genes with each other than
they would with their offspring if they could breed.[174] However Wilson and Hölldobler argue that this
explanation is faulty: for example, it is based on kin selection, but there is no evidence of nepotism in
colonies that have multiple queens. Instead, they write, eusociality evolves only in species that are under
strong pressure from predators and competitors, but in environments where it is possible to build "fortresses";
after colonies have established this security, they gain other advantages though co-operative foraging. In
support of this explanation they cite the appearance of eusociality in bathyergid mole rats,[173] which are not
haplodiploid.[175]
The earliest fossils of insects have been found in Early Devonian rocks from about 400 million years
ago, which preserve only a few varieties of flightless insect. The Mazon Creek lagerstätten from the Late
Carboniferous, about 300 million years ago, include about 200 species, some gigantic by modern standards,
and indicate that insects had occupied their main modern ecological niches as herbivores, detritivores and
insectivores. Social termites and ants first appear in the Early Cretaceous, and advanced social bees have
been found in Late Cretaceous rocks but did not become abundant until the Mid Cenozoic.[176]

[edit] Humans
Main article: Human evolution
Modern humans evolved from a lineage of upright-walking apes that has been traced back over 6
million years ago to Sahelanthropus.[177] The first known stone tools were made about 2.5 million years ago,
apparently by Australopithecus garhi, and were found near animal bones that bear scratches made by these
tools.[178] The earliest hominines had chimp-sized brains, but there has been a fourfold increase in the last
3 million years; a statistical analysis suggests that hominine brain sizes depend almost completely on the
date of the fossils, while the species to which they are assigned has only slight influence.[179] There is a
long-running debate about whether modern humans evolved all over the world simultaneously from existing
advanced hominines or are descendants of a single small population in Africa, which then migrated all over
the world less than 200,000 years ago and
replaced previous hominine species.[180] There is
also debate about whether anatomically-modern
humans had an intellectual, cultural and
technological "Great Leap Forward" under
100,000 years ago and, if so, whether this was
due to neurological changes that are not visible in
fossils.[181]

[edit] Mass extinctions

Main article: Mass extinction
Life on earth has suffered occasional
mass extinctions at least since 542 million years
ago. Although they are disasters at the time, mass
extinctions have sometimes accelerated the
evolution of life on earth. When dominance of
particular ecological niches passes from one
group of organisms to another, it is rarely because
the new dominant group is "superior" to the old

Apparent extinction intensity, i.e. the fraction of

genera going extinct at any given time, as reconstructed
from the fossil record. (Graph not meant to include
recent epoch of Holocene extinction event)
and usually because an extinction event eliminates the old dominant group and makes way for the new one.
[182][183]
The fossil record appears to show that the gaps between mass extinctions are becoming longer and
the average and background rates of extinction are decreasing. Both of these phenomena could be
explained in one or more ways:[184]
• The oceans may have become more hospitable to life over the last 500 million years and less
vulnerable to mass extinctions: dissolved oxygen became more widespread and penetrated to
greater depths; the development of life on land reduced the run-off of nutrients and hence the risk of
eutrophication and anoxic events; and marine ecosystems became more diversified so that food
chains were less likely to be disrupted.[185][186]
• Reasonably complete fossils are very rare, most extinct organisms are represented only by
partial fossils, and complete fossils are rarest in the oldest rocks. So paleontologists have mistakenly
assigned parts of the same organism to different genera which were often defined solely to
accommodate these finds – the story of Anomalocaris is an example of this. The risk of this mistake is
higher for older fossils because these are often unlike parts of any living organism. Many of the
"superfluous" genera are represented by fragments which are not found again and the "superfluous"
genera appear to become extinct very quickly.[184]
 Biodiversity in the fossil record, which is
"the number of distinct genera
alive at any given time; that is, those
whose first occurrence predates and
whose last occurrence postdates that
time"[187]
shows a different trend: a fairly swift rise from
542 to 400 million years ago; a slight decline from
400 to 200 million years ago, in which the devastating
Permian–Triassic extinction event is an important factor;
and a swift rise from 200 million years ago to the
Thousands of genera
present.[187]

[edit] The present

Oxygenic photosynthesis accounts for virtually
all of the production of organic matter from non-organic
ingredients. Production is split about evenly between
land and marine plants, and phytoplankton are the
dominant marine producers.[188]
The processes that drive evolution are still
operating. Well-known examples include the changes in
coloration of the peppered moth over the last 200 years

Phanerozoic biodiversity as shown by the

fossil record
and the more recent appearance of pathogens that are resistant to antibiotics.[189][190] There is even
evidence that humans are still evolving, and possibly at an accelerating rate over the last 40,000 years.[191]

[edit] See also

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[edit] Footnotes
1. ^ Name given as in Butterfield's paper "Bangiomorpha pubescens ..." (2000). A fossil fish,
also from China, has also been named Qingshania. The name of one of these will have to change.
 2. ^ Myxozoa were thought to be an exception, but are now thought to be heavily modified
members of the Cnidaria: Jímenez-Guri, E., Philippe, H., Okamura, B. and Holland, P. W. H. (July
2007). "Buddenbrockia is a cnidarian worm". Science 317 (116): 116–118.
doi:10.1126/science.1142024. PMID 17615357.
http://www.sciencemag.org/cgi/content/abstract/317/5834/116. Retrieved 2008-09-03.

[edit] References
1. ^ Futuyma, Douglas J. (2005). Evolution. Sunderland, Massachusetts: Sinuer Associates,
Inc. ISBN 0-87893-187-2.
2. ^ a b c Nisbet, E.G., and Fowler, C.M.R. (December 7, 1999). "Archaean metabolic evolution
of microbial mats". Proceedings of the Royal Society: Biology 266 (1436): 2375.
doi:10.1098/rspb.1999.0934. - abstract with link to free full content (PDF)
3. ^ Anbar, A.; Duan, Y.; Lyons, T.; Arnold, G.; Kendall, B.; Creaser, R.; Kaufman, A.; Gordon,
G. et al. (2007). "A whiff of oxygen before the great oxidation event?". Science (New York, N.Y.) 317
(5846): 1903–1906. doi:10.1126/science.1140325. PMID 17901330. edit
4. ^ Knoll, Andrew H.; Javaux, E.J, Hewitt, D. and Cohen, P. (2006). "Eukaryotic organisms in
Proterozoic oceans". Philosophical Transactions of the Royal Society of London, Part B 361 (1470):
1023–38. doi:10.1098/rstb.2006.1843. PMID 16754612.
5. ^ a b c d e f Fedonkin, M. A. (March 2003). "The origin of the Metazoa in the light of the
Proterozoic fossil record" (PDF). Paleontological Research 7 (1): 9–41. doi:10.2517/prpsj.7.9.
http://www.vend.paleo.ru/pub/Fedonkin_2003.pdf. Retrieved 2008-09-02.
6. ^ Bonner, J.T. (1998) The origins of multicellularity. Integr. Biol. 1, 27–36
 7. ^ "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician
Period (~450-440 m.y.a.) on the basis of fossil spores" Transition of plants to land
8. ^ Algeo, T.J.; Scheckler, S. E. (1998). "Terrestrial-marine teleconnections in the Devonian:
links between the evolution of land plants, weathering processes, and marine anoxic events".
Philosophical Transactions of the Royal Society B: Biological Sciences 353 (1365): 113–130.
doi:10.1098/rstb.1998.0195.
9. ^ "Metazoa: Fossil Record". http://www.ucmp.berkeley.edu/phyla/metazoafr.html.
10.^ Shu et al.; Luo, H-L.; Conway Morris, S.; Zhang, X-L.; Hu, S-X.; Chen, L.; Han, J.; Zhu, M.
et al. (November 4, 1999). "Lower Cambrian vertebrates from south China". Nature 402: 42–46.
doi:10.1038/46965.
11.^ Hoyt, Donald F. (1997). "Synapsid Reptiles".
http://www.csupomona.edu/~dfhoyt/classes/zoo138/SYNAPSID.HTML.
12.^ Barry, Patrick L. (January 28, 2002). "The Great Dying". Science@NASA. Science and
Technology Directorate, Marshall Space Flight Center, NASA.
http://science.nasa.gov/headlines/y2002/28jan_extinction.htm. Retrieved March 26, 2009.
13.^ Tanner LH, Lucas SG & Chapman MG (2004). "Assessing the record and causes of Late
Triassic extinctions" (PDF). Earth-Science Reviews 65 (1-2): 103–139. doi:10.1016/S0012-
8252(03)00082-5. http://nmnaturalhistory.org/pdf_files/TJB.pdf. Retrieved 2007-10-22.
14.^ Benton, M.J. (2004). Vertebrate Paleontology. Blackwell Publishers. xii-452. ISBN 0-632-
05614-2.
15.^ "Amniota - Palaeos". http://www.palaeos.org/Amniota.
16.^ Fastovsky DE, Sheehan PM (2005). "The extinction of the dinosaurs in North America".
GSA Today 15 (3): 4–10. doi:10.1130/1052-5173(2005)015<4:TEOTDI>2.0.CO;2.
http://www.gsajournals.org/perlserv/?request=get-document&doi=10.1130%2F1052-
5173%282005%29015%3C4%3ATEOTDI%3E2.0.CO%3B2. Retrieved 2007-05-18.
17.^ "Dinosaur Extinction Spurred Rise of Modern Mammals". News.nationalgeographic.com.
http://news.nationalgeographic.com/news/2007/06/070620-mammals-dinos.html. Retrieved 2009-03-
08.
18.^ Van Valkenburgh, B. (1999). "Major patterns in the history of carnivorous mammals".
Annual Review of Earth and Planetary Sciences 26: 463–493. doi:10.1146/annurev.earth.27.1.463.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.27.1.463.
19.^ a b El Albani, Abderrazak; Bengtson, Stefan; Canfield, Donald E.; Bekker, Andrey;
Macchiarelli, Reberto (July 2010). "Large colonial organisms with coordinated growth in oxygenated
environments 2.1 Gyr ago". Nature 466 (7302): 100–104. doi:10.1038/nature09166. PMID 11931142.
20.^ a b
• Dalrymple, G.B. (1991). The Age of the Earth. California: Stanford University Press.
ISBN 0-8047-1569-6.
• Newman, W.L. (July 2007). "Age of the Earth". Publications Services, USGS.
http://pubs.usgs.gov/gip/geotime/age.html. Retrieved 2008-08-29.
• Dalrymple, G.B. (2001). "The age of the Earth in the twentieth century: a problem
(mostly) solved". Geological Society, London, Special Publications 190: 205–221.
doi:10.1144/GSL.SP.2001.190.01.14.
http://sp.lyellcollection.org/cgi/content/abstract/190/1/205. Retrieved 2007-09-20.
21.^ Galimov, E.M. and Krivtsov, A.M. (December 2005). "Origin of the Earth-Moon System". J.
Earth Syst. Sci. 114 (6): 593–600. doi:10.1007/BF02715942. [1]
22.^ a b Cohen, B.A., Swindle, T.D. and Kring, D.A. (December 2000). "Support for the Lunar
Cataclysm Hypothesis from Lunar Meteorite Impact Melt Ages". Science 290 (5497): 1754–1756.
doi:10.1126/science.290.5497.1754. PMID 11099411.
http://www.sciencemag.org/cgi/content/abstract/290/5497/1754. Retrieved 2008-08-31.
23.^
• "Early Earth Likely Had Continents And Was Habitable" . University of Colorado.
2005-11-17. http://www.colorado.edu/news/releases/2005/438.html. Retrieved 2009-01-11.
• Cavosie, A.J., Valley, J.W., Wilde, S. A. and the Edinburgh Ion Microprobe Facility
(July 15, 2005). "Magmatic δ18O in 4400-3900 Ma detrital zircons: A record of the alteration
and recycling of crust in the Early Archean". Earth and Planetary Science Letters 235 (3-4):
663–681. doi:10.1016/j.epsl.2005.04.028. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6V61-4GDKB05-
3&_coverDate=07%2F15%2F2005&_alid=382434001&_rdoc=1&_fmt=&_orig=search&_qd=
1&_cdi=5801&_sort=d&view=c&_acct=C000050221&_version=1&_urlVersion=0&_userid=10
&md5=be47e49c535d059be188b66c6e596dd5.
24.^ Britt, R.R. (2002-07-24). "Evidence for Ancient Bombardment of Earth". Space.com.
http://www.space.com/scienceastronomy/planetearth/earth_bombarded_020724.html. Retrieved
2006-04-15.
25.^ Valley, J.W., Peck, W.H., King, E.M. and Wilde, S.A. (April 2002). "A cool early Earth"
(PDF). Geology 30 (4): 351–354. doi:10.1130/0091-7613(2002)030<0351:ACEE>2.0.CO;2.
http://www.geology.wisc.edu/zircon/Valley2002Cool_Early_Earth.pdf. Retrieved 2008-09-13.
26.^ Dauphas, N., Robert, F. and Marty, B. (December 2000). "The Late Asteroidal and
Cometary Bombardment of Earth as Recorded in Water Deuterium to Protium Ratio". Icarus 148 (2):
508–512. doi:10.1006/icar.2000.6489.
27.^ a b Brasier, M., McLoughlin, N., Green, O. and Wacey, D. (June 2006). "A fresh look at the
fossil evidence for early Archaean cellular life" (PDF). Philosophical Transactions of the Royal
Society: Biology 361 (1470): 887–902. doi:10.1098/rstb.2006.1835. PMID 16754605. PMC 1578727.
http://physwww.mcmaster.ca/~higgsp/3D03/BrasierArchaeanFossils.pdf. Retrieved 2008-08-30.
28.^
• Schopf, J. W. (April 1993). "Microfossils of the Early Archean Apex Chert: New
Evidence of the Antiquity of Life". Science 260 (5108): 640–646.
doi:10.1126/science.260.5108.640. PMID 11539831.
http://www.sciencemag.org/cgi/content/abstract/260/5108/640. Retrieved 2008-08-30.
• Altermann, W. and Kazmierczak, J. (2003). "Archean microfossils: a reappraisal of
early life on Earth". Res Microbiol 154 (9): 611–7. doi:10.1016/j.resmic.2003.08.006.
PMID 14596897.
29.^ Mojzsis, S.J., Arrhenius, G., McKeegan, K.D., Harrison, T.M., Nutman, A.P. and Friend,
C.R.L. (November 1996). "Evidence for life on Earth before 3,800 million years ago". Nature 384
(6604): 55–59. doi:10.1038/384055a0. PMID 8900275.
http://www.nature.com/nature/journal/v384/n6604/abs/384055a0.html. Retrieved 2008-08-30.
30.^ a b Grotzinger, J.P. and Rothman, D.H. (1996). "An abiotic model for stomatolite
morphogenesis". Nature 383: 423–425. doi:10.+1038/383423a0.
31.^
• Fedo, C.M. and Whitehouse, M.J. (May 2002). "Metasomatic Origin of Quartz-
Pyroxene Rock, Akilia, Greenland, and Implications for Earth's Earliest Life". Science 296
(5572): 1448–1452. doi:10.1126/science.1070336. PMID 12029129.
http://www.sciencemag.org/cgi/content/abstract/296/5572/1448. Retrieved 2008-08-30.
• Lepland, A., van Zuilen, M.A., Arrhenius, G., Whitehouse, M.J. and Fedo, C.M.
(January 2005). "Questioning the evidence for Earth's earliest life — Akilia revisited". Geology
 33 (1): 77–79. doi:10.1130/G20890.1.
http://geology.geoscienceworld.org/cgi/content/abstract/33/1/77. Retrieved 2008-08-30.
32.^ Schopf, J. (2006). "Fossil evidence of Archaean life". Philosophical Transactions of the
Royal Society of London: B Biological Sciences 361 (1470): 869–85. doi:10.1098/rstb.2006.1834.
PMID 16754604.
33.^ Ciccarelli, F.D., Doerks, T., von Mering, C., Creevey, C.J., et al (2006). "Toward automatic
reconstruction of a highly resolved tree of life". Science 311 (5765): 1283–7.
doi:10.1126/science.1123061. PMID 16513982.
34.^ Mason, S.F. (1984). "Origins of biomolecular handedness". Nature 311 (5981): 19–23.
doi:10.1038/311019a0. PMID 6472461.
35.^ Orgel, L.E. (October 1994). "The origin of life on the earth" (PDF). Scientific American 271
(4): 76–83. doi:10.1038/scientificamerican1094-76. PMID 7524147.
http://courses.washington.edu/biol354/The%20Origin%20of%20Life%20on%20Earth.pdf. Retrieved
2008-08-30. Also available as a web page
36.^ a b c Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. p. 6.
ISBN 0632044446.
37.^ Villarreal LP, Witzany G (October 2009). "Viruses are essential agents within the roots and
stem of the tree of life". J. Theor. Biol. 262 (4): 698–710. doi:10.1016/j.jtbi.2009.10.014.
PMID 19833132.
38.^ O'Leary, M.R. (2008). Anaxagoras and the Origin of Panspermia Theory. iUniverse, Inc..
ISBN 0595495966.
39.^ a b Arrhenius, S. (1903). "The Propagation of Life in Space". Die Umschau volume=7.
Reprinted in Goldsmith, D.,, ed. The Quest for Extraterrestrial Life. University Science Books.
ISBN 0198557043.
 40.^ Hoyle, F. and Wickramasinghe, C. (1979). "On the Nature of Interstellar Grains".
Astrophysics and Space Science 66: 77–90. doi:10.1007/BF00648361.
41.^ a b Crick, F (1973). "Directed Panspermia". Icarus 19: 341–348. doi:10.1016/0019-
1035(73)90110-3.
42.^ a b c d Warmflash, D. and Weiss, B. (November 2005). "Did Life Come From Another
World?". Scientific American: 64–71. http://www.sciam.com/article.cfm?articleID=00073A97-5745-
1359-94FF83414B7F0000&pageNumber=1&catID=2. Retrieved 2008-09-02.
43.^ Ker, Than (August 2007). "Claim of Martian Life Called 'Bogus'". space.com.
http://www.space.com/news/070823_mars_life.html. Retrieved 2008-09-02.
44.^ Bennett, J. O. (2008). "What is life?". Beyond UFOs: The Search for Extraterrestrial Life
and Its Astonishing Implications for Our Future . Princeton University Press. pp. 82–85.
ISBN 0691135495. http://books.google.com/?
id=lEQKnip7St4C&pg=PA84&dq=life+earth+carbon+water. Retrieved 2009-01-11.
45.^ Schulze-Makuch, D., Irwin, L. N. (April 2006). "The prospect of alien life in exotic forms on
other worlds". Naturwissenschaften 93 (4): 155–72. doi:10.1007/s00114-005-0078-6.
PMID 16525788.
46.^ Peretó, J. (2005). "Controversies on the origin of life" (PDF). Int. Microbiol. 8 (1): 23–31.
PMID 15906258. http://www.im.microbios.org/0801/0801023.pdf. Retrieved 2007-10-07.
47.^ Szathmáry, E. (February 2005). "Life: In search of the simplest cell". Nature 433 (7025):
469–470. doi:10.1038/433469a. PMID 15690023.
http://www.nature.com/nature/journal/v433/n7025/full/433469a.html. Retrieved 2008-09-01.
48.^ Luisi, P. L., Ferri, F. and Stano, P. (2006). "Approaches to semi-synthetic minimal cells: a
review". Naturwissenschaften 93 (1): 1–13. doi:10.1007/s00114-005-0056-z. PMID 16292523.
 49.^ Joyce, G.F. (2002). "The antiquity of RNA-based evolution". Nature 418 (6894): 214–21.
doi:10.1038/418214a. PMID 12110897.
50.^ a b Hoenigsberg, H. (December 2003)). "Evolution without speciation but with selection:
LUCA, the Last Universal Common Ancestor in Gilbert’s RNA world". Genetic and Molecular
Research 2 (4): 366–375. PMID 15011140. http://www.funpecrp.com.br/gmr/year2003/vol4-
2/gmr0070_full_text.htm. Retrieved 2008-08-30. (also available as PDF)
51.^ Trevors, J. T. and Abel, D. L. (2004). "Chance and necessity do not explain the origin of
life". Cell Biol. Int. 28 (11): 729–39. doi:10.1016/j.cellbi.2004.06.006. PMID 15563395.
52.^ Forterre, P., Benachenhou-Lahfa, N., Confalonieri, F., Duguet, M., Elie, C. and Labedan, B.
(1992). "The nature of the last universal ancestor and the root of the tree of life, still open questions".
BioSystems 28 (1-3): 15–32. doi:10.1016/0303-2647(92)90004-I. PMID 1337989.
53.^ Cech, T.R. (August 2000). "The ribosome is a ribozyme". Science 289 (5481): 878–9.
doi:10.1126/science.289.5481.878. PMID 10960319.
http://www.sciencemag.org/cgi/content/short/289/5481/878. Retrieved 2008-09-01.
54.^ Johnston, W. K. et al (2001). "RNA-Catalyzed RNA Polymerization: Accurate and General
RNA-Templated Primer Extension". Science 292 (5520): 1319–1325. doi:10.1126/science.1060786.
PMID 11358999.
55.^ a b
• Levy, M. and Miller, S.L. (July 1998). "The stability of the RNA bases: implications for
the origin of life". Proc. Natl. Acad. Sci. U.S.A. 95 (14): 7933–8. doi:10.1073/pnas.95.14.7933.
PMID 9653118. PMC 20907. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=9653118.
• Larralde, R., Robertson, M. P. and Miller, S. L. (August 1995). "Rates of
decomposition of ribose and other sugars: implications for chemical evolution". Proc. Natl.
 Acad. Sci. U.S.A. 92 (18): 8158–60. doi:10.1073/pnas.92.18.8158. PMID 7667262.
PMC 41115. http://www.pnas.org/cgi/pmidlookup?view=long&pmid=7667262.
• Lindahl, T. (April 1993). "Instability and decay of the primary structure of DNA".
Nature 362 (6422): 709–15. doi:10.1038/362709a0. PMID 8469282.
56.^ Orgel, L. (November 2000). "Origin of life. A simpler nucleic acid". Science (journal) 290
(5495): 1306–7. PMID 11185405.
57.^ Nelson, K.E., Levy, M., and Miller, S.L. (April 2000). "Peptide nucleic acids rather than RNA
may have been the first genetic molecule". Proc. Natl. Acad. Sci. U.S.A. 97 (8): 3868–71.
doi:10.1073/pnas.97.8.3868. PMID 10760258. PMC 18108. http://www.pnas.org/cgi/pmidlookup?
view=long&pmid=10760258.
58.^ Martin, W. and Russell, M.J. (2003). "On the origins of cells: a hypothesis for the
evolutionary transitions from abiotic geochemistry to chemoautotrophic prokaryotes, and from
prokaryotes to nucleated cells". Philosophical Transactions of the Royal Society: Biological 358
(1429): 59–85. doi:10.1098/rstb.2002.1183. PMID 12594918.
59.^ Wächtershäuser, G. (August 2000). "Origin of life. Life as we don't know it". Science
(journal) 289 (5483): 1307–8. PMID 10979855.
60.^ Trevors, J.T. and Psenner, R. (2001). "From self-assembly of life to present-day bacteria: a
possible role for nanocells". FEMS Microbiol. Rev. 25 (5): 573–82. doi:10.1111/j.1574-
6976.2001.tb00592.x. PMID 11742692.
61.^ Segré, D., Ben-Eli, D., Deamer, D. and Lancet, D. (February–April 2001). "The Lipid World"
(PDF). Origins of Life and Evolution of Biospheres 2001 31 (1-2): 119–45.
doi:10.1023/A:1006746807104. PMID 11296516. http://ool.weizmann.ac.il/Segre_Lipid_World.pdf.
Retrieved 2008-09-01.
 62.^ Cairns-Smith, A.G. (1968). "An approach to a blueprint for a primitive organism". in
Waddington, C,H.. Towards a Theoretical Biology. 1. Edinburgh University Press. pp. 57–66
63.^ Ferris, J.P. (June 1999). "Prebiotic Synthesis on Minerals: Bridging the Prebiotic and RNA
Worlds". Biological Bulletin. Evolution: A Molecular Point of View (Biological Bulletin, Vol. 196, No. 3)
196 (3): 311–314. doi:10.2307/1542957. PMID 10390828. http://www.jstor.org/pss/1542957.
Retrieved 2008-09-01.
64.^ Hanczyc, M.M., Fujikawa, S.M. and Szostak, Jack W. (October 2003). "Experimental
Models of Primitive Cellular Compartments: Encapsulation, Growth, and Division". Science 302
(5645): 618–622. doi:10.1126/science.1089904. PMID 14576428.
http://www.sciencemag.org/cgi/content/abstract/302/5645/618. Retrieved 2008-09-01.
65.^ Hartman, H. (October 1998). "Photosynthesis and the Origin of Life". Origins of Life and
Evolution of Biospheres 28 (4–6): 512–521. http://www.springerlink.com/content/t1n325268n01217k/.
Retrieved 2008-09-01.
66.^ a b Krumbein, W.E., Brehm, U., Gerdes, G., Gorbushina, A.A., Levit, G. and Palinska, K.A.
(2003). "Biofilm, Biodictyon, Biomat Microbialites, Oolites, Stromatolites, Geophysiology, Global
Mechanism, Parahistology". in Krumbein, W.E., Paterson, D.M., and Zavarzin, G.A. (PDF). Fossil
and Recent Biofilms: A Natural History of Life on Earth . Kluwer Academic. pp. 1–28.
ISBN 1402015976. Archived from the original on January 6, 2007.
http://web.archive.org/web/20070106201614/http://134.106.242.33/krumbein/htdocs/Archive/397/Kru
mbein_397.pdf. Retrieved 2008-07-09
67.^ a b Risatti, J. B., Capman, W. C. and Stahl, D. A. (October 11, 1994). "Community structure
of a microbial mat: the phylogenetic dimension" (PDF). Proceedings of the National Academy of
Sciences 91 (21): 10173–10177. doi:10.1073/pnas.91.21.10173. PMID 7937858. PMC 44980.
http://www.pnas.org/content/91/21/10173.full.pdf. Retrieved 2008-07-09.
 68.^ (the editor) (June 2006)). "Editor's Summary: Biodiversity rocks". Nature 441.
http://www.nature.com/nature/journal/v441/n7094/edsumm/e060608-01.html. Retrieved 2009-01-10.
69.^ Allwood, A. C., Walter, M. R., Kamber, B. S., Marshall, C. P. and Burch, I. W. (June 2006)).
"Stromatolite reef from the Early Archaean era of Australia". Nature 441 (7094): 714–718.
doi:10.1038/nature04764. PMID 16760969.
http://www.nature.com/nature/journal/v441/n7094/abs/nature04764.html. Retrieved 2008-08-31.
70.^ Blankenship, R.E.; Bebout, BM; Des Marais, DJ (1 January 2001). "Molecular evidence for
the evolution of photosynthesis". Trends in Plant Science 6 (1): 4–6. doi:10.1038/35085554.
PMID 11460161. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6TD1-424KK4J-
3&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_url
Version=0&_userid=10&md5=6f38f9f1d29b24fc90d0145837338b9e. Retrieved 2008-07-14.
71.^ Hoehler, T.M., Bebout, B.M. and Des Marais, D.J. (19 July 2001). "The role of microbial
mats in the production of reduced gases on the early Earth". Nature 412 (6844): 324–327.
doi:10.1038/35085554. PMID 11460161.
http://www.nature.com/nature/journal/v412/n6844/full/412324a0.html. Retrieved 2008-07-14.
72.^ Abele, D. (7 November 2002). "Toxic oxygen: The radical life-giver". Nature 420 (27): 27.
doi:10.1038/420027a. PMID 12422197.
http://www.nature.com/nature/journal/v420/n6911/full/420027a.html. Retrieved 2008-07-14.
73.^ "Introduction to Aerobic Respiration". University of California, Davis. Archived from the
original on October 29, 2007.
http://web.archive.org/web/20071029120120/http://trc.ucdavis.edu/biosci10v/bis10v/week3/06aerobi
crespirintro.html. Retrieved 2008-07-14.
74.^ Goldblatt, C., Lenton, T.M. and Watson, A.J. (2006). "The Great Oxidation at ~2.4 Ga as a
bistability in atmospheric oxygen due to UV shielding by ozone" (PDF). Geophysical Research
Abstracts 8 (00770). http://www.cosis.net/abstracts/EGU06/00770/EGU06-J-00770.pdf. Retrieved
2008-09-01.
75.^ a b Glansdorff, N., Xu, Y. and Labedan, B. (2008). "The Last Universal Common Ancestor:
emergence, constitution and genetic legacy of an elusive forerunner". Biology Direct 3 (29): 29.
doi:10.1186/1745-6150-3-29. PMID 18613974.
76.^ a b Brocks, J. J., Logan, G. A., Buick, R. and Summons, R. E. (1999). "Archaean molecular
fossils and the rise of eukaryotes". Science 285 (5430): 1033–1036.
doi:10.1126/science.285.5430.1033. PMID 10446042.
http://www.sciencemag.org/cgi/content/abstract/285/5430/1033. Retrieved 2008-09-02.
77.^ a b Hedges, S. B., Blair, J. E., Venturi, M. L. and Shoe, J. L (January 2004). "A molecular
timescale of eukaryote evolution and the rise of complex multicellular life". BMC Evolutionary Biology
4 (2): 2. doi:10.1186/1471-2148-4-2. PMID 15005799. PMC 341452.
http://www.biomedcentral.com/1471-2148/4/2/abstract/. Retrieved 2008-07-14.
78.^ Burki, F., Shalchian-Tabrizi, K., Minge, M., Skjæveland, Å., Nikolaev, S. I. et al. (2007).
"Phylogenomics Reshuffles the Eukaryotic Supergroups". PLoS ONE 2 (8): e790.
doi:10.1371/journal.pone.0000790. PMID 17726520.
79.^ Parfrey, L. W., Barbero, E., Lasser, E., Dunthorn, M., Bhattacharya, D., Patterson, D.J. and
Katz, L.A. (December 2006). "Evaluating Support for the Current Classification of Eukaryotic
Diversity". PLoS Genetics 2 (12): e220. doi:10.1371/journal.pgen.0020220. PMID 17194223.
80.^ Margulis, L. (1981). Symbiosis in cell evolution. San Francisco: W.H. Freeman.
ISBN 0716712563.
81.^ Vellai, T. and Vida, G. (1999). "The origin of eukaryotes; the difference between eukaryotic
and prokaryotic cells". Proceedings of the Royal Society: Biology 266 (1428): 1571–1577.
doi:10.1098/rspb.1999.0817. PMID 10467746.
 82.^ Selosse, M-A., Abert, B., and Godelle, B. (2001). "Reducing the genome size of organelles
favours gene transfer to the nucleus". Trends in ecology & evolution 16 (3): 135–141.
doi:10.1016/S0169-5347(00)02084-X. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6VJ1-429XTFM-
H&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=8370ca16bcde45bfa1c050068a2d6e19. Retrieved 2008-09-02.
83.^ Pisani, D., Cotton, J.A. and McInerney, J.O. (2007). "Supertrees disentangle the chimerical
origin of eukaryotic genomes". Mol Biol Evol. 24 (8): 1752–60. doi:10.1093/molbev/msm095.
PMID 17504772.
84.^ Gray, M.W., Burger, G., and Lang, B.F. (1999). "Mitochondrial evolution". Science 283
(5407): 1476–1481. doi:10.1126/science.283.5407.1476. PMID 10066161.
http://www.sciencemag.org/cgi/content/abstract/283/5407/1476. Retrieved 2008-09-02.
85.^ Rasmussen, B., Fletcher, I.R., Brocks, J.R. and Kilburn, M.R. (October 2008).
"Reassessing the first appearance of eukaryotes and cyanobacteria". Nature 455 (7216): 1101–1104.
doi:10.1038/nature07381. PMID 18948954.
86.^ Han, T.M. and Runnegar, B. (July 1992). "Megascopic eukaryotic algae from the 2.1-billion-
year-old negaunee iron-formation, Michigan". Science 257 (5067): 232–235.
doi:10.1126/science.1631544. PMID 1631544.
http://www.sciencemag.org/cgi/content/abstract/257/5067/232. Retrieved 2008-09-02.
87.^ Javaux, E. J., Knoll, A. H. and Walter, M. R. (September 2004). "TEM evidence for
eukaryotic diversity in mid-Proterozoic oceans". Geobiology 2 (3): 121–132. doi:10.1111/j.1472-
4677.2004.00027.x. http://www3.interscience.wiley.com/journal/118814335/abstract. Retrieved
2008-09-02.
 88.^ a b Butterfield, N. J. (2005). "Probable Proterozoic fungi". Paleobiology 31 (1): 165–182.
doi:10.1666/0094-8373(2005)031<0165:PPF>2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/content/abstract/31/1/165. Retrieved 2008-09-02.
89.^ a b Nakagaki, T., Yamada, H. and Tóth, Á. (September 2000). "Intelligence: Maze-solving
by an amoeboid organism". Nature 407 (6803): 470. doi:10.1038/35035159. PMID 11028990.
http://www.nature.com/nature/journal/v407/n6803/abs/407470a0.html. Retrieved 2008-09-03.
90.^ Bell, G. and Mooers, A.O. (1968). "Size and complexity among multicellular organisms".
Biological Journal of the Linnean Society 60 (3): 345–363. doi:10.1111/j.1095-8312.1997.tb01500.x.
http://www3.interscience.wiley.com/journal/119168103/abstract. Retrieved 2008-09-03.
91.^ Kaiser, D. (2001). "Building a multicellular organism". Annual Review of Genetics 35: 103–
123. doi:10.1146/annurev.genet.35.102401.090145. PMID 11700279.
92.^ a b Bonner, J. T. (January 1999). "The Origins of Multicellularity". Integrative Biology 1 (1):
27–36. doi:10.1002/(SICI)1520-6602(1998)1:1<27::AID-INBI4>3.0.CO;2-6.
http://doi.wiley.com/10.1002/(SICI)1520-6602(1998)1:1%3C27::AID-INBI4%3E3.0.CO;2-6. Retrieved
2008-09-03.
93.^ a b c d e Butterfield, N. J. (September 2000). "Bangiomorpha pubescens n. gen., n. sp.:
implications for the evolution of sex, multicellularity, and the Mesoproterozoic/Neoproterozoic
radiation of eukaryotes". Paleobiology 26 (3): 386–404. doi:10.1666/0094-
8373(2000)026<0386:BPNGNS>2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/content/abstract/26/3/386. Retrieved 2008-09-02.
94.^ a b c d e Jokela, J. (2001). "Sex: Advantage". Encyclopedia of Life Sciences. John Wiley &
Sons, Ltd.. doi:10.1038/npg.els.0001716
 95.^ a b Holmes, R.K. and Jobling, M.G. (1996). "Genetics: Exchange of Genetic Information". in
Baron, S.. Baron's Medical Microbiology (4th ed.). Galveston: University of Texas Medical Branch.
ISBN 0-9631172-1-1. http://www.ncbi.nlm.nih.gov/books/bv.fcgi?
highlight=conjugation&rid=mmed.section.468. Retrieved 2008-09-02
96.^ a b Christie, P. J. (April 2001). "Type IV secretion: intercellular transfer of macromolecules
by systems ancestrally related to conjugation machines". Molecular Microbiology 40 (22): 294–305.
doi:10.1046/j.1365-2958.2001.02302.x. http://lib.bioinfo.pl/meid:10183. Retrieved 2008-09-02.
97.^ Ramesh, M. A., Malik, S-B. and Logsdon, J. M. Jr. (January 2005). "A phylogenomic
inventory of meiotic genes; evidence for sex in Giardia and an early eukaryotic origin of meiosis"
(PDF). Current Biology 15 (2): 185–91. doi:10.1016/j.cub.2005.01.003. PMID 15668177.
http://euplotes.biology.uiowa.edu/web/jmlpubls/rml05.pdf. Retrieved 2008-12-22.
98.^ a b Otto, S. P., and Gerstein, A. C. (2006). "Why have sex? The population genetics of sex
and recombination". Biochemical Society Transactions 34 (Pt 4): 519–522. doi:10.1042/BST0340519.
PMID 16856849. http://www.biochemsoctrans.org/bst/034/0519/bst0340519.htm. Retrieved 2008-12-
22.
99.^ Bernstein, H., Byerly, H., Hopf, F. and Michod, R. (1984). "Origin of sex". Journal of
Theoretical Biology 110 (3): 323–351. doi:10.1016/S0022-5193(84)80178-2. PMID 6209512.
100.^ a b c Judson, O. (2002). Dr. Tatiana's sex advice to all creation. New York: Metropolitan
Books. pp. 233–4. ISBN 0-8050-6331-5.
101.^ Hickey, D. (1982). "Selfish DNA: a sexually-transmitted nuclear parasite". Genetics 101
(3–4): 519–31. PMID 6293914.
102.^ DasSarma, S. (2007). Extreme Microbes. "Extreme Microbes". American Scientist 95:
224–231.
 103.^ Sterrer, W. (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology
216 (4): 387–96. doi:10.1006/jtbi.2002.3008. PMID 12151256. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6WMD-46DM0JD-
1&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=3f5b76ba0b7fff999ee670479f06e5af. Retrieved 2008-08-02.
104.^ a b Dong, L., Xiao, S., Shen, B. and Zhou, C. (January 2008). "Silicified Horodyskia and
Palaeopascichnus from upper Ediacaran cherts in South China: tentative phylogenetic interpretation
and implications for evolutionary stasis". Journal of the Geological Society 165: 367–378.
doi:10.1144/0016-76492007-074.
http://findarticles.com/p/articles/mi_qa3721/is_200801/ai_n24394476/pg_1?tag=artBody;col1.
Retrieved 2008-09-02.
105.^ Dickey, Gwyneth. "African fossils suggest complex life arose early", Science News,
Washington, D.C., Wednesday, June 30th, 2010. Retrieved on 2010-07-02.
106.^ Gaidos, E., Dubuc, T., Dunford, M., McAndrew, P., Padilla-gamiño, J., Studer, B.,
Weersing, K. and Stanley, S. (2007). "The Precambrian emergence of animal life: a geobiological
perspective" (PDF). Geobiology 5: 351. doi:10.1111/j.1472-4669.2007.00125.x.
http://www.soest.hawaii.edu/GG/FACULTY/GAIDOS/geobiology2007.pdf. Retrieved 2008-09-03.
107.^ Davidson, M.W.. "Animal Cell Structure". Florida State University.
http://micro.magnet.fsu.edu/cells/animalcell.html. Retrieved 2008-09-03.
108.^ Saupe, S.G. "Concepts of Biology". College of St. Benedict / St. John's University.
http://employees.csbsju.edu/SSAUPE/biol116/Zoology/digestion.htm. Retrieved 2008-09-03.
109.^ Hinde, R. T. (1998). "The Cnidaria and Ctenophora". in Anderson, D.T.,. Invertebrate
Zoology. Oxford University Press. pp. 28–57. ISBN 0195513681.
 110.^ Chen, J.-Y., Oliveri, P., Gao, F., Dornbos, S.Q., Li, C-W., Bottjer, D.J. and Davidson, E.H.
(August 2002). "Precambrian Animal Life: Probable Developmental and Adult Cnidarian Forms from
Southwest China" (PDF). Developmental Biology 248 (1): 182–196. doi:10.1006/dbio.2002.0714.
PMID 12142030. http://www.uwm.edu/~sdornbos/PDF's/Chen%20et%20al.%202002.pdf. Retrieved
2008-09-03.
111.^ Grazhdankin, D. (2004). "Patterns of distribution in the Ediacaran biotas: facies versus
biogeography and evolution". Paleobiology 30: 203. doi:10.1666/0094-
8373(2004)030<0203:PODITE>2.0.CO;2. ISSN 0094–8373.
112.^ Seilacher, A. (1992). "Vendobionta and Psammocorallia: lost constructions of
Precambrian evolution" (abstract). Journal of the Geological Society, London 149 (4): 607–613.
doi:10.1144/gsjgs.149.4.0607. ISSN 0016–7649.
http://jgs.lyellcollection.org/cgi/content/abstract/149/4/607. Retrieved 2007-06-21.
113.^ Martin, M.W.; Grazhdankin, D. V., Bowring, S. A., Evans, D. A. D., Fedonkin, M. A. and
Kirschvink, J. L. (2000-05-05). "Age of Neoproterozoic Bilaterian Body and Trace Fossils, White Sea,
Russia: Implications for Metazoan Evolution" (abstract). Science 288 (5467): 841.
doi:10.1126/science.288.5467.841. PMID 10797002.
http://www.scienceonline.org/cgi/content/abstract/288/5467/841. Retrieved 2008-07-03.
114.^ Fedonkin, M. A. and Waggoner, B. (1997). "The late Precambrian fossil Kimberella is a
mollusc-like bilaterian organism" (abstract). Nature 388: 868–871. doi:10.1038/42242.
http://www.nature.com/nature/journal/v388/n6645/abs/388868a0.html. Retrieved 2008-07-03.
115.^ Mooi, R. and Bruno, D. (1999). "Evolution within a bizarre phylum: Homologies of the first
echinoderms" (PDF). American Zoologist 38: 965–974.
http://icb.oxfordjournals.org/cgi/reprint/38/6/965.pdf. Retrieved 2007-11-24.
 116.^ McMenamin, M. A. S (2003). "Spriggina is a trilobitoid ecdysozoan" (abstract). Abstracts
with Programs (Geological Society of America) 35 (6): 105.
http://gsa.confex.com/gsa/2003AM/finalprogram/abstract_62056.htm. Retrieved 2007-11-24.
117.^ Lin, J. P.; Gon, S.M.; Gehling, J.G.; Babcock, L.E.; Zhao, Y.L.; Zhang, X.L.; Hu, S.X.;
Yuan, J.L.; Yu, M.Y.; Peng, J. (2006). "A Parvancorina-like arthropod from the Cambrian of South
China". Historical Biology 18 (1): 33–45. doi:10.1080/08912960500508689. edit
118.^ Butterfield, N. J. (2006). "Hooking some stem-group "worms": fossil lophotrochozoans in
the Burgess Shale". Bioessays 28 (12): 1161–6. doi:10.1002/bies.20507. PMID 17120226.
119.^ a b c Bengtson, S. (2004). "Early skeletal fossils". in Lipps, J.H., and Waggoner, B.M.
(PDF). Neoproterozoic - Cambrian Biological Revolutions. 10. 67–78.
http://www.cosmonova.org/download/18.4e32c81078a8d9249800021554/Bengtson2004ESF.pdf.
Retrieved 2008-07-18
120.^ Gould, S. J. (1989). Wonderful Life. Hutchinson Radius. pp. 124–136 and many others.
ISBN 0091742714.
121.^ Gould, S. J. (1989). Wonderful Life: The Burgess Shale and the Nature of History . W.W.
Norton & Company. ISBN 039330700X.
122.^ Budd, G. E. (2003). "The Cambrian Fossil Record and the Origin of the Phyla" (Free full
text). Integrative and Comparative Biology 43 (1): 157–165. doi:10.1093/icb/43.1.157. http://intl-
icb.oxfordjournals.org/cgi/content/abstract/43/1/157. Retrieved 2008-07-15.
123.^ Budd, G. E. (1996). "The morphology of Opabinia regalis and the reconstruction of the
arthropod stem-group". Lethaia 29 (1): 1–14. doi:10.1111/j.1502-3931.1996.tb01831.x.
124.^ Marshall, C. R. (2006). "Explaining the Cambrian “Explosion” of Animals". Annu. Rev.
Earth Planet. Sci. 34: 355–384. doi:10.1146/annurev.earth.33.031504.103001.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.33.031504.103001?
journalCode=earth. Retrieved 2007-11-06.
125.^ Janvier, P. (2001). "Vertebrata (Vertebrates)". Encyclopedia of Life Sciences. Wiley
InterScience. doi:10.1038/npg.els.0001531
126.^ Conway Morris, S. (August 2, 2003). "Once we were worms". New Scientist 179 (2406):
34. http://cas.bellarmine.edu/tietjen/Evolution/once_we_were_worms.htm. Retrieved 2008-09-05.
127.^ Shu, D-G., Luo, H-L., Conway Morris, S., Zhang, X-L., Hu, S-X., Chen, L., J. Han, J., Zhu,
M., Li, Y. and Chen, L-Z. (November 1999). "Lower Cambrian vertebrates from south China" (PDF).
Nature 402: 42–46. doi:10.1038/46965. http://www.bios.niu.edu/davis/bios458/Shu1.pdf. Retrieved
2008-09-05.
128.^ Shu, D.-G., Conway Morris, S., Han, J., Zhang, Z.-F., Yasui, K., Janvier, P., Chen, L.,
Zhang, X.-L., Liu, J.-N., Li, Y. and Liu, H.-Q. (January 2003). "Head and backbone of the Early
Cambrian vertebrate Haikouichthys". Nature 421 (6922): 526–529. doi:10.1038/nature01264.
PMID 12556891. http://www.nature.com/nature/journal/v421/n6922/abs/nature01264.html. Retrieved
2008-09-05.
129.^ Sansom I. J., Smith, M. M. and Smith, M. P. (2001). "The Ordovician radiation of
vertebrates". in Ahlberg, P.E.. Major Events in Early Vertebrate Evolution. Taylor and Francis.
pp. 156–171. ISBN 0-415-23370-4
130.^ a b Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. pp. 120–122.
ISBN 0632044446.
131.^ a b c Selden, P. A. (2001). ""Terrestrialization of Animals"". in Briggs, D.E.G., and
Crowther, P.R.. Palaeobiology II: A Synthesis. Blackwell. pp. 71–74. ISBN 0632051493.
http://books.google.com/?id=AHsrhGOTRM4C&pg=PA71&lpg=PA71&dq=
%22Terrestrialization+of+Animals%22+selden. Retrieved 2008-09-05
 132.^ a b c d e f g Shear, W.A. (2000). "The Early Development of Terrestrial Ecosystems". in
Gee, H.. Shaking the Tree: Readings from Nature in the History of Life . University of Chicago Press.
pp. 169–184. ISBN 0226284964. http://books.google.com/?id=ZJe_Dmdbm-
QC&pg=PA233&dq=evolution+flowering+plant+angiosperm. Retrieved 2008-09-09
133.^ a b Hawksworth, D.L. (2001). "Lichens". Encyclopedia of Life Sciences. John Wiley &
Sons, Ltd.. doi:10.1038/npg.els.0000368
134.^ Retallack, G.J.; Feakes, C.R. (1987). "Trace Fossil Evidence for Late Ordovician Animals
on Land". Science 235 (4784): 61–63. doi:10.1126/science.235.4784.61. PMID 17769314.
135.^ a b Kenrick, P. and Crane, P. R. (September 1997). "The origin and early evolution of
plants on land" (PDF). Nature 389: 33. doi:10.1038/37918.
http://botit.botany.wisc.edu/courses/botany_940/06EvidEvol/papers/KendrickCrane1997.pdf.
Retrieved 2008-09-05.
136.^ Scheckler, S. E. (2001). ""Afforestation – the First Forests"". in Briggs, D.E.G., and
Crowther, P.R.. Palaeobiology II: A Synthesis. Blackwell. pp. 67–70. ISBN 0632051493.
http://books.google.com/?
id=AHsrhGOTRM4C&pg=PA69&lpg=PA69&dq=devonian+meandering+plants+trees. Retrieved
2008-09-05
137.^ The phrase "Late Devonian wood crisis" is used at "Palaeos – Tetrapoda: Acanthostega".
PALAEOS: The Trace of Life on Earth.
http://www.palaeos.com/Vertebrates/Units/150Tetrapoda/150.150.html. Retrieved 2008-09-05.
138.^ a b Algeo, T. J. and Scheckler, S. E. (1998). "Terrestrial-marine teleconnections in the
Devonian: links between the evolution of land plants, weathering processes, and marine anoxic
events". Philosophical Transactions of the Royal Society: Biology 353: 113–130.
doi:10.1098/rstb.1998.0195.
 139.^ Taylor T. N. and Osborn J. M. (1996). "The importance of fungi in shaping the
paleoecosystem". Review of Paleobotany and Palynology 90: 249–262. doi:10.1016/0034-
6667(95)00086-0. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B6V6W-454YDFK-
7&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_acct=C000050221&_version=1&_url
Version=0&_userid=10&md5=9d5d008d99d044684e947ad74b05514d. Retrieved 2008-09-05.
140.^ MacNaughton, R. B., Cole, J. M., Dalrymple, R. W., Braddy, S. J., Briggs, D. E. G. and
Lukie, T. D. (May 2002). "First steps on land: Arthropod trackways in Cambrian-Ordovician eolian
sandstone, southeastern Ontario, Canada". Geology 30 (5): 391–394. doi:10.1130/0091-
7613(2002)030<0391:FSOLAT>2.0.CO;2.
http://geology.geoscienceworld.org/cgi/content/abstract/30/5/391. Retrieved 2008-09-05.
141.^ Vaccari, N. E., Edgecombe, G. D. and Escudero, C. (2004). "Cambrian origins and
affinities of an enigmatic fossil group of arthropods". Nature 430 (6999): 554–557.
doi:10.1038/nature02705. PMID 15282604.
142.^ Buatois, L. A., Mangano, M. G., Genise, J. F. and Taylor, T. N. (June 1998). "The
ichnologic record of the continental invertebrate invasion; evolutionary trends in environmental
expansion, ecospace utilization, and behavioral complexity". PALAIOS (PALAIOS, Vol. 13, No. 3) 13
(3): 217–240. doi:10.2307/3515447. http://palaios.sepmonline.org/cgi/content/abstract/13/3/217.
Retrieved 2008-09-05.
143.^ Cowen, R. (2000). History of Life (3rd ed.). Blackwell Science. p. 126. ISBN 0632044446.
144.^ Grimaldi, D. and Engel, M. (2005). "Insects Take to the Skies". Evolution of the Insects.
Cambridge University Press. pp. 155–160. ISBN 0521821495. http://books.google.com/?
id=Ql6Jl6wKb88C&dq=%22Evolution+of+the+Insects%22+grimaldi&printsec=frontcover. Retrieved
2009-01-11.
 145.^ Grimaldi, D. and Engel, M. (2005). "Diversity of evolution". Evolution of the Insects.
Cambridge University Press. p. 12. ISBN 0521821495. http://books.google.com/?
id=Ql6Jl6wKb88C&dq=%22Evolution+of+the+Insects%22+grimaldi&printsec=frontcover. Retrieved
2009-01-11.
146.^ a b c Clack, J. A. (November, 2005). "Getting a Leg Up on Land". Scientific American.
http://www.sciam.com/article.cfm?id=getting-a-leg-up-on-land. Retrieved 2008-09-06.
147.^ a b Ahlberg, P. E. and Milner, A. R. (April 1994). "The Origin and Early Diversification of
Tetrapods". Nature 368: 507–514. doi:10.1038/368507a0.
http://www.nature.com/nature/journal/v368/n6471/abs/368507a0.html. Retrieved 2008-09-06.
148.^ Gordon, M. S., Graham, J. B. and Wang, T. (September/October 2004). "Revisiting the
Vertebrate Invasion of the Land". Physiological and Biochemical Zoology 77 (5): 697–699.
doi:10.1086/425182.
149.^ Daeschler, E. B., Shubin, N. H. and Jenkins, F. A. (April 2006). "A Devonian tetrapod-like
fish and the evolution of the tetrapod body plan" (PDF). Nature 440 (7085): 757–763.
doi:10.1038/nature04639. PMID 16598249. http://www.com.univ-
mrs.fr/~boudouresque/Publications_DOM_2006_2007/Daeschler_et_al_2006_Nature.pdf. Retrieved
2008-09-06.
150.^ Debraga, M. and Rieppel, O. (July 1997). "Reptile phylogeny and the interrelationships of
turtles". Zoological Journal of the Linnean Society 120 (3): 281–354. doi:10.1111/j.1096-
3642.1997.tb01280.x. http://www3.interscience.wiley.com/journal/119830935/abstract. Retrieved
2008-09-07.
151.^ a b Benton M. J. and Donoghue, P. C. J. (2007). "Paleontological Evidence to Date the
Tree of Life". Molecular Biology and Evolution 24 (1): 26–53. doi:10.1093/molbev/msl150.
PMID 17047029. http://mbe.oxfordjournals.org/cgi/content/full/24/1/26. Retrieved 2008-09-07.
 152.^ a b Benton, M. J. (May 1990). "Phylogeny of the Major Tetrapod Groups: Morphological
Data and Divergence Dates". Journal of Molecular Evolution 30 (5): 409–424.
doi:10.1007/BF02101113. PMID 2111854. http://www.springerlink.com/content/k152294003652458/.
Retrieved 2008-09-07.
153.^ Sidor, C. A., O'Keefe, F. R., Damiani, R., Steyer, J. S., Smith, R. M. H., Larsson, H. C. E.,
Sereno, P. C., Ide, O., and Maga, A. (April 2005). "Permian tetrapods from the Sahara show climate-
controlled endemism in Pangaea". Nature 434 (7035): 886–889. doi:10.1038/nature03393.
PMID 15829962. http://www.nature.com/nature/journal/v434/n7035/full/nature03393.html. Retrieved
2008-09-08.
154.^ Smith, R. and Botha, J. (September–October 2005). "The recovery of terrestrial vertebrate
diversity in the South African Karoo Basin after the end-Permian extinction". Comptes Rendus
Palevol 4 (6-7): 623–636. doi:10.1016/j.crpv.2005.07.005. http://www.sciencedirect.com/science?
_ob=ArticleURL&_udi=B6X1G-4GYH7VN-
1&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=add24b0622f2aff0b41e7c42a3160fa7. Retrieved 2008-09-08.
155.^ Benton, M. J. (2005). When Life Nearly Died: The Greatest Mass Extinction of All Time .
Thames & Hudson. ISBN 978-0500285732.
156.^ Sahney, S. and Benton, M.J. (2008). "Recovery from the most profound mass extinction of
all time" (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759.
doi:10.1098/rspb.2007.1370. PMID 18198148. PMC 2596898.
http://journals.royalsociety.org/content/qq5un1810k7605h5/fulltext.pdf.
157.^ Gauthier, J., Cannatella, D. C., de Queiroz, K., Kluge, A. G. and Rowe, T. (1989).
"Tetrapod Phylogeny". in B. Fernholm, B., Bremer K., and Jörnvall, H. (PDF). The Hierarchy of Life.
Elsevier Science. p. 345. http://si-
pddr.si.edu/dspace/bitstream/10088/4689/1/VZ_1989GauthieretalHierLife.pdf. Retrieved 2008-09-08
158.^ Benton, M. J. (March 1983). "Dinosaur Success in the Triassic: a Noncompetitive
Ecological Model" (PDF). Quarterly Review of Biology 58 (1).
http://palaeo.gly.bris.ac.uk/Benton/reprints/1983success.pdf. Retrieved 2008-09-08
159.^ a b Padian, K. (2004). "Basal Avialae". in Weishampel, David B.; Dodson, Peter; &
Osmólska, Halszka (eds.). The Dinosauria (Second ed.). Berkeley: University of California Press.
pp. 210–231. ISBN 0-520-24209-2.
160.^ Hou, L., Zhou, Z., Martin, L. D. and Feduccia, A. (October 2002). "A beaked bird from the
Jurassic of China". Nature 377: 616–618. doi:10.1038/377616a0.
http://www.nature.com/nature/journal/v377/n6550/abs/377616a0.html. Retrieved 2008-09-08.
161.^ Clarke, J. A., Zhou, Z. and Zhang, F. (2006). "Insight into the evolution of avian flight from
a new clade of Early Cretaceous ornithurines from China and the morphology of Yixianornis grabaui".
Journal of Anatomy 208 (3): 287–308. doi:10.1111/j.1469-7580.2006.00534.x. PMID 16533313.
PMC 2100246. http://www3.interscience.wiley.com/journal/118559634/abstract?
CRETRY=1&SRETRY=0. Retrieved 2008-09-08.
162.^ Ruben, J. A. and Jones, T. D. (2000). "Selective Factors Associated with the Origin of Fur
and Feathers". American Zoologist 40 (4): 585–596. doi:10.1093/icb/40.4.585.
http://icb.oxfordjournals.org/cgi/content/full/40/4/585.
163.^ Luo, Z-X., Crompton, A. W. and Sun, A-L. (May 2001). "A New Mammaliaform from the
Early Jurassic and Evolution of Mammalian Characteristics". Science 292 (5521): 1535–1540.
doi:10.1126/science.1058476. PMID 11375489.
http://www.sciencemag.org/cgi/content/full/292/5521/1535. Retrieved 2008-09-08.
 164.^ Cifelli, R.L. (November 2001). "Early mammalian radiations". Journal of Paleontology 75:
1214. doi:10.1666/0022-3360(2001)075<1214:EMR>2.0.CO;2.
http://findarticles.com/p/articles/mi_qa3790/is_200111/ai_n8958762/pg_6.
165.^ Flynn, J. J., Parrish, J. M. Rakotosamimanana, B., Simpson, W. F. and Wyss, A.R.
(September 1999). "A Middle Jurassic mammal from Madagascar". Nature 401: 57–60.
doi:10.1038/43420. http://www.nature.com/nature/journal/v401/n6748/abs/401057a0.html. Retrieved
2008-09-08.
166.^ MacLeod, N., Rawson, P. F., Forey, P. L., Banner. F. T., Boudagher-Fadel, M. K., Bown,
P. R., Burnett, J. A., Chambers, P., Culver, S., Evans, S. E., Jeffery, C., Kaminski, M. A., Lord, A. R.,
Milner, A. C., Milner, A. R., Morris, N., Owen, E., Rosen, B. R., ,Smith, A. B., Taylor, P. D., Urquhart,
E. and Young, J. R. (1997). "The Cretaceous–Tertiary biotic transition". Journal of the Geological
Society 154 (2): 265–292. doi:10.1144/gsjgs.154.2.0265.
http://findarticles.com/p/articles/mi_qa3721/is_199703/ai_n8738406/print.
167.^ Alroy, J. (March 1999). "The fossil record of North American mammals: evidence for a
Paleocene evolutionary radiation". Systematic biology 48 (1): 107–18.
doi:10.1080/106351599260472. PMID 12078635.
168.^ Archibald, J. D. and Deutschman, D. H. (June 2001). "Quantitative Analysis of the Timing
of the Origin and Diversification of Extant Placental Orders". Journal of Mammalian Evolution 8 (2):
107–124. doi:10.1023/A:1011317930838.
http://www.ingentaconnect.com/content/klu/jomm/2001/00000008/00000002/00342277. Retrieved
2008-09-24.
169.^ Simmons, N. B., Seymour, K. L., Habersetzer, J. and Gunnell, G. F. (February 2008).
"Primitive Early Eocene bat from Wyoming and the evolution of flight and echolocation". Nature 451
(7180): 818–821. doi:10.1038/nature06549. PMID 18270539.
 170.^ Thewissen, J. G. M., Madar, S. I. and Hussain, S. T. (1996). " Ambulocetus natans, an
Eocene cetacean (Mammalia) from Pakistan". Courier Forschungsinstitut Senckenberg 191: 1–86.
ISBN 978-3-510-61084-6.
171.^ a b c d Crane, P. R., Friis, E. M. and Pedersen, K. R. (2000). "The Origin and Early
Diversification of Angiosperms". in Gee, H.. Shaking the Tree: Readings from Nature in the History of
Life. University of Chicago Press. pp. 233–250. ISBN 0226284964. http://books.google.com/?
id=ZJe_Dmdbm-QC&pg=PA233&dq=evolution+flowering+plant+angiosperm. Retrieved 2008-09-09
172.^ a b c d Crepet, W. L. (November 2000). "Progress in understanding angiosperm history,
success, and relationships: Darwin’s abominably "perplexing phenomenon"". Proceedings of the
National Academy of Sciences 97 (24): 12939–12941. doi:10.1073/pnas.97.24.12939.
PMID 11087846. PMC 34068. http://www.pnas.org/content/97/24/12939.full.pdf+html. Retrieved
2008-09-09.
173.^ a b Wilson, E. O. and Hölldobler, B. (September 2005). "Eusociality: Origin and
consequences". Proceedings of the National Academy of Sciences 102 (38): 13367–13371.
doi:10.1073/pnas.0505858102. PMID 16157878. PMC 1224642.
http://www.pnas.org/content/102/38/13367.full.pdf+html. Retrieved 2008-09-07.
174.^ Hughes, W. O. H., Oldroyd, B. P., Beekman, M. and Ratnieks, F. L. W. (2008-05-30).
"Ancestral Monogamy Shows Kin Selection Is Key to the Evolution of Eusociality". Science
(American Association for the Advancement of Science) 320 (5880): 1213–1216.
doi:10.1126/science.1156108. PMID 18511689.
http://www.sciencemag.org/cgi/content/abstract/320/5880/1213. Retrieved 2008-08-04.
175.^ Lovegrove, B. G. (January 1991). "The evolution of eusociality in molerats (Bathyergidae):
a question of risks, numbers, and costs". Behavioral Ecology and Sociobiology 28 (1): 37–45.
doi:10.1007/BF00172137. http://www.springerlink.com/content/k4n52v522l816125/. Retrieved 2008-
09-07.
176.^ Labandeira, C. and Eble, G. J. (2000). "The Fossil Record of Insect Diversity and
Disparity". in Anderson, J., Thackeray, F., van Wyk, B., and de Wit, M. (PDF). Gondwana Alive:
Biodiversity and the Evolving Biosphere. Witwatersrand University Press.
http://www.santafe.edu/research/publications/workingpapers/00-08-044.pdf. Retrieved 2008-09-07
177.^ Brunet, M., Guy, F., Pilbeam, D., Mackaye, H. T. et al (July 2002). "A new hominid from
the Upper Miocene of Chad, Central Africa". Nature 418 (6894): 145–151. doi:10.1038/nature00879.
PMID 12110880. http://www.nature.com/nature/journal/v418/n6894/abs/nature00879.html. Retrieved
2008-09-09.
178.^ de Heinzelin, J., Clark, J. D., White, T. et al (April 1999). "Environment and Behavior of
2.5-Million-Year-Old Bouri Hominids". Science 284 (5414): 625–629.
doi:10.1126/science.284.5414.625. PMID 10213682.
http://www.sciencemag.org/cgi/content/full/sci;284/5414/625. Retrieved 2008-09-09.
179.^ De Miguel, C. and Henneberg, M. (2001). "Variation in hominid brain size: How much is
due to method?". HOMO - Journal of Comparative Human Biology 52 (1): 3–58. doi:10.1078/0018-
442X-00019. http://www.sciencedirect.com/science?_ob=ArticleURL&_udi=B7GW4-4DPCHXC-
2&_user=10&_rdoc=1&_fmt=&_orig=search&_sort=d&view=c&_version=1&_urlVersion=0&_userid=
10&md5=aef79dbca1f189c885cfe9f36636b131. Retrieved 2008-09-09.
180.^ Leakey, Richard (1994). The Origin of Humankind. Science Masters Series. New York,
NY: Basic Books. pp. 87–89. ISBN 0465053130.
181.^ Mellars, Paul (2006). "Why did modern human populations disperse from Africa ca.
60,000 years ago?". Proceedings of the National Academy of Sciences 103 (25): 9381.
doi:10.1073/pnas.0510792103. PMID 16772383. PMC 1480416.
http://www.pnas.org/cgi/reprint/0510792103v1.
182.^ Benton, M. J. (2004). "6. Reptiles Of The Triassic". Vertebrate Palaeontology (3rd ed.).
Blackwell. ISBN 978-0-632-05637-8. http://www.blackwellpublishing.com/book.asp?
ref=0632056371.
183.^ Van Valkenburgh, B. (1999). "Major patterns in the history of xarnivorous mammals".
Annual Review of Earth and Planetary Sciences 26: 463–493. doi:10.1146/annurev.earth.27.1.463.
http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.earth.27.1.463.
184.^ a b MacLeod, N. (2001-01-06). "Extinction!".
http://www.firstscience.com/home/articles/earth/extinction-page-2-1_1258.html. Retrieved 2008-09-
11.
185.^ Martin, R. E. (1995). "Cyclic and secular variation in microfossil biomineralization: clues to
the biogeochemical evolution of Phanerozoic oceans". Global and Planetary Change 11 (1): 1.
doi:10.1016/0921-8181(94)00011-2.
186.^ Martin, R.E. (1996). "Secular increase in nutrient levels through the Phanerozoic:
Implications for productivity, biomass, and diversity of the marine biosphere". PALAIOS (PALAIOS,
Vol. 11, No. 3) 11 (3): 209–219. doi:10.2307/3515230. http://jstor.org/stable/3515230.
187.^ a b Rohde, R. A. and Muller, R. A. (March 2005). "Cycles in fossil diversity" (PDF). Nature
434 (7030): 208–210. doi:10.1038/nature03339. PMID 15758998. http://muller.lbl.gov/papers/Rohde-
Muller-Nature.pdf. Retrieved 2008-09-22.
188.^ Field, C. B., Behrenfeld, M. J., Randerson, J. T. and Falkowski, P. (July 1998). "Primary
Production of the Biosphere: Integrating Terrestrial and Oceanic Components". Science 281 (5374):
237–240. doi:10.1126/science.281.5374.237. PMID 9657713.
http://www.sciencemag.org/cgi/content/full/sci;281/5374/237. Retrieved 2008-09-13.
 189.^ Grant, B. S., and Wiseman, L. L. (2002). "Recent History of Melanism in American
Peppered Moths". Journal of Heredity 93 (2): 86–90. doi:10.1093/jhered/93.2.86. ISSN 1465-7333.
PMID 12140267. http://jhered.oxfordjournals.org/cgi/content/abstract/93/2/86. Retrieved 2008-09-11.
190.^ Levin, B. R., Perrot, V. and Walker, N. (March 1, 2000). "Compensatory Mutations,
Antibiotic Resistance and the Population Genetics of Adaptive Evolution in Bacteria". Genetics 154
(3): 985–997. PMID 10757748. PMC 1460977.
http://www.genetics.org/cgi/content/abstract/154/3/985. Retrieved 2008-09-11.
191.^ Hawks, J., Wang, E. T., Cochran, G. M., Harpending, H. C. and Moyzis, R. K. (December
2007). "Recent acceleration of human adaptive evolution". Proceedings of the National Academy of
Sciences 104 (52): 20753–20758. doi:10.1073/pnas.0707650104. PMID 18087044. PMC 2410101.
http://www.pnas.org/content/104/52/20753.full. Retrieved 2008-09-11.

[edit] Further reading

• Cowen, R. (2004). History of Life (4th ed.). Blackwell Publishing Limited. ISBN 978-
1405117562.
• The Ancestor's Tale, A Pilgrimage to the Dawn of Life . Boston: Houghton Mifflin Company.
2004. ISBN 0-618-00583-8.
• Richard Dawkins. (1990). The Selfish Gene. Oxford University Press. ISBN 0192860925.
• Smith, John Maynard; Eörs Szathmáry (1997). The Major Transitions in Evolution.
Oxfordshire: Oxford University Press. ISBN 0-198-50294-X.
[edit] External links
General information
• General information on evolution- Fossil Museum nav.
• Understanding Evolution from University of California, Berkeley
• National Academies Evolution Resources
• Evolution poster- PDF format "tree of life"
• Everything you wanted to know about evolution by New Scientist
• Howstuffworks.com — How Evolution Works
• Synthetic Theory Of Evolution: An Introduction to Modern Evolutionary Concepts and
Theories
History of evolutionary thought
• The Complete Work of Charles Darwin Online
• Understanding Evolution: History, Theory, Evidence, and Implications
Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_life"

Categories: History by topic | Paleontology | Evolutionary biology

 W000

Histoire évolutive du vivant

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Cet article est une ébauche concernant l'origine et évolution de la vie.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
En biologie, l'évolution est le processus par lequel des populations d'organismes vivants acquièrent
et transmettent des traits biologiques nouveaux de génération en génération. Sur une grande échelle de
temps, la répétition de ce processus explique l'origine de nouvelles espèces et la vaste diversité du monde
vivant. Les espèces biologiques contemporaines sont reliées entre elles par une descendance commune et
sont le produit de l'évolution et de la spécialisation sur plusieurs milliards d'années.
Sommaire
[masquer]
• 1 Chronologie simple de l'évolution
• 2 L'origine de la vie
• 3 L'archéen et l'évolution des premières cellules
• 4 Le protérozoïque et l'apparition de pluricellularité
• 5 L'explosion cambrienne
• 6 Le paléozoïque et la colonisation du milieu terrestre
• 7 Le mésosoïque
• 8 Le cénozoïque : histoire évolutive récente
• 9 L'évolution aujourd'hui
• 10 Dans l'enseignement,
• 11 Notes et références
• 12 Voir aussi
• 12.1 Articles connexes

Chronologie simple de l'évolution [modifier]

• - 4 600 millions d'années : formation de la planète Terre,
• - 4 000 millions d'années : apparition des cellules procaryotes,
 • - 3 000 millions d'années : apparition de la photosynthèse,
• - 2 100 millions d'années : apparition d'une vie multicellulaire,
• - 2 000 millions d'années : apparition des cellules eucaryotes,
• - 600 millions d'années : apparition d'animaux simples,
• - 570 millions d'années : apparition des arthropodes,
• - 550 millions d'années : apparition d'animaux complexes
• - 500 millions d'années : apparition des poissons et des proto-amphibiens,
• - 475 millions d'années : apparition des plantes terrestres,
• - 400 millions d'années : apparition des insectes et des graines,
• - 360 millions d'années : apparition des amphibiens,
• - 300 millions d'années : apparition des reptiles,
• - 200 millions d'années : apparition des mammifères,
• - 150 millions d'années : apparition des oiseaux,
• - 100 millions d'années : apparition des fleurs
• - 65 millions d'années : disparition des dinosaures non-aviaires (voir article Extinction du
Crétacé).

L'origine de la vie [modifier]

Article détaillé : Origine de la vie.
L'archéen et l'évolution des premières cellules [modifier]
Il y a 3,5 milliards d'années les cyanobactéries apparaissent et constituent les stromatolithes
actuelles. Ces procaryotes (unicellulaires) immobiles (sans flagelle) longtemps appelés algues bleue-vertes
sont les premiers organismes à posséder les 2 photosystèmes leur permettant de pratiquer la
photosynthèse. Elles participent de ce fait à la réduction de la concentration en CO 2 atmosphérique et à
l'augmentation de la proportion en oxygène. Selon la théorie de l'endosymbiose plastidiale, elles seraient à
l'origine des plastes dans les végétaux supérieurs intégrés par symbiose dans leurs cellules.

Le protérozoïque et l'apparition de pluricellularité [modifier]

Article détaillé : Évolution de la pluricellularité.
L'existence d'ères glaciaires généralisées il y a de 850 à 630 millions d'années environ, une ère que
l'on appelle le cryogénien, aurait favorisé le regroupement des colonies bactériennes (ou des animalcules les
plus primitifs) autour « d'oasis » de vies dans des sources hydrothermales sous-marines ou affleurant en
surface. La planète étant plongée dans un manteau de glaces, ce qui rend les comportements
« individualistes » des micro-organismes défavorables.

L'explosion cambrienne [modifier]

Article détaillé : Explosion cambrienne.
Le paléozoïque et la colonisation du milieu terrestre [modifier]
• Histoire évolutive des poissons cartilagineux
Lignées passées au milieu terrestre :
• Histoire évolutive des végétaux
• Histoire évolutive des gastéropodes

Le mésosoïque [modifier]
• Histoire évolutive des mammifères
• Histoire évolutive des dinosaures
• Histoire évolutive des oiseaux

Le cénozoïque : histoire évolutive récente [modifier]

• Histoire évolutive des équidés
• Histoire évolutive des cétacés
• Histoire évolutive des hippopotamidés
• Histoire évolutive des siréniens
• Histoire évolutive des primates
• Histoire évolutive des homininés
 L'évolution aujourd'hui [modifier]
L'évolution, bien que lente et rarement observable à l'échelle humaine, peut parfois être décelée
chez les espèces à reproduction rapide. Ainsi, les moustiques sont parfois localement devenus résistants à
des insecticides. Il en va de même pour les bactéries responsables de pathologies humaines, souvent
multirésistantes aux antibiotiques.
Enfin, on peut constater le résultat d'une évolution divergente récente dans le cas de la colonie de crabes
découverte sous Rome[1]. Ainsi qu'un changement de régime alimentaire et l'apparition de sous-espèces
chez certains lézards, comme Podarcis sicula, avec modifications anatomiques internes et externes.

Dans l'enseignement, [modifier]

L'histoire évolutive est en Europe généralement, comme au collège de France [2] à Paris abordée
sous un double angle (Biologie historique et Évolutionnisme), en tentant d'à la fois enseigner ce qu'on sait de
l’histoire de la vie sur Terre et des mécanismes sous-jacents à cette histoire, et ce que l’on peut penser des
différentes interprétations humaines relatives à cette histoire et de ces mécanismes, interprétations qui ont
varié dans l'espace et dans le temps selon les contexte et les cultures ou religions.

Notes et références [modifier]

1. ↑ Sous les vestiges de Rome, une colonie de crabes prospère. [archive]
2. ↑ Leçon inaugurale d'Armand de Ricqlès au Collège de France [archive], Chaire de Biologie
historique et évolutionnisme, Lundi 6 mai 1996
 Voir aussi [modifier]
Articles connexes [modifier]
• Évolution
• Histoire de la Terre
• Biodiversité
• Biosphère

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle
 Adaptation · Convergence évolutive · Exaptation · Extinction des
Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux
 [Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_du_vivant ».

Catégorie : Histoire évolutive | [+]

 W000

Evolution of Mollusca
From Wikipedia, the free encyclopedia

Jump to: navigation, search

This article has multiple issues. Please help improve it or discuss these issues on
the talk page.
 Diagram of primitive mollusc viewed from left side. Hypothetical mollusc ancestor. From
Encyclopædia Britannica Eleventh Edition.
The current understanding of molluscan phylogeny is shown below.
Horizontal cladogram (source:[1]):
Mollusca
Aplacophora

Solenogastres

Caudofoeveata
 Testaria
Polyplacophora

Palaeoloricata

Neoloricata

Conchifera

Monoplacophora

(unnamed)
Cyrtosoma
Gastropoda
 Cephalopoda

Diasoma

Rostroconchia

(unnamed)

Scaphopoda

Bivalvia
 Basal Mollusca phylogeny.
Early organisms which have been compared to molluscs include Kimberella and Odontogriphus.

[edit] References
1. ^ Zoology 250 Phylogenetic Trees (2006)
http://www.biology.ualberta.ca/courses.hp/zool250/Clades/clade08-Mollusca.htm
 [edit] Further reading
• From sea to land and beyond – New insights into the evolution of euthyneuran Gastropoda
(Mollusca)

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts
 Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_Mollusca"

Categories: Evolution by taxon | Molluscs | Mollusc stubs

 w000

Evolutionary history of plants

From Wikipedia, the free encyclopedia
(Redirected from Evolution of plants)
Jump to: navigation, search
Contents
[hide]
• 1 Colonisation of land
• 2 Changing life cycles
• 3 Water transport
• 4 Evolution of leaves
• 5 Evolution of trees
• 6 Evolution of roots
• 6.1 Arbuscular mycorrhizae
• 7 Evolution of seeds
• 8 Evolution of flowers
• 9 Advances in metabolism
• 9.1 Concentrating carbon
• 9.2 Evolutionary record
• 9.3 When is C4 an advantage?
• 10 Evolutionary trends
• 11 See also
• 12 References
 The evolution of plants occurred through increasing levels of complexity, from the earliest algal mats,
through bryophytes, lycopods, ferns and gymnosperms to the complex angiosperms of today. While the
simple plants continue to thrive, especially in the environments in which they evolved, each new grade of
organisation has eventually become more "successful" than its predecessors by most measures. Further,
most cladistic analyses, where they agree, suggest that each "more complex" group arose from the most
complex group at the time.
Evidence suggests that an algal scum formed on the land 1,200 million years ago, but it was not until
the Ordovician period, around 450 million years ago, that land plants appeared[1]. These began to diversify
in the late Silurian period, around 420 million years ago, and the fruits of their diversification are displayed in
remarkable detail in an early Devonian fossil assemblage known as the Rhynie chert. This chert preserved
early plants in cellular detail, petrified in volcanic springs. By the middle of the Devonian period most of the
features recognised in plants today are present, including roots, leaves and secondary wood, and by late
Devonian times seeds had evolved.[2] Late Devonian plants had thereby reached a degree of sophistication
that allowed them to form forests of tall trees. Evolutionary innovation continued after the Devonian period.
Most plant groups were relatively unscathed by the Permo-Triassic extinction event, although the structures
of communities changed. This may have set the scene for the evolution of flowering plants in the Triassic
(~200 million years ago), which exploded in the Cretaceous and Tertiary. The latest major group of plants to
evolve were the grasses, which became important in the mid Tertiary, from around 40 million years ago. The
grasses, as well as many other groups, evolved new mechanisms of metabolism to survive the low CO 2 and
warm, dry conditions of the tropics over the last 10 million years.
[edit] Colonisation of land

A late Silurian sporangium. Green: A spore tetrad. Blue: A spore bearing a trilete mark – the Y-
shaped scar. The spores are about 30-35 μm across
 The Devonian period marks the beginning of extensive land colonization by plants, which through
their effects on erosion and sedimentation brought about significant climatic change.
Land plants evolved from chlorophyte algae, perhaps as early as 510 million years ago;[3] their
closest living relatives are the charophytes, specifically Charales. Assuming that the Charales' habit has
changed little since the divergence of lineages, this means that the land plants evolved from a branched,
filamentous, haplontic alga, dwelling in shallow fresh water,[4] perhaps at the edge of seasonally desiccating
pools.[3] Co-operative interactions with fungi may have helped early plants adapt to the stresses of the
terrestrial realm.[5]
Plants were not the first photosynthesisers on land, though: consideration of weathering rates
suggests that organisms were already living on the land 1,200 million years ago,[3] and microbial fossils have
been found in freshwater lake deposits from 1,000 million years ago,[6] but the carbon isotope record
suggests that they were too scarce to impact the atmospheric composition until around 850 million years ago.
[7] These organisms were probably small and simple, forming little more than an "algal scum".[3]
The first evidence of plants on land comes from spore tetrads [8] attributed to land plants from the
Mid-Ordovician early Llanvirn, ~470 million years ago).[9] Spore tetrads consist of four identical, connected
spores, produced when a single cell undergoes meiosis. Spore tetrads are borne by all land plants, and
some algae.[8] The microstructure of the earliest spores resembles that of modern liverwort spores,
suggesting they share an equivalent grade of organisation.[10] It could be that atmospheric 'poisoning'
prevented eukaryotes from colonising the land prior to this,[11] or it could simply have taken a great time for
the necessary complexity to evolve.[12]
Trilete spores, the progeny of spore tetrads, appear soon afterwards, in the Late Ordovician.[13]
Depending exactly when the tetrad splits, each of the four spores may bear a "trilete mark", a Y-shape,
reflecting the points at which each cell was squashed up against its neighbours.[8] However, in order for this
to happen, the spore walls must be sturdy and resistant at an early stage. This resistance is closely
associated with having a desiccation-resistant outer wall – a trait only of use when spores have to survive out
of water. Indeed, even those embryophytes that have returned to the water lack a resistant wall, thus don't
bear trilete marks.[8] A close examination of algal spores shows that none have trilete spores, either because
their walls are not resistant enough, or in those rare cases where it is, the spores disperse before they are
squashed enough to develop the mark, or don't fit into a tetrahedral tetrad.[8]
The earliest megafossils of land plants were thalloid organisms, which dwelt in fluvial wetlands and
are found to have covered most of an early Silurian flood plain. They could only survive when the land was
waterlogged.[14]
 Once plants had reached the land, there were two approaches to desiccation. The bryophytes avoid
it or give in to it, restricting their ranges to moist settings, or drying out and putting their metabolism "on hold"
until more water arrives. Tracheophytes resist desiccation. They all bear a waterproof outer cuticle layer
wherever they are exposed to air (as do some bryophytes), to reduce water loss – but since a total covering
would cut them off from CO2 in the atmosphere, they rapidly evolved stomata – small openings to allow gas
exchange. Tracheophytes also developed vascular tissue to aid in the movement of water within the
organisms (see below), and moved away from a gametophyte dominated life cycle (see below).
The establishment of a land-based flora permitted the accumulation of oxygen in the atmosphere as
never before, as the new hordes of land plants pumped it out as a waste product. When this concentration
rose above 13%, it permitted the possibility of wildfire. This is first recorded in the early Silurian fossil record
by charcoalified plant fossils.[15] Apart from a controversial gap in the Late Devonian, charcoal is present
ever since.
Charcoalification is an important taphonomic mode. Wildfire drives off the volatile compounds,
leaving only a shell of pure carbon. This is not a viable food source for herbivores or detritovores, so is prone
to preservation; it is also robust, so can withstand pressure and display exquisite, sometimes sub-cellular,
detail.

[edit] Changing life cycles

Further information: Alternation of generations
 Angiosperm life cycle
All multicellular plants have a life cycle comprising two phases (often confusingly referred to as
"generations").[16] One is termed the gametophyte, has a single set of chromosomes (denoted 1n), and
produces gametes (sperm and eggs). The other is termed the sporophyte, has paired chromosomes
(denoted 2n), and produces spores. The two phases may be identical, or phenomenally different.
 The overwhelming pattern in plant evolution is for a reduction of the gametophytic phase, and the
increase in sporophyte dominance. The algal ancestors to land plants were almost certainly haplobiontic,
being haploid for all their life cycles, with a unicellular zygote providing the 2n stage. All land plants (i.e.
embryophytes) are diplobiontic – that is, both the haploid and diploid stages are multicellular.[16]
There are two competing theories to explain the appearance of a diplobiontic lifecycle.
The interpolation theory (also known as the antithetic or intercalary theory)[17] holds that the
sporophyte phase was a fundamentally new invention, caused by the mitotic division of a freshly germinated
zygote, continuing until meiosis produces spores. This theory implies that the first sporophytes would bear a
very different morphology to the gametophyte, on which they would have been dependent.[17] This seems to
fit well with what we know of the bryophytes, in which a vegetative thalloid gametophyte is parasitised by
simple sporophytes, which often comprise no more than a sporangium on a stalk. Increasing complexity of
the ancestrally simple sporophyte, including the eventual acquisition of photosynthetic cells, would free it
from its dependence on a gametophyte, as we see in some hornworts ( Anthoceros), and eventually result in
the sporophyte developing organs and vascular tissue, and becoming the dominant phase, as in the
tracheophytes (vascular plants).[16] This theory may be supported by observations that smaller Cooksonia
individuals must have been supported by a gametophyte generation. The observed appearance of larger
axial sizes, with room for photosynthetic tissue and thus self-sustainability, provides a possible route for the
development of a self-sufficient sporophyte phase.[17]
The alternative hypothesis is termed the transformation theory (or homologous theory). This posits
that the sporophyte appeared suddenly by a delay in the occurrence of meiosis after the zygote germinated.
Since the same genetic material would be employed, the haploid and diploid phases would look the same.
This explains the behaviour of some algae, which produce alternating phases of identical sporophytes and
gametophytes. Subsequent adaption to the desiccating land environment, which makes sexual reproduction
difficult, would result in the simplification of the sexually active gametophyte, and elaboration of the
sporophyte phase to better disperse the waterproof spores.[16] The tissue of sporophytes and gametophytes
preserved in the Rhynie chert is of similar complexity, which is taken to support this hypothesis.[17]

[edit] Water transport

In order to photosynthesise, plants must uptake CO 2 from the atmosphere. However, this comes at a
price: while stomata are open to allow CO2 to enter, water can evaporate.[18] Water is lost much faster than
CO2 is absorbed, so plants need to replace it, and have developed systems to transport water from the moist
soil to the site of photosynthesis.[18] Early plants sucked water between the walls of their cells, then evolved
the ability to control water loss (and CO2 acquisition) through the use of stomata. Specialised water transport
tissues soon evolved in the form of hydroids, tracheids, then secondary xylem, followed by an endodermis
and ultimately vessels.[18]
The high CO2 levels of the Silu-Devonian, when early plants were colonising land, meant that the
need for water was relatively low in the early days. As CO 2 was withdrawn from the atmosphere by plants,
more water was lost in its capture, and more elegant transport mechanisms evolved.[18] As water transport
mechanisms, and waterproof cuticles, evolved, plants could survive without being continually covered by a
film of water. This transition from poikilohydry to homoiohydry opened up new potential for colonisation.[18]
Plants were then faced with a balance, between transporting water as efficiently as possible and preventing
transporting vessels to implode and cavitate.
 During the Silurian, CO2 was readily available, so little water needed expending to acquire it. By the
end of the Carboniferous, when CO2 levels had lowered to something approaching today's, around 17 times
more water was lost per unit of CO2 uptake.[18] However, even in these "easy" early days, water was at a
premium, and had to be transported to parts of the plant from the wet soil to avoid desiccation. This early
water transport took advantage of the cohesion-tension mechanism inherent in water. Water has a tendency
to diffuse to areas that are drier, and this process is accelerated when water can be wicked along a fabric
with small spaces. In small passages, such as that between the plant cell walls (or in tracheids), a column of
water behaves like rubber – when molecules evaporate from one end, they literally pull the molecules behind
them along the channels. Therefore transpiration alone provided the driving force for water transport in early
plants.[18] However, without dedicated transport vessels, the cohesion-tension mechanism cannot transport
water more than about 2 cm, severely limiting the size of the earliest plants.[18] This process demands a
steady supply of water from one end, to maintain the chains; to avoid exhausing it, plants developed a
waterproof cuticle. Early cuticle may not have had pores but did not cover the entire plant surface, so that gas
exchange could continue.[18] However, dehydration at times was inevitable; early plants cope with this by
having a lot of water stored between their cell walls, and when it comes to it sticking out the tough times by
putting life "on hold" until more water is supplied.[18]
 A banded tube from the late Silurian/early Devonian. The bands are difficult to see on this specimen,
as an opaque carbonaceous coating conceals much of the tube. Bands are just visible in places on the left
half of the image – click on the image for a larger view. Scale bar: 20 μm
In order to be free from the constraints of small size and constant moisture that the parenchymatic
transport system inflicted, plants needed a more efficient water transport system. During the early Silurian,
they developed specialized cells, which were lignified (or bore similar chemical compounds)[18] to avoid
implosion; this process coincided with cell death, allowing their innards to be emptied and water to be passed
through them.[18] These wider, dead, empty cells were a million times more conductive than the inter-cell
method, giving the potential for transport over longer distances, and higher CO 2 diffusion rates.

The first macrofossils to bear water-transport tubes in situ are the early Devonian pretracheophytes
Aglaophyton and Horneophyton, which have structures very similar to the hydroids of modern mosses. Plants
continued to innovate new ways of reducing the resistance to flow within their cells, thereby increasing the
efficiency of their water transport. Bands on the walls of tubes, in fact apparent from the early Silurian
onwards,[19] are an early improvisation to aid the easy flow of water.[20] Banded tubes, as well as tubes with
pitted ornamentation on their walls, were lignified[21] and, when they form single celled conduits, are
considered to be tracheids. These, the "next generation" of transport cell design, have a more rigid structure
than hydroids, allowing them to cope with higher levels of water pressure.[18] Tracheids may have a single
evolutionary origin, possibly within the hornworts,[22] uniting all tracheophytes (but they may have evolved
more than once).[18]
Water transport requires regulation, and dynamic control is provided by stomata.[23] By adjusting the
amount of gas exchange, they can restrict the amount of water lost through transpiration. This is an important
role where water supply is not constant, and indeed stomata appear to have evolved before tracheids, being
present in the non-vascular hornworts.[18]
An endodermis probably evolved during the Silu-Devonian, but the first fossil evidence for such a
structure is Carboniferous.[18] This structure in the roots covers the water transport tissue and regulates ion
exchange (and prevents unwanted pathogens etc. from entering the water transport system). The
endodermis can also provide an upwards pressure, forcing water out of the roots when transpiration is not
enough of a driver.
Once plants had evolved this level of controlled water transport, they were truly homoiohydric, able to
extract water from their environment through root-like organs rather than relying on a film of surface moisture,
enabling them to grow to much greater size.[18] As a result of their independence from their surroundings,
they lost their ability to survive desiccation – a costly trait to retain.[18]
During the Devonian, maximum xylem diameter increased with time, with the minimum diameter
remaining pretty constant.[20] By the middle Devonian, the tracheid diameter of some plant lineages[24] had
plateaued.[20] Wider tracheids allow water to be transported faster, but the overall transport rate depends
also on the overall cross-sectional area of the xylem bundle itself.[20] The increase in vascular bundle
thickness further seems to correlate with the width of plant axes, and plant height; it is also closely related to
the appearance of leaves[20] and increased stomatal density, both of which would increase the demand for
water.[18]
While wider tracheids with robust walls make it possible to achieve higher water transport pressures,
this increases the problem of cavitation.[18] Cavitation occurs when a bubble of air forms within a vessel,
breaking the bonds between chains of water molecules and preventing them from pulling more water up with
their cohesive tension. A tracheid, once cavitated, cannot have its embolism removed and return to service
(except in a few advanced angiosperms[verification needed] which have developed a mechanism of doing
so). Therefore it is well worth plants' while to avoid cavitation occurring. For this reason, pits in tracheid walls
have very small diameters, to prevent air entering and allowing bubbles to nucleate.[18] Freeze-thaw cycles
are a major cause of cavitation.[18] Damage to a tracheid's wall almost inevitably leads to air leaking in and
cavitation, hence the importance of many tracheids working in parallel.[18]
Cavitation is hard to avoid, but once it has occurred plants have a range of mechanisms to contain
the damage.[18] Small pits link adjacent conduits to allow fluid to flow between them, but not air – although
ironically these pits, which prevent the spread of embolisms, are also a major cause of them.[18] These
pitted surfaces further reduce the flow of water through the xylem by as much as 30%.[18] Conifers, by the
Jurassic, developed an ingenious improvement, using valve-like structures to isolate cavitated elements.
These torus-margo structures have a blob floating in the middle of a donut; when one side depressurises the
blob is sucked into the torus and blocks further flow.[18] Other plants simply accept cavitation; for instance,
oaks grow a ring of wide vessels at the start of each spring, none of which survive the winter frosts. Maples
use root pressure each spring to force sap upwards from the roots, squeezing out any air bubbles.
Growing to height also employed another trait of tracheids – the support offered by their lignified
walls. Defunct tracheids were retained to form a strong, woody stem, produced in most instances by a
secondary xylem. However, in early plants, tracheids were too mechanically vulnerable, and retained a
central position, with a layer of tough sclerenchyma on the outer rim of the stems.[18] Even when tracheids
do take a structural role, they are supported by sclerenchymatic tissue.
Tracheids end with walls, which impose a great deal of resistance on flow;[20] vessel members have
perforated end walls, and are arranged in series to operate as if they were one continuous vessel.[20] The
function of end walls, which were the default state in the Devonian, was probably to avoid embolisms. An
embolism is where an air bubble is created in a tracheid. This may happen as a result of freezing, or by
gases dissolving out of solution. Once an embolism is formed, it usually cannot be removed (but see later);
the affected cell cannot pull water up, and is rendered useless.
End walls excluded, the tracheids of prevascular plants were able to operate under the same
hydraulic conductivity as those of the first vascular plant, Cooksonia.[20]
The size of tracheids is limited as they comprise a single cell; this limits their length, which in turn
limits their maximum useful diameter to 80 μm.[18] Conductivity grows with the fourth power of diameter, so
increased diameter has huge rewards; vessel elements, consisting of a number of cells, joined at their ends,
overcame this limit and allowed larger tubes to form, reaching diameters of up to 500 μm, and lengths of up
to 10 m.[18]
Vessels first evolved during the dry, low CO2 periods of the late Permian, in the horsetails, ferns and
Selaginellales independently, and later appeared in the mid Cretaceous in angiosperms and gnetophytes.
[18] Vessels allow the same cross-sectional area of wood to transport around a hundred times more water
than tracheids![18] This allowed plants to fill more of their stems with structural fibres, and also opened a new
niche to vines, which could transport water without being as thick as the tree they grew on.[18] Despite these
advantages, tracheid-based wood is a lot lighter, thus cheaper to make, as vessels need to be much more
reinforced to avoid cavitation.[18]
[edit] Evolution of leaves
 The lycopod Isoetes bears microphylls with a single vascular trace.

The branching pattern of megaphyll veins may belie their origin as webbed, dichotomising branches.
Leaves today are, in almost all instances, an adaptation to increase the amount of sunlight that can
be captured for photosynthesis. Leaves certainly evolved more than once, and probably originated as spiny
outgrowths to protect early plants from herbivory.
The rhyniophytes of the Rhynie chert comprised nothing more than slender, unornamented axes.
The early to middle Devonian trimerophytes, therefore, are the first evidence we have of anything that could
be considered leafy. This group of vascular plants are recognisable by their masses of terminal sporangia,
which adorn the ends of axes which may bifurcate or trifurcate.[23] Some organisms, such as Psilophyton,
bore enations. These are small, spiny outgrowths of the stem, lacking their own vascular supply.
Around the same time, the zosterophyllophytes were becoming important. This group is recognisable
by their kidney-shaped sporangia, which grew on short lateral branches close to the main axes. They
sometimes branched in a distinctive H-shape.[23] The majority of this group bore pronounced spines on their
axes. However, none of these had a vascular trace, and the first evidence of vascularised enations occurs in
the Rhynie genus Asteroxylon. The spines of Asteroxylon had a primitive vasuclar supply – at the very least,
leaf traces could be seen departing from the central protostele towards each individual "leaf". A fossil known
as Baragwanathia appears in the fossil record slightly earlier, in the late Silurian.[25] In this organism, these
leaf traces continue into the leaf to form their mid-vein.[26] One theory, the "enation theory", holds that the
leaves developed by outgrowths of the protostele connecting with existing enations, but it is also possible
that microphylls evolved by a branching axis forming "webbing".[23]
Asteroxylon[27] and Baragwanathia are widely regarded as primitive lycopods.[23] The lycopods are
still extant today, familiar as the quillwort Isoetes and the club mosses. Lycopods bear distinctive microphylls
– leaves with a single vascular trace. Microphylls could grow to some size – the Lepidodendrales boasted
microphylls over a meter in length – but almost all just bear the one vascular bundle. (An exception is the
branching Selaginella).
The more familiar leaves, megaphylls, are thought to have separate origins – indeed, they appeared
four times independently, in the ferns, horsetails, progymnosperms, and seed plants.[28] They appear to
have originated from dichotomising branches, which first overlapped (or "overtopped") one another, and
eventually developed "webbing" and evolved into gradually more leaf-like structures.[26] So megaphylls, by
this "teleome theory", are composed of a group of webbed branches[26] – hence the "leaf gap" left where the
leaf's vascular bundle leaves that of the main branch resembles two axes splitting.[26] In each of the four
groups to evolve megaphylls, their leaves first evolved during the late Devonian to early Carboniferous,
diversifying rapidly until the designs settled down in the mid Carboniferous.[28]
The cessation of further diversification can be attributed to developmental constraints,[28] but why
did it take so long for leaves to evolve in the first place? Plants had been on the land for at least 50 million
years before megaphylls became significant. However, small, rare mesophylls are known from the early
Devonian genus Eophyllophyton – so development could not have been a barrier to their appearance.[29]
The best explanation so far incorporates observations that atmospheric CO 2 was declining rapidly during this
time – falling by around 90% during the Devonian.[30] This corresponded with an increase in stomatal density
by 100 times. Stomata allow water to evaporate from leaves, which causes them to curve. It appears that the
low stomatal density in the early Devonian meant that evaporation was limited, and leaves would overheat if
they grew to any size. The stomatal density could not increase, as the primitive steles and limited root
systems would not be able to supply water quickly enough to match the rate of transpiration.[31]
Clearly, leaves are not always beneficial, as illustrated by the frequent occurrence of secondary loss
of leaves, famously exemplified by cacti and the "whisk fern" Psilotum.
Secondary evolution can also disguise the true evolutionary origin of some leaves. Some genera of
ferns display complex leaves which are attached to the pseudostele by an outgrowth of the vascular bundle,
leaving no leaf gap.[26] Further, horsetail (Equisetum) leaves bear only a single vein, and appear for all the
world to be microphyllous; however, in the light of the fossil record and molecular evidence, we conclude that
their forbears bore leaves with complex venation, and the current state is a result of secondary simplification.
[32]
Deciduous trees deal with another disadvantage to having leaves. The popular belief that plants shed
their leaves when the days get too short is misguided; evergreens prospered in the Arctic circle during the
most recent greenhouse earth.[33] The generally accepted reason for shedding leaves during winter is to
cope with the weather – the force of wind and weight of snow are much more comfortably weathered without
leaves to increase surface area. Seasonal leaf loss has evolved independently several times and is exhibited
in the ginkgoales, pinophyta and angiosperms.[34] Leaf loss may also have arisen as a response to pressure
from insects; it may have been less costly to lose leaves entirely during the winter or dry season than to
continue investing resources in their repair.[35]

[edit] Evolution of trees

 The trunk of early tree fern Psaronius, showing internal structure. The top of the plant would have
been to the left of the image

External mold of Lepidodendron trunk showing leaf scars from the Upper Carboniferous of Ohio
The early Devonian landscape was devoid of vegetation taller than waist height. Without the
evolution of a robust vascular system, taller heights could not be obtained. There was, however, a constant
evolutionary pressure to attain greater height. The most obvious advantage is the harvesting of more sunlight
for photosynthesis – by overshadowing competitors – but a further advantage is present in spore distribution,
as spores (and, later, seeds) can be blown greater distances if they start higher. This may be demonstrated
by Prototaxites, thought to be a late Silurian fungus reaching eight metres in height.[36]
In order to attain arborescence, early plants needed to develop woody tissue that would act as both
support and water transport. To understand wood, we must know a little of vascular behaviour. The stele of
plants undergoing "secondary growth" is surrounded by the vascular cambium, a ring of cells which produces
more xylem (on the inside) and phloem (on the outside). Since xylem cells comprise dead, lignified tissue,
subsequent rings of xylem are added to those already present, forming wood.
The first plants to develop this secondary growth, and a woody habit, were apparently the ferns, and
as early as the middle Devonian one species, Wattieza, had already reached heights of 8 m and a tree-like
habit.[37]
Other clades did not take long to develop a tree-like stature; the late Devonian Archaeopteris, a
precursor to gymnosperms which evolved from the trimerophytes,[38] reached 30 m in height. These
progymnosperms were the first plants to develop true wood, grown from a bifacial cambium, of which the first
appearance is in the mid Devonian Rellimia.[39] True wood is only thought to have evolved once, giving rise
to the concept of a "lignophyte" clade.
These Archaeopteris forests were soon supplemented by lycopods, in the form of lepidodendrales,
which topped 50m in height and 2m across at the base. These lycopods rose to dominate late Devonian and
Carboniferous coal deposits.[40] Lepidodendrales differ from modern trees in exhibiting determinate growth:
after building up a reserve of nutrients at a low height, the plants would "bolt" to a genetically determined
height, branch at that level, spread their spores and die.[41] They consisted of "cheap" wood to allow their
rapid growth, with at least half of their stems comprising a pith-filled cavity.[23] Their wood was also
generated by a unifacial vascular cambium – it did not produce new phloem, meaning that the trunks could
not grow wider over time.[verification needed]
The horsetail Calamites was next on the scene, appearing in the Carboniferous. Unlike the modern
horsetail Equisetum, Calamites had a unifacial vascular cambium, allowing them to develop wood and grow
to heights in excess of 10 m. They also branched multiple times.
 While the form of early trees was similar to that of today's, the groups containing all modern trees
had yet to evolve.
The dominant groups today are the gymnosperms, which include the coniferous trees, and the
angiosperms, which contain all fruiting and flowering trees. It was long thought that the angiosperms arose
from within the gymnosperms, but recent molecular evidence suggests that their living representatives form
two distinct groups.[42][43][44] It must be noted that the molecular data has yet to be fully reconciled with
morphological data,[45][46][47] but it is becoming accepted that the morphological support for paraphyly is
not especially strong.[48] This would lead to the conclusion that both groups arose from within the
pteridosperms, probably as early as the Permian.[48]
The angiosperms and their ancestors played a very small role until they diversified during the
Cretaceous. They started out as small, damp-loving organisms in the understory, and have been diversifying
ever since the mid[verification needed]-Cretaceous, to become the dominant member of non-boreal forests
today.
[edit] Evolution of roots
Roots are important to plants for two main
reasons: Firstly, they provide anchorage to the substrate;
more importantly, they provide a source of water and
nutrients from the soil. Roots allowed plants to grow taller
and faster.
The onset of roots also had effects on a global
scale. By disturbing the soil, and promoting its
acidification (by taking up nutrients such as nitrate and
phosphate[verification needed]), they enabled it to
weather more deeply, promoting the draw-down of
CO2[49] with huge implications for climate.[50] These
effects may have been so profound they led to a mass
extinction.[51]
But how and when did roots evolve in the first
place? While there are traces of root-like impressions in
fossil soils in the late Silurian,[52] body fossils show the
earliest plants to be devoid of roots. Many had tendrils
which sprawled along or beneath the ground, with upright
axes or thalli dotted here and there, and some even had
non-photosynthetic subterranean branches which lacked
stomata. The distinction between root and specialised
The roots (bottom image) of
lepidodendrales are thought to be functionally
equivalent to the stems (top), as the similar
appearance of "leaf scars" and "root scars" on
these specimens from different species
demonstrates.
branch is developmental; true roots follow a different developmental trajectory to stems. Further, roots differ
in their branching pattern, and in possession of a root cap.[3] So while Silu-Devonian plants such as Rhynia
and Horneophyton possessed the physiological equivalent of roots, roots – defined as organs differentiated
from stems – did not arrive until later.[3] Unfortunately, roots are rarely preserved in the fossil record, and our
understanding of their evolutionary origin is sparse.[3]
Rhizoids – small structures performing the same role as roots, usually a cell in diameter – probably
evolved very early, perhaps even before plants colonised the land; they are recognised in the Characeae, an
algal sister group to land plants.[3] That said, rhizoids probably evolved more than once; the rhizines of
lichens, for example, perform a similar role. Even some animals ( Lamellibrachia) have root-like structures![3]
More advanced structures are common in the Rhynie chert, and many other fossils of comparable
early Devonian age bear structures that look like, and acted like, roots.[3] The rhyniophytes bore fine
rhizoids, and the trimerophytes and herbaceous lycopods of the chert bore root-like structure penetrating a
few centimetres into the soil.[53] However, none of these fossils display all the features borne by modern
roots.[3] Roots and root-like structures became increasingly more common and deeper penetrating during
the Devonian period, with lycopod trees forming roots around 20 cm long during the Eifelian and Givetian.
These were joined by progymnosperms, which rooted up to about a metre deep, during the ensuing Frasnian
stage.[53] True gymnosperms and zygopterid ferns also formed shallow rooting systems during the
Famennian period.[53]
The rhizomorphs of the lycopods provide a slightly approach to rooting. They were equivalent to
stems, with organs equivalent to leaves performing the role of rootlets.[3] A similar construction is observed
in the extant lycopod Isoetes, and this appears to be evidence that roots evolved independently at least
twice, in the lycophytes and other plants.[3]
 A vascular system is indispensable to a rooted plants, as non-photosynthesising roots need a supply
of sugars, and a vascular system is required to transport water and nutrients from the roots to the rest of the
plant.[4] These plants are little more advanced than their Silurian forbears, without a dedicated root system;
however, the flat-lying axes can be clearly seen to have growths similar to the rhizoids of bryophytes today.
[54]
By the mid-to-late Devonian, most groups of plants had independently developed a rooting system of
some nature.[54] As roots became larger, they could support larger trees, and the soil was weathered to a
greater depth.[51] This deeper weathering had effects not only on the aforementioned drawdown of CO 2, but
also opened up new habitats for colonisation by fungi and animals.[53]
Roots today have developed to the physical limits. They penetrate many[ quantify] metres of soil to
tap the water table.[verification needed] The narrowest roots are a mere 40 μm in diameter, and could not
physically transport water if they were any narrower.[3] The earliest fossil roots recovered, by contrast,
narrowed from 3 mm to under 700 μm in diameter; of course, taphonomy is the ultimate control of what
thickness we can see.[3]

[edit] Arbuscular mycorrhizae

The efficiency of many plants' roots is increased via a symbiotic relationship with a fungal partner.
The most common are arbuscular mycorrhizae (AM), literally "tree-like fungal roots". These comprise fungi
which invade some root cells, filling the cell membrane with their hyphae. They feed on the plant's sugars,
but return nutrients generated or extracted from the soil (especially phosphate), which the plant would
otherwise have no access to.
 This symbiosis appears to have evolved early in plant history. AM are found in all plant groups, and
80% of extant vascular plants,[55] suggesting an early ancestry; a "plant"-fungus symbiosis may even have
been the step that enabled them to colonise the land,[56] and indeed AM are abundant in the Rhynie chert;
[57] the association occurred even before there were true roots to colonise, and is has even been suggested
that roots evolved in order to provide a more comfortable habitat for mycorrhizal fungi.[58]

[edit] Evolution of seeds

The fossil seed Trigonocarpus

Early land plants reproduced in the fashion of ferns: spores germinated into small gametophytes,
which produced sperm. These would swim across moist soils to find the female organs (archegonia) on the
same or another gametophyte, where they would fuse with an ovule to produce an embryo, which would
germinate into a sporophyte.[53]
This mode of reproduction restricted early plants to damp environments, moist enough that the
sperm could swim to their destination. Therefore, early land plants were constrained to the lowlands, near
shores and streams. The development of heterospory freed them from this constraint.
Heterosporic organisms, as their name suggests, bear spores of two sizes – microspores and
megaspores. These would germinate to form microgametophytes and megagametophytes, respectively. This
system paved the way for seeds: taken to the extreme, the megasporangia could bear only a single
megaspore tetrad, and to complete the transition to true seeds, three of the megaspores in the original tetrad
could be aborted, leaving one megaspore per megasporangium.
The transition to seeds continued with this megaspore being "boxed in" to its sporangium while it
germinates. Then, the megagametophyte is contained within a waterproof integument, which forms the bulk
of the seed. The microgametophyte – a pollen grain which has germinated from a microspore – is employed
for dispersal, only releasing its desiccation-prone sperm when it reaches a receptive megagametophyte.[23]
Lycopods go a fair way down the path to seeds without ever crossing the threshold. Fossil lycopod
megaspores reaching 1 cm in diameter, and surrounded by vegetative tissue, are known – these even
germinate into a megagametophyte in situ. However, they fall short of being seeds, since the nucellus, an
inner spore-covering layer, does not completely enclose the spore. A very small slit remains, meaning that
the seed is still exposed to the atmosphere. This has two consequences – firstly, it means it is not fully
resistant to desiccation, and secondly, sperm do not have to "burrow" to access the archegonia of the
megaspore.[23]
 The first "spermatophytes" (literally:seed plants) – that is, the first plants to bear true seeds – are
called pteridosperms: literally, "seed ferns", so called because their foliage consisted of fern-like fronds,
although they were not closely related to ferns. The oldest fossil evidence of seed plants is of Late Devonian
age and they appear to have evolved out of an earlier group known as the progymnosperms. These early
seed plants ranged from trees to small, rambling shrubs; like most early progymnosperms, they were woody
plants with fern-like foliage. They all bore ovules, but no cones, fruit or similar. While it is difficult to track the
early evolution of seeds, we can trace the lineage of the seed ferns from the simple trimerophytes through
homosporous Aneurophytes.[23]
This seed model is shared by basically all gymnosperms (literally: "naked seeds"), most of which
encase their seeds in a woody or fleshy (the yew, for example) cone, but none of which fully enclose their
seeds. The angiosperms ("vessel seeds") are the only group to fully enclose the seed, in a carpel.
Fully enclosed seeds opened up a new pathway for plants to follow: that of seed dormancy. The
embryo, completely isolated from the external atmosphere and hence protected from desiccation, could
survive some years of drought before germinating. Gymnosperm seeds from the late Carboniferous have
been found to contain embryos, suggesting a lengthy gap between fertilisation and germination.[59] This
period is associated with the entry into a greenhouse earth period, with an associated increase in aridity. This
suggests that dormancy arose as a response to drier climatic conditions, where it became advantageous to
wait for a moist period before germinating.[59] This evolutionary breakthrough appears to have opened a
floodgate: previously inhospitable areas, such as dry mountain slopes, could now be tolerated, as were soon
covered by trees.[59]
Seeds offered further advantages to their bearers: they increased the success rate of fertilised
gametophytes, and because a nutrient store could be "packaged" in with the embryo, the seeds could
germinate rapidly in inhospitable environments, reaching a size where it could fend for itself more quickly.[53]
For example, without an endosperm, seedlings growing in arid environments would not have the reserves to
grow roots deep enough to reach the water table before they expired.[53] Likewise, seeds germinating in a
gloomy understory require an additional reserve of energy to quickly grow high enough to capture sufficient
light for self-sustenance.[53] A combination of these advantages gave seed plants the ecological edge over
the previously dominant genus Archaeopteris, this increasing the biodiversity of early forests.[53]

[edit] Evolution of flowers

The pollen bearing organs of the early "flower" Crossotheca

 The evolution of syncarps.
a: sporangia borne at tips of leaf
b: Leaf curls up to protect sporangia
c: leaf curls to form enclosed roll
d: grouping of three rolls into a syncarp
 Flowers are modified leaves possessed only by the group known as the angiosperms, which are
relatively late to appear in the fossil record. Colourful and/or pungent structures surround the cones of plants
such as cycads and gnetales, making a strict definition of the term "flower" elusive.[47]
The flowering plants have long been assumed to have evolved from within the gymnosperms;
according to the traditional morphological view, they are closely allied to the gnetales. However, as noted
above, recent molecular evidence is at odds to this hypothesis,[43][44] and further suggests that gnetales are
more closely related to some gymnosperm groups than angiosperms,[42] and that extant gymnosperms form
a distinct clade to the angiosperms,[43][44][42] the two clades diverging some 300 million years ago.[60]
Phylogeny of anthophytes and gymnosperms, from [61]

Cycads

Ginkgo

Conifers

Anthophytes

Bennettitales
Gnetales

Angiosperms

Angiosperms

Cycads

Bennettitales
 Ginkgo

Conifers

Gnetales

Traditional view Modern view

 Further information: Gnetophyta#Classification
The relationship of stem groups to the angiosperms is of utmost importance in determining the
evolution of flowers; stem groups provide an insight into the state of earlier "forks" on the path to the current
state. If we identify an unrelated group as a stem group, then we will gain an incorrect image of the lineages'
history. The traditional view that flowers arose by modification of a structure similar to that of the gnetales, for
example, no longer bears weight in the light of the molecular data.
Convergence increases our chances of misidentifying stem groups. Since the protection of the
megagametophyte is evolutionarily desirable, it would be unsurprising if many separate groups stumbled
upon protective encasements independently. Distinguishing ancestry in such a situation, especially where we
usually only have fossils to go on, is tricky – to say the least.[ clarification needed]
In flowers, this protection is offered by the carpel, an organ believed to represent an adapted leaf,
recruited into a protective role, shielding the ovules. These ovules are further protected by a double-walled
integument.
Penetration of these protective layers needs something more that a free-floating microgametophyte.
Angiosperms have pollen grains comprising just three cells. One cell is responsible for drilling down through
the integuments, and creating a conduit for the two sperm cells to flow down. The megagametophyte has just
seven cells; of these, one fuses with a sperm cell, forming the nucleus of the egg itself, and another other
joins with the other sperm, and dedicates itself to forming a nutrient-rich endosperm. The other cells take
auxiliary roles.[clarification needed] This process of "double fertilisation" is unique and common to all
angiosperms.

The inflorescences of the Bennettitales are strikingly similar to flowers

In the fossil record, there are three intriguing groups which bore flower-like structures. The first is the
Permian pteridosperm Glossopteris, which already bore recurved leaves resembling carpels. The Triassic
Caytonia is more flower-like still, with enclosed ovules – but only a single integument. Further, details of their
pollen and stamens set them apart from true flowering plants.
 The Bennettitales bore remarkably flower-like organs, protected by whorls of bracts which may have
played a similar role to the petals and sepals of true flowers; however, these flower-like structures evolved
independently, as the Bennettitales are more closely related to cycads and ginkgos than to the angiosperms.
[61]
However, no true flowers are found in any groups save those extant today. Most morphological and
molecular analyses place Amborella, the nymphaeales and Austrobaileyaceae in a basal clade dubbed
"ANA". This clade appear to have diverged in the early Cretaceous, around 130 million years ago – around
the same time as the earliest fossil angiosperm,[62][63] and just after the first angiosperm-like pollen, 136
million years ago.[48] The magnoliids diverged soon after, and a rapid radiation had produced eudicots and
monocots by 125 million years ago.[48] By the end of the Cretaceous 65.5 million years ago, over 50% of
today's angiosperm orders had evolved, and the clade accounted for 70% of global species.[64] It was
around this time that flowering trees became dominant over conifers [65]
The features of the basal "ANA" groups suggest that angiosperms originated in dark, damp,
frequently disturbed areas.[66] It appears that the angiosperms remained constrained to such habitats
throughout the Cretaceous – occupying the niche of small herbs early in the successional series.[64] This
may have restricted their initial significance, but given them the flexibility that accounted for the rapidity of
their later diversifications in other habitats.[66]
[edit] Advances in metabolism
 The C4 carbon concentrating mechanism

The most recent major innovation by the plants is the development of the C4 metabolic pathway.

Photosynthesis is not quite as simple as adding water to CO2 to produce sugars and oxygen. A
complex chemical pathway is involved, facilitated along the way by a range of enzymes and co-enzymes.
The enzyme RuBisCO is responsible for "fixing" CO2 – that is, it attaches it to a carbon-based molecule to
form a sugar, which can be used by the plant, releasing an oxygen molecule along the way. However, the
enzyme is notoriously inefficient, and just as effectively will also fix oxygen instead of CO 2 in a process called
photorespiration. This is energetically costly as the plant has to use energy to turn the products of
photorepsiration back into a form that can react with CO 2.

[edit] Concentrating carbon

To work around this inefficiency, C4 plants evolved carbon concentrating mechanisms. These work
by increasing the concentration of CO2 around RuBisCO, thereby increasing the amount of photosynthesis
and decreasing photorespiration. The process of concentrating CO 2 around RuBisCO requires more energy
than allowing gases to diffuse, but under certain conditions – i.e. warm temperatures (>25°C), low CO 2
concentrations, or high oxygen concentrations – pays off in terms of the decreased loss of sugars through
photorespiration.
 One, C4 metabolism, employs a so-called Kranz anatomy. This transports CO2 through an outer
mesophyll layer, via a range of organic molecules, to the central bundle sheath cells, where the CO 2 is
released. In this way, CO2 is concentrated near the site of RuBisCO operation. Because RuBisCO is
operating in an environment with much more CO2 than it otherwise would be, it performs more efficiently.

A second method, CAM photosynthesis, temporally separates photosynthesis from the action of
RuBisCO. RuBisCO only operates during the day, when stomata are sealed and CO 2 is provided by the
breakdown of the chemical malate. More CO2 is then harvested from the atmosphere when stomata open,
during the cool, moist nights, reducing water loss.

[edit] Evolutionary record

These two pathways, with the same effect on RuBisCO, evolved a number of times independently –
indeed, C4 alone arose in 18 different plant families. The C4 construction is most famously used by a subset
of grasses, while CAM is employed by many succulents and cacti. The trait appears to have emerged during
the Oligocene, around 25 to 32 million years ago;[67] however, they did not become ecologically significant
until the Miocene, -1 million years ago.[68] Remarkably, some charcoalified fossils preserve tissue organised
into the Kranz anatomy, with intact bundle sheath cells,[69] allowing the presence C4 metabolism to be
identified without doubt at this time. In deducing their distribution and significance, we resort to the use of
isotopic markers. C3 plants preferentially use the lighter of two isotopes of carbon in the atmosphere, 12C,
which is more readily involved in the chemical pathways involved in its fixation. Because C 4 metabolism
involves a further chemical step, this effect is accentuated. Plant material can be analysed to deduce the
ratio of the heavier 13C to 12C. This ratio is denoted δ13C. C3 plants are on average around 14‰ (parts per
thousand) lighter than the atmospheric ratio, while C4 plants are about 28‰ lighter. The δ13C of CAM plants
depends on the percentage of carbon fixed at night relative to what is fixed in the day, being closer to C 3
plants if they fix most carbon in the day and closer to C4 plants if they fix all their carbon at night.[70]

It's troublesome procuring original fossil material in sufficient quantity to analyse the grass itself, but
fortunately we have a good proxy: horses. Horses were globally widespread in the period of interest, and
browsed almost exclusively on grasses. There's an old phrase in isotope palæontology, "you are what you
eat (plus a little bit)" – this refers to the fact that organisms reflect the isotopic composition of whatever they
eat, plus a small adjustment factor. There is a good record of horse teeth throughout the globe, and their
δ13C has been measured. The record shows a sharp negative inflection around -1 million years ago, during
the Messinian, and this is interpreted as the rise of C4 plants on a global scale.[68]

[edit] When is C4 an advantage?

While C4 enhances the efficiency of RuBisCO, the concentration of carbon is highly energy intensive.
This means that C4 plants only have an advantage over C3 organisms in certain conditions: namely, high
temperatures and low rainfall. C4 plants also need high levels of sunlight in order to thrive.[71] Models
suggest that without wildfires removing shade-casting trees and shrubs, there would be no space for C 4
plants.[72] But wildfires have occurred for 400 million years – why did C 4 take so long to arise, and then
appear independently so many times? The Carboniferous period (~300 million years ago) had notoriously
high oxygen levels – almost enough to allow spontaneous combustion[73] – and very low CO2, but there is no
C4 isotopic signature to be found. And there doesn't seem to be a sudden trigger for the Miocene rise.

During the Micoene, the atmosphere and climate was relatively stable. If anything, CO 2 increased
gradually from 14 to 9 million years ago before settling down to concentrations similar to the Holocene.[74]
This suggests that it did not have a key role in invoking C 4 evolution.[67] Grasses themselves (the group
which would give rise to the most occurrences of C4) had probably been around for 60 million years or more,
so had had plenty of time to evolve C4,[75][76] which in any case is present in a diverse range of groups and
thus evolved independently. There is a strong signal of climate change in South Asia;[67] increasing aridity –
hence increasing fire frequency and intensity – may have led to an increase in the importance of grasslands.
[77] However, this is difficult to reconcile with the North American record.[67] It is possible that the signal is
entirely biological, forced by the fire- (and elephant?)-[78] driven acceleration of grass evolution – which, both
by increasing weathering and incorporating more carbon into sediments, reduced atmospheric CO 2 levels.
[78] Finally, there is evidence that the onset of C 4 from 9 to 7 million years ago is a biased signal, which only
holds true for North America, from where most samples originate; emerging evidence suggests that
grasslands evolved to a dominant state at least 15Ma earlier in South America.

[edit] Evolutionary trends

The process of evolution works slightly differently in plants than animals. Differences in plant
physiology and reproduction mean that while the same evolutionary principles of natural selection apply, the
finer nuances of their effect are radically different.
 One major difference is the ability of plants to reproduce clonally, and the totipotent nature of their
cells, allowing them to reproduce asexually much more easily than most animals. They are also capable of
polyploidy – where more than two chromosome sets are inherited from parents. This allows relatively fast
bursts of evolution to occur. The long periods of dormancy that seed plants can employ also makes them less
vulnerable to extinction, as they can "sit out" the tough periods and wait until more clement times to leap back
to life.
The effect of these differences is most profoundly seen during extinction events. These events, which
wiped out between 6 and 62% of terrestrial animal families, had "negligible" effect on plant families.[79]
However, the ecosystem structure is significantly rearranged, with the abundances and distributions of
different groups of plants changing profoundly.[79] These effects are perhaps due to the higher diversity
within families, as extinction – which was common at the species level – was very selective. For example,
wind-pollinated species survived better than insect-pollinated taxa, and specialised species generally lost
out.[79] In general, the surviving taxa were rare before the extinction, suggesting that they were generalists
who were poor competitors when times were easy, but prospered when specialised groups went extinct and
left ecological niches vacant.[79]

[edit] See also

Plants
• Plant
• Flora
General evolution
 • Evolutionary history of life
• Timeline of plant evolution
• Timeline of evolution
Study of plants
• Paleobotany
• Plant evolutionary developmental biology
• Cryptospores
Plant interactions
• Evolution of herbivory

[edit] References
1. ^ "The oldest fossils reveal evolution of non-vascular plants by the middle to late Ordovician
Period (~450-440 m.y.a.) on the basis of fossil spores" Transition of plants to land
2. ^ Rothwell, G. W., Scheckler, S. E. & Gillespie, W. H. (1989). " Elkinsia gen. nov., a Late
Devonian gymnosperm with cupulate ovules." Botanical Gazette, 150: 170-189.
3. ^ a b c d e f g h i j k l m n o Raven, J.A.; Edwards, D. (2001). "Roots: evolutionary origins and
biogeochemical significance" (in active DOI due to publisher error (2008-04-30)). Journal of
Experimental Botany 52 (90001): 381–401. doi:10.1093/jexbot/52.suppl_1.381.
http://jxb.oxfordjournals.org/cgi/content/full/52/suppl_1/381.
 4. ^ a b P. Kenrick, P.R. Crane (1997) (1997). The origin and early diversification of land plants.
A cladistic study. Smithsonian Institution Press, Washington & London . Washington: Smithsonian
Inst. Press. ISBN 1-56098-729-4.
5. ^ Heckman; Geiser, D. M.; Eidell, B.R.; Stauffer, R. L.; Kardos, N. L; Hedges, S. B. (Aug
2001). "Molecular evidence for the early colonization of land by fungi and plants". Science (New
York, N.Y.) 293 (5532): 1129–1133. doi:10.1126/science.1061457. ISSN 0036-8075.
PMID 11498589. edit
6. ^ Battison, Leila; Brasier, Martin D. (August 2009). "Exceptional Preservation of Early
Terrestrial Communities in Lacustrine Phosphate One Billion Years Ago". in Smith, Martin R.;
O'Brien, Lorna J.; Caron, Jean-Bernard. International Conference on the Cambrian Explosion
(Walcott 2009). Abstract Volume. Toronto, Ontario, Canada: The Burgess Shale Consortium. 31st
July 2009. ISBN 978-0-9812885-1-2. http://burgess-shale.info/abstract/brasier-1.
7. ^ Knauth, P.; Kennedy, J. (Aug 2009). "The late Precambrian greening of the Earth.". Nature
460 (7256): 728–732. doi:10.1038/nature08213. ISSN 0028-0836. PMID 19587681. edit
8. ^ a b c d e Gray, J.; Chaloner, W. G.; Westoll, T. S. (1985). "The Microfossil Record of Early
Land Plants: Advances in Understanding of Early Terrestrialization, 1970-1984". Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences (1934-1990) 309 (1138):
167–195. doi:10.1098/rstb.1985.0077. http://links.jstor.org/sici?sici=0080-
4622(19850402)309%3A1138%3C167%3ATMROEL%3E2.0.CO%3B2-E. Retrieved 2008-04-26.
9. ^ Wellman, H.; Gray, J. (Jun 2000). "The microfossil record of early land plants".
Philosophical Transactions: Biological Sciences 355 (1398): 717–732. doi:10.1098/rstb.2000.0612.
ISSN 0962-8436. PMID 10905606. PMC 1692785.
http://rstb.royalsocietypublishing.org/content/355/1398/717. edit
 10.^ Wellman, C.H.; Osterloff, P.L.; Mohiuddin, U. (2003). "Fragments of the earliest land
plants". Nature 425 (6955): 282–285. doi:10.1038/nature01884. PMID 13679913.
11.^ Kump, L. R.; Pavlov, A.; Arthur, M. A. (2005). "Massive release of hydrogen sulfide to the
surface ocean and atmosphere during intervals of oceanic anoxia". Geology 33: 397.
doi:10.1130/G21295.1. edit
12.^ Butterfield, N. J. (Jan 2009). "Oxygen, animals and oceanic ventilation: an alternative view".
Geobiology 7 (1): 1–7. doi:10.1111/j.1472-4669.2009.00188.x. ISSN 1472-4677. PMID 19200141.
edit
13.^ Steemans, P.; Hérissé, L.; Melvin, J.; Miller, A.; Paris, F.; Verniers, J.; Wellman, H. (Apr
2009). "Origin and Radiation of the Earliest Vascular Land Plants". Science 324 (5925): 353.
doi:10.1126/science.1169659. ISSN 0036-8075. PMID 19372423. edit
14.^ Greb, S. F.; Rothwell, G.W. (2006). "Wetlands before tracheophytes: Thalloid terrestrial
communities of the Early Silurian Passage Creek biota (Virginia)" (Google books). Wetlands Through
Time. doi:10.1130/2006.2399(02). ISBN 9780813723990. http://books.google.com/?
id=A1p8FQ0VnIUC&pg=PA41&dq=tomescu+rothwell. Retrieved 2008-05-06. edit
15.^ Scott, C.; Glasspool, J. (Jul 2006). "The diversification of Paleozoic fire systems and
fluctuations in atmospheric oxygen concentration" (Free full text). Proceedings of the National
Academy of Sciences of the United States of America 103 (29): 10861–10865.
doi:10.1073/pnas.0604090103. ISSN 0027-8424. PMID 16832054. PMC 1544139.
http://www.pnas.org/cgi/pmidlookup?view=long&pmid=16832054. edit
16.^ a b c d Stewart, W.N. and Rothwell, G.W. 1993. Paleobotany and the evolution of plants,
Second edition. Cambridge University Press, Cambridge, UK. ISBN 0-521-38294-7
 17.^ a b c d C. Kevin Boyce (2008). "How green was Cooksonia? The importance of size in
understanding the early evolution of physiology in the vascular plant lineage". Paleobiology 34: 179.
doi:10.1666/0094-8373(2008)034[0179:HGWCTI]2.0.CO;2.
18.^ a b c d e f g h i j k l m n o p q r s t u v w x y z aa ab ac ad ae af ag ah Sperry, J.S. (2003). "Evolution
of Water Transport and Xylem Structure". International Journal of Plant Sciences 164 (S3): 115–127.
doi:10.1086/36839.
19.^ Niklas, K.J.; Smocovitis, V. (1983). "Evidence for a conducting strand in Early Silurian
(Llandoverian) plants; implications for the evolution of the land plants". Paleobiology 9 (2): 126–137.
http://links.jstor.org/sici?sici=0094-8373(198321)9%3A2%3C126%3AEFACSI%3E2.0.CO%3B2-6.
Retrieved 2008-04-26.
20.^ a b c d e f g h Niklas, K.J. (1985). "The evolution of tracheid diameter in early vascular plants
and its implications on the hydraulic conductance of the primary xylem strand". Evolution 39 (5):
1110–1122. doi:10.2307/2408738. http://links.jstor.org/sici?sici=0014-
3820(198509)39%3A5%3C1110%3ATEOTDI%3E2.0.CO%3B2-A. Retrieved 2008-04-26.
21.^ Niklas, K.J.; Pratt, L.M. (1980). "Evidence for Lignin-Like Constituents in Early Silurian
(Llandoverian) Plant Fossils". Science 209 (4454): 396–397. doi:10.1126/science.209.4454.396.
PMID 17747811.
22.^ Qiu, Y.L.; Li, L.; Wang, B.; Chen, Z.; Knoop, V.; Groth-malonek, M.; Dombrovska, O.; Lee,
J.; Kent, L.; Rest, J.; Others, (2006). "The deepest divergences in land plants inferred from
phylogenomic evidence". Proceedings of the National Academy of Sciences 103 (42): 15511.
doi:10.1073/pnas.0603335103. PMID 17030812.
23.^ a b c d e f g h i Stewart, W.N.; Rothwell, G.W. (1993). Paleobiology and the evolution of
plants. Cambridge University Press. pp. 521pp.
24.^ Zosterophyllophytes
 25.^ Rickards, R.B. (2000). "The age of the earliest club mosses: the Silurian Baragwanathia
flora in Victoria, Australia" (abstract). Geological Magazine 137 (2): 207–209.
doi:10.1017/S0016756800003800.
http://geolmag.geoscienceworld.org/cgi/content/abstract/137/2/207. Retrieved 2007-10-25.
26.^ a b c d e Kaplan, D.R. (2001). "The Science of Plant Morphology: Definition, History, and
Role in Modern Biology". American Journal of Botany 88 (10): 1711–1741. doi:10.2307/3558347.
http://links.jstor.org/sici?sici=0002-9122(200110)88%3A10%3C1711%3ATSOPMD%3E2.0.CO
%3B2-T. Retrieved 2008-01-31.
27.^ Taylor, T.N.; Hass, H.; Kerp, H.; Krings, M.; Hanlin, R.T. (2005). "Perithecial ascomycetes
from the 400 million year old Rhynie chert: an example of ancestral polymorphism" (abstract).
Mycologia 97 (1): 269–285. doi:10.3852/mycologia.97.1.269. PMID 16389979.
http://www.mycologia.org/cgi/content/abstract/97/1/269. Retrieved 2008-04-07.
28.^ a b c Boyce, C.K.; Knoll, A.H. (2002). "Evolution of developmental potential and the multiple
independent origins of leaves in Paleozoic vascular plants". Paleobiology 28 (1): 70–100.
doi:10.1666/0094-8373(2002)028<0070:EODPAT>2.0.CO;2.
29.^ Hao, S.; Beck, C.B.; Deming, W. (2003). "Structure of the Earliest Leaves: Adaptations to
High Concentrations of Atmospheric CO2". International Journal of Plant Sciences 164 (1): 71–75.
doi:10.1086/344557.
30.^ Berner, R.A.; Kothavala, Z. (2001). "Geocarb III: A Revised Model of Atmospheric CO 2 over
Phanerozoic Time" (abstract). American Journal of Science 301 (2): 182. doi:10.2475/ajs.301.2.182.
http://ajsonline.org/cgi/content/abstract/301/2/182. Retrieved 2008-04-07.
 31.^ Beerling, D.J.; Osborne, C.P.; Chaloner, W.G. (2001). "Evolution of leaf-form in land plants
linked to atmospheric CO2 decline in the Late Palaeozoic era". Nature 410 (6826): 287–394.
doi:10.1038/35066546. PMID 11268207.
32.^ Taylor, T.N.; Taylor, E.L. (1993). The biology and evolution of fossil plants.
33.^ Shellito, C.J.; Sloan, L.C. (2006). "Reconstructing a lost Eocene paradise: Part I.
Simulating the change in global floral distribution at the initial Eocene thermal maximum". Global and
Planetary Change 50 (1-2): 1–17. doi:10.1016/j.gloplacha.2005.08.001.
http://linkinghub.elsevier.com/retrieve/pii/S0921818105001475. Retrieved 2008-04-08.
34.^ Aerts, R. (1995). "The advantages of being evergreen". Trends in Ecology & Evolution 10
(10): 402–407. doi:10.1016/S0169-5347(00)89156-9.
35.^ Labandeira, C.C.; Dilcher, D.L.; Davis, D.R.; Wagner, D.L. (1994). "Ninety-seven million
years of angiosperm-insect association: paleobiological insights into the meaning of coevolution".
Proceedings of the National Academy of Sciences of the United States of America 91 (25): 12278–
12282. doi:10.1073/pnas.91.25.12278. PMID 11607501.
36.^ Boyce, K.C.; Hotton, C.L.; Fogel, M.L.; Cody, G.D.; Hazen, R.M.; Knoll, A.H.; Hueber, F.M.
(May 2007). "Devonian landscape heterogeneity recorded by a giant fungus" (PDF). Geology 35 (5):
399–402. doi:10.1130/G23384A.1. http://geology.geoscienceworld.org/cgi/reprint/35/5/399.pdf.
37.^ Stein, W.E.; Mannolini, F.; Hernick, L.V.; Landing, E.; Berry, C.M. (2007). "Giant
cladoxylopsid trees resolve the enigma of the Earth's earliest forest stumps at Gilboa.". Nature 446
(7138): 904–7. doi:10.1038/nature05705. PMID 17443185.
38.^ Retallack, G.J.; Catt, J.A.; Chaloner, W.G. (1985). "Fossil Soils as Grounds for Interpreting
the Advent of Large Plants and Animals on Land [and Discussion"]. Philosophical Transactions of the
Royal Society of London. Series B, Biological Sciences 309 (1138): 105–142.
doi:10.1098/rstb.1985.0074. http://links.jstor.org/sici?sici=0080-
4622(19850402)309%3A1138%3C105%3AFSAGFI%3E2.0.CO%3B2-5. Retrieved 2008-04-07.
39.^ Dannenhoffer, J.M.; Bonamo, P.M. (1989). "Rellimia thomsonii from the Givetian of New
York: Secondary Growth in Three Orders of Branching". American Journal of Botany 76 (9): 1312–
1325. doi:10.2307/2444557. http://links.jstor.org/sici?sici=0002-
9122(198909)76:9%3C1312:RTFTGO%3E2.0.CO;2-S. Retrieved 2008-04-07.
40.^ Davis, P; Kenrick, P. (2004). Fossil Plants. Smithsonian Books, Washington D.C..
41.^ Donoghue, M.J. (2005). "Key innovations, convergence, and success: macroevolutionary
lessons from plant phylogeny" (abstract). Paleobiology 31 (2): 77–93. doi:10.1666/0094-
8373(2005)031[0077:KICASM]2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/content/abstract/31/2_Suppl/77. Retrieved 2008-04-07.
42.^ a b c Bowe, L.M.; Coat, G.; Depamphilis, C.W. (2000). "Phylogeny of seed plants based on
all three genomic compartments: Extant gymnosperms are monophyletic and Gnetales' closest
relatives are conifers". Proceedings of the National Academy of Sciences 97 (8): 4092.
doi:10.1073/pnas.97.8.4092.
43.^ a b c Chaw, S.M.; Parkinson, C.L.; Cheng, Y.; Vincent, T.M.; Palmer, J.D. (2000). "Seed
plant phylogeny inferred from all three plant genomes: Monophyly of extant gymnosperms and origin
of Gnetales from conifers". Proceedings of the National Academy of Sciences 97 (8): 4086.
doi:10.1073/pnas.97.8.4086.
44.^ a b c Soltis, D.E.; Soltis, P.S.; Zanis, M.J. (2002). "Phylogeny of seed plants based on
evidence from eight genes" (abstract). American Journal of Botany 89 (10): 1670.
doi:10.3732/ajb.89.10.1670. http://amjbot.org/cgi/content/abstract/89/10/1670. Retrieved 2008-04-
08.
 45.^ Friis, E.M.; Pedersen, K.R.; Crane, P.R. (2006). "Cretaceous angiosperm flowers:
Innovation and evolution in plant reproduction". Palaeogeography, Palaeoclimatology,
Palaeoecology 232 (2-4): 251–293. doi:10.1016/j.palaeo.2005.07.006.
46.^ Hilton, J.; Bateman, R.M. (2006). "Pteridosperms are the backbone of seed-plant
phylogeny". The Journal of the Torrey Botanical Society 133 (1): 119–168. doi:10.3159/1095-
5674(2006)133[119:PATBOS]2.0.CO;2.
47.^ a b Bateman, R.M.; Hilton, J.; Rudall, P.J. (2006). "Morphological and molecular
phylogenetic context of the angiosperms: contrasting the 'top-down' and 'bottom-up' approaches
used to infer the likely characteristics of the first flowers". Journal of Experimental Botany 57 (13):
3471. doi:10.1093/jxb/erl128. PMID 17056677.
48.^ a b c d Frohlich, M.W.; Chase, M.W. (2007). "After a dozen years of progress the origin of
angiosperms is still a great mystery.". Nature 450 (7173): 1184–9. doi:10.1038/nature06393.
PMID 18097399.
49.^ Mora, C.I.; Driese, S.G.; Colarusso, L.A. (1996). "Middle to Late Paleozoic Atmospheric
CO2 Levels from Soil Carbonate and Organic Matter". Science 271 (5252): 1105–1107.
doi:10.1126/science.271.5252.1105.
50.^ Berner, R.A. (1994). "GEOCARB II: A revised model of atmospheric CO 2 over Phanerozoic
time". Am. J. Sci 294: 56–91.
51.^ a b Algeo, T.J.; Berner, R.A.; Maynard, J.B.; Scheckler, S.E.; Archives, G.S.A.T. (1995).
"Late Devonian Oceanic Anoxic Events and Biotic Crises: "Rooted" in the Evolution of Vascular Land
Plants?" ([dead link]). GSA Today 5 (3). https://rock.geosociety.org/pubs/gsatoday/gsat9503.htm.
52.^ Retallack, G. J. (1986). Wright, V. P.. ed. Paleosols: their Recognition and Interpretation .
Oxford: Blackwell.
 53.^ a b c d e f g h i Algeo, T.J.; Scheckler, S.E. (1998). "Terrestrial-marine teleconnections in the
Devonian: links between the evolution of land plants, weathering processes, and marine anoxic
events". Philosophical Transactions of the Royal Society B: Biological Sciences 353 (1365): 113–130.
doi:10.1098/rstb.1998.0195.
54.^ a b Kenrick, P.; Crane, P.R. (1997). "The origin and early evolution of plants on land".
Nature 389 (6646): 33. doi:10.1038/37918.
55.^ Schüßler, A. et al. (2001). "A new fungal phlyum, the Glomeromycota: phylogeny and
evolution.". Mycol. Res. 105 (12): 1416. doi:10.1017/S0953756201005196.
http://journals.cambridge.org/action/displayAbstract?fromPage=online&aid=95091.
56.^ Simon, L., Bousquet, J., Levesque, C., Lalonde, M. (1993). "Origin and diversification of
endomycorrhizal fungi and coincidence with vascular land plants". Nature 363: 67–69.
doi:10.1038/363067a0.
57.^ Remy, W., Taylor, T., Hass, H., Kerp, H. (1994). "Four hundred-million-year-old vesicular
arbuscular mycorrhizae". Proceedings of the National Academy of Science USA 91: 11841–11843.
doi:10.1073/pnas.91.25.11841.
58.^ * Brundrett, M.C. (2002). "Coevolution of roots and mycorrhizas of land plants". New
Phytologist 154 (2): 275–304. doi:10.1046/j.1469-8137.2002.00397.x.
59.^ a b c Mapes, G.; Rothwell, G.W.; Haworth, M.T. (1989). "Evolution of seed dormancy".
Nature 337 (6208): 645–646. doi:10.1038/337645a0.
60.^ Nam, J.; Depamphilis, CW; Ma, H; Nei, M (2003). "Antiquity and Evolution of the MADS-
Box Gene Family Controlling Flower Development in Plants". Mol. Biol. Evol. 20 (9): 1435–1447.
doi:10.1093/molbev/msg152. PMID 12777513.
http://mbe.oxfordjournals.org/cgi/content/full/20/9/1435.
 61.^ a b Crepet, W. L. (2000). "Progress in understanding angiosperm history, success, and
relationships: Darwin's abominably "perplexing phenomenon"". Proceedings of the National
Academy of Sciences 97: 12939. doi:10.1073/pnas.97.24.12939. PMID 11087846.
http://www.pnas.org/cgi/reprint/97/24/12939.
62.^ Sun, G.; Ji, Q.; Dilcher, D.L.; Zheng, S.; Nixon, K.C.; Wang, X. (2002). "Archaefructaceae,
a New Basal Angiosperm Family". Science 296 (5569): 899. doi:10.1126/science.1069439.
PMID 11988572.
63.^ In fact, Archaeofructus probably didn't bear true flowers: see
• Friis, E.M.; Doyle, J.A.; Endress, P.K.; Leng, Q. (2003). "Archaefructus--angiosperm
precursor or specialized early angiosperm?". Trends in Plant Science 8 (8): 369–373.
doi:10.1016/S1360-1385(03)00161-4. PMID 12927969.
64.^ a b Wing, S.L.; Boucher, L.D. (1998). "Ecological Aspects Of The Cretaceous Flowering
Plant Radiation". Annual Reviews in Earth and Planetary Sciences 26 (1): 379–421.
doi:10.1146/annurev.earth.26.1.379.
65.^ Wilson Nichols Stewart & Gar W. Rothwell, Paleobotany and the evolution of plants, 2nd
ed., Cambridge Univ. Press 1993, p. 498
66.^ a b Feild, T.S.; Arens, N.C.; Doyle, J.A.; Dawson, T.E.; Donoghue, M.J. (2004). "Dark and
disturbed: a new image of early angiosperm ecology" (abstract). Paleobiology 30 (1): 82–107.
doi:10.1666/0094-8373(2004)030<0082:DADANI>2.0.CO;2.
http://paleobiol.geoscienceworld.org/cgi/content/abstract/30/1/82. Retrieved 2008-04-08.
67.^ a b c d Osborne, C.P.; Beerling, D.J. (2006). "Review. Nature's green revolution: the
remarkable evolutionary rise of C4 plants". Philosophical Transactions: Biological Sciences 361
(1465): 173–194. doi:10.1098/rstb.2005.1737. PMID 16553316. PMC 1626541.
http://www.journals.royalsoc.ac.uk/index/YTH8204514044972.pdf. Retrieved 2008-02-11.
68.^ a b Retallack, G. J. (1 August 1997). "Neogene Expansion of the North American Prairie".
PALAIOS 12 (4): 380–390. doi:10.2307/3515337. ISSN 08831351. http://links.jstor.org/sici?
sici=0883-1351(199708)12%3A4%3C380%3ANEOTNA%3E2.0.CO%3B2-Q. Retrieved 2008-02-11.
edit
69.^ Thomasson, J.R.; Nelson, M.E.; Zakrzewski, R.J. (1986). "A Fossil Grass (Gramineae:
Chloridoideae) from the Miocene with Kranz Anatomy". Science 233 (4766): 876–878.
doi:10.1126/science.233.4766.876. PMID 17752216.
70.^ O'Leary, Marion (May, 1988). "Carbon Isotopes in Photosynthesis". BioScience 38 (5): 328–
336. doi:10.2307/1310735. http://www.jstor.org/pss/1310735.
71.^ Osborne, P.; Freckleton, P. (Feb 2009). "Ecological selection pressures for C4
photosynthesis in the grasses". Proceedings. Biological sciences / the Royal Society 276 (1663):
1753. doi:10.1098/rspb.2008.1762. ISSN 0962-8452. PMID 19324795. edit
72.^ Bond, W.J.; Woodward, F.I.; Midgley, G.F. (2005). "The global distribution of ecosystems in
a world without fire". New Phytologist 165 (2): 525–538. doi:10.1111/j.1469-8137.2004.01252.x.
PMID 15720663.
73.^ Above 35% atmospheric oxygen, the spread of fire is unstoppable. Many models have
predicted higher values and had to be revised, because there was not a total extinction of plant life.
74.^ Pagani, M.; Zachos, J.C.; Freeman, K.H.; Tipple, B.; Bohaty, S. (2005). "Marked Decline in
Atmospheric Carbon Dioxide Concentrations During the Paleogene". Science 309 (5734): 600–603.
doi:10.1126/science.1110063. PMID 15961630.
75.^ Piperno, D.R.; Sues, H.D. (2005). "Dinosaurs Dined on Grass". Science 310 (5751): 1126.
doi:10.1126/science.1121020. PMID 16293745.
 76.^ Prasad, V.; Stroemberg, C.A.E.; Alimohammadian, H.; Sahni, A. (2005). "Dinosaur
Coprolites and the Early Evolution of Grasses and Grazers". Science(Washington) 310 (5751): 1177–
1180. doi:10.1126/science.1118806. PMID 16293759.
77.^ Keeley, J.E.; Rundel, P.W. (2005). "Fire and the Miocene expansion of C4 grasslands".
Ecology Letters 8 (7): 683–690. doi:10.1111/j.1461-0248.2005.00767.x.
78.^ a b Retallack, G.J. (2001). "Cenozoic Expansion of Grasslands and Climatic Cooling". The
Journal of Geology 109 (4): 407–426. doi:10.1086/320791.
79.^ a b c d McElwain, J.C.; Punyasena, S.W. (2007). "Mass extinction events and the plant
fossil record". Trends in Ecology & Evolution 22 (10): 548–557. doi:10.1016/j.tree.2007.09.003.
PMID 17919771.

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection
 Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_history_of_plants"

Categories: Plants | Prehistoric plants | Evolution by taxon

 W000

Histoire évolutive des végétaux

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Cet article est une ébauche concernant la flore.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
Les végétaux ont évolué à travers des niveaux croissants de complexité, depuis les premières
algues, puis les bryophytes, lycopodes, fougères, gymnospermes jusqu'au complexes angiospermes
d'aujourd'hui.
Alors que les végétaux plus primitifs continuent de prospérer, chaque nouveau degré d'organisation
développe de nouvelles capacités pour mieux s'adapter à son milieu. En outre, la plupart des analyses
cladistiques (bien que celles-ci soient souvent contradictoires) suggèrent que chaque groupe « plus
complexe » nait du plus complexe de son époque.
Tout porte à croire que les algues (pluricellulaires) sont apparues sur Terre il y a 1,2 milliard
d'années. Pendant l'Ordovicien, il y a près de 500 millions d'années, certaines formes d'algues, qui
produisaient de la chlorophylle a et de la chlorophylle b, donnèrent naissance aux algues vertes. C'est à
cette époque que les premières plantes terrestres apparurent. Celles-ci ont commencé à se diversifier à la fin
du Silurien, il y a environ 420 millions d'années, et les fruits de cette diversification sont visibles en détail
dans un fossile de début du Dévonien connu sous le nom de Flore de Rhynie.
Vers le milieu de la période du Dévonien, la plupart des caractéristiques connues des plantes
actuelles sont déjà présentes, y compris les racines, les feuilles et les graines.
À la fin du Dévonien, les plantes ont atteint un degré de sophistication qui leur a permis de former
des forêts de grands arbres.
L'innovation évolutionnaire continue après la période du Dévonien. La plupart des groupes de végétaux ont
été relativement épargnée par les extinctions Permo-Triassique, bien que les structures des communautés
aient changé. Ceci a contribué à l'évolution des plantes à fleur au Trias (il y a environ 200 millions d'années),
ce groupe s'étant fortement diversifié au Crétacé et au Tertiaire.
L'évolution la plus récente est celle des graminées, groupe qui est devenu important au milieu du
tertiaire, il y a environ 40 millions d'années. Les graminées, comme de nombreux autres groupes, ont créé
de nouveaux mécanismes de leur métabolisme pour survivre au faible taux de CO2, relativement aux ères
précédentes. Ainsi qu'aux conditions sèches et chaudes des tropiques au cours des 10 dernières millions
d'années.
 [Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives
 Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

[Dérouler]
v·d·m
Branches de la biologie
 Anatomie · Biochimie · Bio-informatique · Biologie cellulaire · Biologie de l'évolution · Biologie
humaine · Biologie marine · Biologie moléculaire · Botanique · Écologie · Exobiologie · Génétique ·
Génomique · Géobiologie · Microbiologie · Origine de la vie · Paléontologie · Parasitologie · Physiologie ·
Taxinomie · Virologie · Zoologie

• Portail de l’origine et de l’évolution du vivant

• Portail de la botanique
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_v%C3%A9g
%C3%A9taux ».

Catégories : Histoire évolutive | Plante | [+]

 W000

Evolution of sirenians
From Wikipedia, the free encyclopedia

Jump to: navigation, search

 A manatee's toenails. Manatee share common ancestry with Elephants.
Sirenia is the order of placental mammals which comprise modern "sea cows" (manatees and the
Dugong) and their extinct relatives. They are the only extant herbivorous marine mammals and the only
group of herbivorous mammals to have become completely aquatic. Sirenians are thought to have a 50-
million-year-old fossil record (early Eocene-recent). They attained modest diversity during the Oligocene and
Miocene, but have since declined as a result of climatic cooling, oceanographic changes, and human
interference. Two genera and four species are extant: Trichechus which includes the three species of
manatee that live along the Atlantic coasts and in rivers and coastlines of the Americas and western Africa,
and Dugong which is found in the Indian and Pacific oceans.
Contents
[hide]
• 1 Origins
• 2 Fossil history
• 3 See also
• 4 References
• 5 External links

[edit] Origins

Evolution of Sirenian Locomotion, based on Berta and Sumich, 1999.

Evolution of Sirenia, based on Daryl P. Domning and Encyclopedia of Marine Mammals.

Amazonian Manatee.

Dugong.
 West Indian Manatee.

Steller's Sea Cow (extinct).

Sirenians, along with Proboscidea (elephants), group together with the extinct Desmostylia and likely
the extinct Embrithopoda to form the Tethytheria. Tethytheria is thought to have evolved from primitive
hoofed mammals ("condylarths") along the shores of the ancient Tethys Ocean.
Tethytheria, combined with Hyracoidea (hyraxes) forms a clade called Paenungulata. Paenungulata
and Tethytheria (especially the latter) are among the least controversial mammalian clades, with strong
support from morphological and molecular interpretations. The ancestry of Sirenia is remote from that of
Cetacea and Pinnipedia, although they are thought to have evolved an aquatic lifestyle around the same
time.
[edit] Fossil history

Prorastomus, an early sirenian from the Eocene

The first appearance of sirenians in the fossil record was during the early Eocene, and by the late
Eocene, sirenians had significantly diversified. Inhabitants of rivers, estuaries, and nearshore marine waters,
they were able to spread rapidly. The most primitive sirenian known to date, Prorastomus, was found in
Jamaica, not the Old World.
The earliest known sea cows, of the families Prorastomidae and Protosirenidae, are both confined to
the Eocene, and were about the size of a pig, four legged amphibious creatures. By the time the Eocene
drew to a close, came the appearance of the Dugongidae; sirenians had acquired their familiar fully-aquatic
streamlined body with flipper-like front legs with no hind limbs, powerful tail with horizontal caudal fin, with up
and down movements which move them through the water, like cetaceans.
The last of the sirenian families who made their appearance, Trichechidae, apparently arose from
early dugongids in the late Eocene or early Oligocene. The current fossil record documents all major stages
in hindlimb and pelvic reduction to the extreme reduction in the modern manatee pelvis, providing an
example of dramatic morphological change among fossil vertebrates.
 Since sirenians first evolved, they have been herbivores, likely depending on seagrasses and aquatic
angiosperms (flowering plants) for food. To the present, almost all have remained tropical, marine and
angiosperm consumers. Sea cows are shallow divers with large lungs. They have heavy skeletons to help
them stay submerged; the bones are pachyostotic (swollen) and osteosclerotic (dense), especially the ribs
which are often found as fossils.
Eocene sirenians, like Mesozoic mammals but in contrast to other Cenozoic ones, have five instead
of four premolars, giving them a 3.1.5.3 dental formula. Whether this condition is truly a primitive retention in
sirenians is still under debate.
Although cheek teeth are relied on for identifying species in other mammals, they do not vary to a
significant degree among sirenians in their morphology, but are almost always low-crowned (brachyodont)
with two rows of large, rounded cusps (bunobilophodont). The most easily identifiable parts of sirenian
skeletons are the skull and mandible, especially the frontal and other skull bones. With the exception of a pair
of tusk-like first upper incisors present in most species, front teeth (incisors and canines) are lacking in all,
except the earliest sirenians.

[edit] See also

• Evolution of cetaceans

[edit] References
• Sirenian Evolution, Daryl P. Domning; Encyclopedia of Marine Mammals Perrin, Würsig and
Thewissen
 • Origin of Sirenians

[edit] External links

• Manatee and Elephant Fossils

[hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary Canalisation · Inversion · Modularity · Phenotypic plasticity

developmental
biology (Evo-devo)
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_sirenians"

 W000

Histoire évolutive des siréniens

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Les siréniens sont des mammifères marins qui, comme les cétacés, n'ont pas besoin de regagner la
terre ferme à une période quelconque de leur existence. Pourtant, l'origine de ces mammifères est
nécessairement terrestre. Comme plusieurs caractéristiques de leur anatomie rappellent celles des
éléphants, les scientifiques pensent que l'ancêtre herbivore des siréniens qui est devenu aquatique est
différent de celui des cétacés qui était par ailleurs carnivore.
Sommaire
[masquer]
• 1 L'origine
• 2 Analyse des fossiles
• 3 Genèse de cette théorie
• 4 Notes et références
• 5 Voir aussi
• 5.1 Article connexe
• 5.2 Lien externe

L'origine [modifier]
La séparation du groupe des siréniens avec les autres Tethytheria est plus ancienne que celle des
Cetacea et des Artiodactyla (10 Ma) ou de celle des Caniformia marins (2-4 Ma).
Prorastomus, le plus ancien des fossiles de siréniens trouvés, est un animal semi-aquatique qui
pouvait encore se déplacer sur la terre ferme et a été daté de 50 Ma.
Durant l'Éocène, le nombre de fossiles est assez diversifié. Ces animaux vivaient près des côtes, le long des
fleuves et les estuaires. Les Prorastomidae et Protosirenidae étaient de la grosseur d'un cochon avec des
pattes adaptées au milieu amphibie. Vers la fin de l'Éocène, d'autres siréniens ayant davantage l'allure de
Dugongidae sont apparus, leurs pattes avant étant devenues des nageoires et la queue transformée en
nageoire caudale horizontale. Comme chez les cétacés, les déplacements se faisaient par ondulation de
haut en bas. Les Trichechidae semblent les plus modernes des siréniens et issus des Dugongidae. On
dispose de nombreux fossiles qui correspondent à plusieurs étapes évolutives, de la réduction des membres
arrière à la disparition du pelvis.

Analyse des fossiles [modifier]

Bien que l'on compte les dents pour identifier de nombreuses espèces de mammifères, leur nombre
ne varie pas significativement parmi les siréniens. Les parties les plus facilement identifiables des squelettes
sont la mâchoire inférieure et le crâne, surtout les os crâniens frontaux. À l'exception d'une paire d'incisives
supérieures semblables à des défenses, les dents de devant (les incisives et les canines) manquent, sauf
chez les premiers siréniens.

Genèse de cette théorie [modifier]

Prorastomus a été découvert en Jamaïque.
Les analyses génétiques des années 1990 montrent que les Siréniens sont des Afrotheria[1], et sont
donc très éloignés des Perissodactyla et des Artiodactyla bien qu'ils aient été rangés avec les Proboscidea
dans les ongulés. Les siréniens n'ont donc pas évolué à partir de mammifères à sabots primitifs comme
Condylarths.
Cladogramme simplifié :
─o Afrotheria
└─o Paenungulata
 ├─o Hyracoidea
└─? Tethytheria
├─o Proboscidea
└─o Sirenia

Notes et références [modifier]

1. ↑ Frédéric Delsuc, Jean-François Mauffrey, Emmanuel Douzery, « Une nouvelle
classification des mammifères », dans La Science, vol. 303, Janvier 2003 [ texte intégral [archive] ]

Voir aussi [modifier]

Article connexe [modifier]
• Histoire évolutive des cétacés

Lien externe [modifier]

• Manatee and Elephant Fossils

[Enrouler]
v·d·m
Évolution biologique
 [Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

 Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire
évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

• Portail de la zoologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_sir
%C3%A9niens ».
Catégories : Sirénien | Histoire évolutive | [+]
 W000

Evolution of spiders
From Wikipedia, the free encyclopedia

Jump to: navigation, search

 A spider in Baltic amber
The evolution of spiders has been going on for at least 400 million years, since the first true spiders
(thin-waisted arachnids) evolved from crab-like chelicerate ancestors. Today, there are over 40,000
described spider species within the diverse phylum of arthropods.
Major developments in spider evolution include the development of spinnerets and silk secretion, as
well as different adaptions for its use.

[edit] Early spider-like arachnids

Among the oldest known land arthropods are Trigonotarbids, members an extinct order of spider-like
arachnids.
 Sharing many superficial characteristics with spiders, Trigonotarbida were terrestrial, respired
through book lungs, and walked on eight legs with two additional legs adapted to use around their mouth.
Arguments still remain open as to whether they possessed the ability to create silk. This had been popular
thought for quite some time, until an unpublished fossil was described with distinct microtubercles on its hind
legs, akin to those used by spiders to direct and manipulate their silk.
Regardless Trigonotarbida are not considered true-spiders. They are generally accepted as an
independent early offshoot within the Arachnida clade, and not directly ancestral to modern spiders.

[edit] Emergence of true spiders

At one stage the oldest fossil spider was believed to be Attercopus which lived 380 million years ago
during the Devonian. Attercopus was placed as the sister-taxon to all living spiders, but has now been
reinterpreted as a member of a separate, extinct order Uraraneida which could produce silk, but did not have
true spinnerets.
 Microscopic structure of spider silk, possibly the most important adaption developed during the
evolution of these arachnids
 The oldest true spiders are thus Carboniferous in age, or about 300 million years. Most of these early
segmented fossil spiders from the Coal Measures of Europe and North America probably belonged to the
Mesothelae, or something very similar, a group of primitive spiders with the spinnerets placed underneath the
middle of the abdomen, rather than at the end as in modern spiders. They were probably ground dwelling
predators, living in the giant clubmoss and fern forests of the mid-late Palaeozoic, where they were
presumably predators of other primitive arthropods. Silk may have been used simply as a protective covering
for the eggs, a lining for a retreat hole, and later perhaps for simple ground sheet web and trapdoor
construction.
As plant and insect life diversified so also did the spider's use of silk. Spiders with spinnerets at the
end of the abdomen (Mygalomorphae and Araneomorphae) appeared more than 250 million years ago,
presumably promoting the development of more elaborate sheet and maze webs for prey capture both on
ground and foliage, as well as the development of the safety dragline. The oldest mygalomorph,
Rosamygale, was described from the Triassic of France and belongs to the modern family Hexathelidae.
Megarachne servinei from the Permo-Carboniferous was once thought to be a giant mygalomorph spider
and, with its body length of 1 foot (34 cm) and leg span of above 20 inches (50 cm), the largest known spider
ever to have lived on Earth, but subsequent examination by an expert revealed that it was actually a
middling-sized sea scorpion.
By the Jurassic, the sophisticated aerial webs of the orb-weaver spiders had already developed to
take advantage of the rapidly diversifying groups of insects. A spider web preserved in amber, thought to be
110 million years old, shows evidence of a perfect "orb" web, the most famous, circular kind one thinks of
when imagining spider webs. An examination of the drift of those genes thought to be used to produce the
web-spinning behavior suggests that orb spinning was in an advanced state as many as 136 million years
ago.
 The 110-million-year-old amber-preserved web is also the oldest to show trapped insects, containing
a beetle, a mite, a wasp's leg, and a fly.[1] The ability to weave orb webs is thought to have been "lost", and
sometimes even re-evolved or evolved separately, in different breeds of spiders since its first appearance.
A complete list of fossil spiders, down to species level, can be found at the American Museum of
Natural History's website.[1]

[edit] See also

• Spider taxonomy
• Insect evolution

[edit] References
1. ^ "LiveScience.com - Oldest Known Spider Web Discovered in Amber".
http://www.livescience.com/animalworld/060622_spider_webs.html. Retrieved June 25, 2006.
Penney, D. 2008. Dominican Amber Spiders: a comparative neontological approach to identification
faunistics ecology and biogeography. Siri Scientific Press, Manchester. ISBN 978-0-9558636-0-8.

[edit] External links

• Picture of spider fossil
 [hide]
v•d•e
Spiders

Arachnology Ballooning · Behavior · Cannibalism ·

Evolution · Classification

Araneomorphae · Mesothelae ·
Taxonomy Mygalomorphae · Opisthothelae · List of
families of spiders · Lists of spider species

Arthropod leg · Book lung ·

Calamistrum · Cephalothorax · Chelicerae ·
Anatomy Cheliceral fang · Cribellum · Epigyne ·
Exuvia · Opisthosoma · Pedipalp · Scopulae ·
Silk · Spinneret · Urticating hair

Human Arachnophobia · Cultural depictions ·

interaction Spider bite · Spider fighting

Webs Spider web · Web decorations

 Category

[show]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_spiders"

Categories: Spiders | Evolution by taxon

 W000

Evolution of complexity
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Part of the Biology series on

Evolution
Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Selection
Speciation

Research and history

Evidence
History
Modern synthesis
Social effect / Objections

Evolutionary biology fields

Cladistics
Ecological genetics
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Evolution Portal · v • d • e
The evolution of complexity is an important outcome of the process of evolution. Evolution has
produced some remarkably complex organisms - although the actual level of complexity is very hard to
define or measure accurately in biology, with properties such as gene content, the number of cell types or
morphology all being used to assess an organism's complexity.[1][2] This observation that complex
organisms can be produced from simpler ones has led to the common misperception of evolution being
progressive and having a direction that leads towards what are viewed as "higher organisms".[3]
Nowadays, this idea of "progression" in evolution is regarded as misleading, with natural selection
having no intrinsic direction and organisms selected for either increased or decreased complexity in
response to local environmental conditions.[4] Although there has been an increase in the maximum level of
complexity over the history of life, there has always been a large majority of small and simple organisms and
the most common level of complexity (the mode) has remained constant.

Contents
[hide]
• 1 Selection for simplicity and complexity
• 2 Types of trends in complexity
• 3 History
• 4 See also
• 5 References

[edit] Selection for simplicity and complexity

Organisms that reproduce more quickly and plentifully than their competitors have an evolutionary
advantage. Consequently, organisms can evolve to become simpler and thus multiply faster and produce
more offspring, as they require fewer resources to reproduce. A good example are parasites such as malaria
and mycoplasma; these organisms often dispense with traits that are made unnecessary through parasitism
on a host.[5].
A lineage can also dispense with complexity when a particular complex trait merely provides no
selective advantage in a particular environment. Loss of this trait need not necessarily confer a selective
advantage, but may be lost by genetic drift if its loss does not confer an immediate selective disadvantage.
For example, a parasitic organism may dispense with the synthetic pathway of a metabolite where it can
readily scavenge that metabolite from its host. Discarding this synthesis may not necessarily allow the
parasite to conserve significant energy or resources and grow faster, but may be fixed in the population
through genetic drift if no disadvantage is incurred by loss of that pathway.
However, evolution can also produce more complex organisms. Complexity often arises in the co-
evolution of hosts and pathogens [6], with each side developing ever more sophisticated adaptations, such
as the immune system and the many techniques pathogens have developed to evade it. For example, the
parasite Trypanosoma brucei, which causes sleeping sickness, has evolved so many copies of its major
surface antigen that about 10% of its genome is devoted to different versions of this one gene. This
tremendous complexity allows the parasite to constantly change its surface and thus evade the immune
system through antigenic variation.[7]
More generally, the growth of complexity may be driven by the co-evolution between an organism
and the ecosystem of predators, prey and parasites to which it tries to stay adapted: as any of these become
more complex in order to cope better with the diversity of threats offered by the ecosystem formed by the
others, the others too will have to adapt by becoming more complex, thus triggering an on-going evolutionary
arms race[6] towards more complexity [8]. This trend may be reinforced by the fact that ecosystems
themselves tend to become more complex over time, as species diversity increases, together with the
linkages or dependencies between species.
[edit] Types of trends in complexity

Passive versus active trends in the evolution of complexity. Organisms at the beginning of the
processes are colored red. Numbers of organisms are shown by the height of the bars, with the graphs
moving up in a time series.
 If evolution possessed an active trend toward complexity, then we would expect to see an increase
over time in the most common value (the mode) of complexity among organisms, as shown to the right.[9]
Indeed, some computer models have suggested that the generation of complex organisms is an inescapable
feature of evolution.[10][11] This is sometimes referred to as evolutionary self-organization. Self-organization
is the spontaneous internal organization of a system. This process is accompanied by an increase in
systemic complexity, resulting in an emergent property that is distinctly different from any of the constituent
parts.
However, the idea of increasing production of complexity in evolution can also be explained through
a passive process.[9] As shown on the left, this involves an increase in variance but the mode does not
change. Thus, the maximum level of complexity increases over time, but only as an indirect product of there
being more organisms in total.
In this hypothesis, any appearance of evolution acting with an intrinsic direction towards increasingly-
complex organisms is a result of people concentrating on the small number of large, complex organisms that
inhabit the right-hand tail of the complexity distribution and ignoring simpler and much more common
organisms. This passive model predicts that the majority of species are microscopic prokaryotes, which is
supported by estimates of 10^6 to 10^9 extant prokaryotes [12] compared to diversity estimates of 10^6 to
3x10^6 for eukaryotes .[13][14] Consequently, in this view, microscopic life dominates Earth, and large
organisms only appear more diverse due to sampling bias.

[edit] History
In the 19th century, some scientists such as Jean-Baptiste Lamarck and Ray Lankester believed that
all Nature had an innate striving to become more complex with evolution. This belief may reflect then-current
ideas of Georg Hegel and Herbert Spencer that all creation was gradually evolving to a higher, more perfect
state.
According to this view, the evolution of parasites from an independent organism to parasite was seen
as "devolution" or "degeneration", and contrary to Nature. This view has sometimes been used
metaphorically by social theorists and propagandists to decry a class of people as "degenerate parasites".
Today, "devolution" is regarded as nonsense; rather, lineages will become simpler or more complicated
according to whatever forms have a selective advantage.[15]

[edit] See also

• Biocomplexity
• Biodiversity
• Biosphere
• Complex adaptive system
• Complexity
• Ecosystem
• Orthogenesis

[edit] References
1. ^ Adami C (2002). "What is complexity?". Bioessays 24 (12): 1085–94.
doi:10.1002/bies.10192. PMID 12447974.
2. ^ Waldrop M. et al. (2008). "Language: Disputed definitions". Nature 455 (7216): 1023–1028.
doi:10.1038/4551023a. PMID 18948925.
 3. ^ McShea D (1991). "Complexity and evolution: What everybody knows". Biology and
Philosophy 6 (3): 303–324. doi:10.1007/BF00132234.
4. ^ Ayala FJ (2007). "Darwin's greatest discovery: design without designer". Proc. Natl. Acad.
Sci. U.S.A. 104 Suppl 1: 8567–73. doi:10.1073/pnas.0701072104. PMID 17494753. PMC 1876431.
http://www.pnas.org/cgi/content/full/104/suppl_1/8567.
5. ^ Sirand-Pugnet P, Lartigue C, Marenda M, et al. (2007). "Being Pathogenic, Plastic, and
Sexual while Living with a Nearly Minimal Bacterial Genome". PLoS Genet. 3 (5): e75.
doi:10.1371/journal.pgen.0030075. PMID 17511520.
6. ^ a b R. Dawkins and J. R. Krebs (1979). "Arms Races between and within Species". Proc. R.
Soc. Lond.B 205 (1161): 489–511. doi:10.1098/rspb.1979.0081. PMID 42057.
7. ^ Pays E (2005). "Regulation of antigen gene expression in Trypanosoma brucei". Trends
Parasitol. 21 (11): 517–20. doi:10.1016/j.pt.2005.08.016. PMID 16126458.
8. ^ Heylighen, F. (1999a) "The Growth of Structural and Functional Complexity during
Evolution", in F. Heylighen, J. Bollen & A. Riegler (eds.) The Evolution of Complexity Kluwer
Academic, Dordrecht, 17-44.
9. ^ a b Carroll SB (2001). "Chance and necessity: the evolution of morphological complexity
and diversity". Nature 409 (6823): 1102–9. doi:10.1038/35059227. PMID 11234024.
10.^ Furusawa C, Kaneko K (2000). "Origin of complexity in multicellular organisms". Phys.
Rev. Lett. 84 (26 Pt 1): 6130–3. doi:10.1103/PhysRevLett.84.6130. PMID 10991141.
11.^ Adami C, Ofria C, Collier TC (2000). "Evolution of biological complexity". Proc. Natl. Acad.
Sci. U.S.A. 97 (9): 4463–8. doi:10.1073/pnas.97.9.4463. PMID 10781045. PMC 18257.
http://www.pnas.org/cgi/content/full/97/9/4463.
12.^ Oren A (2004). "Prokaryote diversity and taxonomy: current status and future challenges".
Philos. Trans. R. Soc. Lond., B, Biol. Sci. 359 (1444): 623–38. doi:10.1098/rstb.2003.1458.
 PMID 15253349. PMC 1693353. http://www.pubmedcentral.nih.gov/picrender.fcgi?
artid=1693353&blobtype=pdf.
13.^ May, R. M.; Beverton, R. J. H. (1990). "How Many Species?". R. M. May, Philos. Trans. R.
Soc. London Ser. B 330, 293 (1990) 330: 293. doi:10.1098/rstb.1990.0200.
http://rstb.royalsocietypublishing.org/content/330/1257/293.abstract?
ijkey=01b04eee73eda3609fb8f928b58db87298a825a0&keytype2=tf_ipsecsha.
14.^ Schloss P, Handelsman J (2004). "Status of the microbial census". Microbiol Mol Biol Rev
68 (4): 686–91. doi:10.1128/MMBR.68.4.686-691.2004. PMID 15590780. PMC 539005.
http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=15590780#r6.
15.^ Scientific American; Biology: Is the human race evolving or devolving? retrieved 2007-06-
11

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W000
Evolution of mammalian auditory ossicles
From Wikipedia, the free encyclopedia

Jump to: navigation, search

The evolution of mammalian auditory ossicles is one of the most well-documented[1] and important
evolutionary events, demonstrating both numerous transitional forms as well as an excellent example of
exaptation, the re-purposing of existing structures during evolution.
In reptiles, the eardrum is connected to the inner ear via a single bone, the stapes or stirrup, while
the upper and lower jaws contain several bones not found in mammals. Over the course of the evolution of
mammals, one lower and one upper jaw bone (the articular and quadrate) lost their purpose in the jaw joint
and were put to new use in the middle ear, connecting to the stapes and forming a chain of three bones
(collectively called the ossicles) which amplify sounds and allow more acute hearing. In mammals, these
three bones are known as the malleus, incus, and stapes (hammer, anvil, and stirrup respectively).
The evidence that the malleus and incus are homologous to the reptilian articular and quadrate was
originally embryological, and since this discovery an abundance of transitional fossils has both supported the
conclusion and given a detailed history of the transition.[2] The evolution of the stapes was an earlier and
distinct event.[3][4]
Contents
[hide]
• 1 Reichert–Gaupp theory
• 2 Definitive mammalian middle ear
• 3 Evolutionary history
• 3.1 Definition of "mammal"
• 3.2 Summary of the fossil evidence
• 3.2.1 Early tetrapod and amniote ears
• 3.2.2 Early therapsid jaws and ears
• 3.2.3 Twin-jointed jaws
• 3.2.4 Mammal-like jaws and ears
• 3.2.5 How these changes affected hearing
• 3.2.6 Natural selection
• 4 Summary
• 5 See also
• 6 References
• 7 Further reading
• 8 External links
[edit] Reichert–Gaupp theory
The relationship between the reptilian jaw bones and mammalian middle-ear bones was first
established on the basis of embryology and comparative anatomy by Reichert (in 1837, before the
publication of On the Origin of Species in 1859) and advanced by Gaupp, and this is known as the Reichert–
Gaupp Theory.
In the course of the development of the embryo, the incus and malleus arise from the same First
Pharyngeal arch as the Mandible and Maxilla, and are served by mandibular and maxillary division of the
Trigeminal Cranial nerve.[5]
...the discovery that the mammalian malleus and incus were actually homologues of visceral
elements of the "reptilian" jaw articulation ... ranks as one of the milestones in the history of
comparative biology.[6]

... it is one of the triumphs of the long series of researches on the extinct Theromorph reptiles,
begun by Owen (1845), and continued by Seeley, Broom, and Watson, to have revealed the
intermediate steps by which the change may have occurred from an inner quadrate to an outer
squamosal articulation ...[7]

There are also more recent studies in the genetic basis for the development of the ossicles from the
embryonic arch.[8] and relating this to evolutionary history.[9]
"Bapx1, also known as Nkx3.2, is the vertebrate homologue of the Drosophila gene Bagpipe. A
member of the NK2 class of homeobox genes ..."[10] and this gene is implicated in the change from the jaw
bones of non-mammals to the ossicles of mammals.[11] Yet the transition between the "reptilian" jaw and the
"mammalian" inner ear was not bridged in the fossil record until the 1950s[12] with the elaboration of such
fossils as the now-famous Morganucodon.[12]

A typical mammalian middle ear: sound makes the tympanum (ear-drum) vibrate; 3 small bones, the
malleus, incus and stapes, transmit the vibrations to the Labyrinth (inner ear), which transforms the vibrations
into nerve signals.
[edit] Definitive mammalian middle ear
The mammalian middle ear contains three tiny bones known as the ossicles: malleus, incus, and
stapes. The ossicles are a complex system of levers whose functions include: reducing the amplitude of the
vibrations; increasing the amount of energy transmitted. The details of these effects vary noticeably between
different mammal species, even when the species are as closely related as humans and chimpanzees.[13]

[edit] Evolutionary history

[edit] Definition of "mammal"
Living mammal species can be identified by the presence in females of mammary glands which
produce milk. Other features are required when classifying fossils, since mammary glands and other soft-
tissue features are not visible in fossils. Paleontologists therefore use a distinguishing feature that is shared
by all living mammals (including monotremes), but is not present in any of the early Triassic therapsids
("mammal-like reptiles"): mammals use two bones for hearing that all other amniotes use for eating. The
earliest amniotes had a jaw joint composed of the articular (a small bone at the back of the lower jaw) and the
quadrate (a small bone at the back of the upper jaw). All non-mammalian amniotes use this system including
lizards, crocodilians, dinosaurs (and their descendants the birds) and therapsids; so the only ossicle in their
middle ears is the stapes. But mammals have a different jaw joint, composed only of the dentary (the lower
jaw bone which carries the teeth) and the squamosal (another small skull bone). And in mammals the
quadrate and articular bones have become the incus and malleus bones in the middle ear.[14][15]
[edit] Summary of the fossil evidence
Here is a very simplified "family tree" of the various lineages involved:
--Tetrapods------
| ("4 legged"; the earliest breathed via gills)
|
+-- Amphibians ----------------------------------------------
|
`--------Reptiliomorphs-----
| ("reptile-like" amphibians)
|
`--Amniotes------
|
+--Sauropsids ("lizard faces")---------------
| (lizards, crocodilians, dinosaurs, birds
| Testudines; and some extinct groups)
|
`--Synapsids------
|
`--Pelycosaurs----
|
`--Therapsids-----
|
`--Mammals---------------
 The first fully terrestrial vertebrates were amniotes - their eggs had internal membranes which
allowed the developing embryo to breathe but kept water in. This allowed amniotes to lay eggs on dry land,
while amphibians generally need to lay their eggs in water. The first amniotes apparently arose in the late
Carboniferous from the ancestral reptiliomorphs (a group of amphibians whose only living descendants are
amniotes). Within a few million years two important amniote lineages became distinct: mammals' synapsid
ancestors and the sauropsids, from which lizards, snakes, crocodilians, dinosaurs and birds are descended.
[16]
The earliest known fossils of all these groups date from about 320 to 315M years ago. Unfortunately
it is difficult to be sure about when each of them evolved, since vertebrate fossils from the late Carboniferous
are very rare, and therefore the actual first occurrences of each of these types of animal might have been
considerably earlier.[17][15]
The pattern in most of the following sections is that each successive more "advanced" group started
with the more "primitive" jaws and ears of its predecessors, then developed more mammal-like jaws and
ears, and so on. The evolution of mammalian jaw joints and ears did not proceed neatly in step with the
evolution of other mammalian features; or, to put it another way, all but the last of the various stages into
which paleontologists divide the evolution towards the mammalian condition are not defined by their jaw
joints and ears.
 Mammalian and non-mammalian jaws. In the mammal configuration, the quadrate and articular
bones are much smaller and form part of the middle ear. Note that in mammals the lower jaw consists of only
the dentary bone.
 [edit] Early tetrapod and amniote ears
In modern amniotes (including mammals), the middle ear collects airborne sounds through an ear
drum and transmits the vibrations to the inner ear via thin cartilaginous and ossified structures, which usually
include the stapes (a stirrup-shaped auditory ossicle). But the earliest tetrapods, amphibians and amniotes
probably did not have ear drums. In fact ear drums apparently evolved independently three to six times, in:
stegocephalians (very primitive amphibians); in anurans (the amphibian group that includes frogs and toads);
in synapsids (mammals and their extinct relatives), in diapsids (the most important sauropsid group, including
lizards, crocodiles, dinosaurs and birds); perhaps separately in anapsids (turtles and their extinct relatives), if
turtles are not modified diapsids; probably in seymouriamorphs (a group of reptiliomorphs); and possibly in
some temnospondyls (primitive amphibians).[18][19] In all basal members of the 3 major clades of amniotes
(synapsids, eureptiles, and parareptiles) the stapes bones are relatively massive props that support the
braincase, and this function prevents them from being used as part of the hearing system. But there is
increasing evidence that synapsids, eureptiles and parareptiles developed eardrums connected to the inner
ear by stapes during the Permian.[20]

[edit] Early therapsid jaws and ears

The jaws of early synapsids, including the ancestors of mammals, were similar to those of other
tetrapods of the time, with a lower jaw consisting of a tooth-bearing dentary bone and several smaller
posterior bones. The jaw joint consisted of the articular bone in the lower jaw and the quadrate in the upper
jaw. The early pelycosaurs (late Carboniferous and early Permian) most probably did not have tympanic
membranes (external eardrums), and their massive stapes bones supported the braincase, with the lower
ends resting on the quadrates. But their descendants the therapsids (including mammals' ancestors)
probably did have tympanic membranes and these probably were in contact with the quadrate bones; and
the stapes bones were still in contact with the quadrates but functioned as auditory ossicles rather than
braincase supports; so the therapsids' quadrates had a dual function, as part of the jaw joint and as parts of
the hearing system.[21][22]

[edit] Twin-jointed jaws

Morganucodontidae and other transitional forms had both types of jaw joint: dentary-squamosal
(front) and articular-quadrate (rear).
During the Permian and early Triassic the dentary of therapsids, including the ancestors of
mammals, continually enlarged while other jaw bones were reduced.[23][23] Eventually, the dentary was
able to make contact with the squamosal, a bone in the upper jaw located anterior to the quadrate, allowing
two simultaneous jaw joints[24] - an anterior "mammalian" joint between the dentary and squamosal and a
posterior "reptilian" joint between the quadrate and articular. This "twin-jointed jaw" can be seen in late
cynodonts and early mammaliforms[25]. Morganucodon is one of the first discovered and most thoroughly
studied of the mammaliforms, since an unusually large number of morganucodont fossils have been found,
and
Morganucodon is an almost perfect intermediate in this respect (the "twin-jointed jaw") between
the higher mammal-like reptiles on the one hand and the typical mammals on the other.[26]

(note: "mammal-like reptiles" is an obsolete term for the therapsids)

[edit] Mammal-like jaws and ears

As the dentary continued to enlarge during the Triassic, the older quadrate-articular joint fell out of
use. Some of the bones were lost, but the quadrate (which is directly connected to the stapes), the articular
(connected to the quadrate) and the angular (connected to the articular) became free-floating and associated
with the stapes. This occurred at least twice in the mammaliformes ("almost-mammals"). The
Multituberculates, which lived from about 160M years ago (mid-Jurassic) to about 35M years ago (early
Oligocene) had jaw joints that consisted of only the dentary and squamosal bones, and the quadrate and
articular bones were part of the middle ear; but other features of their teeth, jaws and skulls are significantly
different from those of mammals.[27][15]
In the lineage most closely related to mammals, the jaws of Hadrocodium (about 195M years ago in
the very early Jurassic) suggest that it or a very close ancestor may have been the first to have a nearly fully
mammalian middle ear: it lacks the trough at the rear of the lower jaw, over which the eardrum stretched in
therapsids and earlier mammaliformes, and the absence of this trough which suggests that Hadrocodium’s
ear was part of the cranium, as it is in mammals, and hence that the former articular and quadrate had
migrated to the middle ear and become the malleus and incus; but Hadrocodium’s dentary has a "bay" at the
rear which mammals lack, a hint that that its dentary bone retained the same shape that it would have had if
the articular and quadrate had remained part of the jaw joint.[27] It has been suggested that a relatively large
trough in the jaw bone of the early Cretaceous monotreme Teinolophos provides evidence of a pre-
mammalian jaw joint, because therapsids and many mammaliforms had such troughs, in which the articular
and angular bones "docked", and therefore that Teinolophos had a pre-mammalian middle ear; and therefore
that the mammalian middle ear ossicles evolved indepedendently in monotremes and in other mammals.[28]
But a more recent analysis of Teinolophos concluded that the animal was a full-fledged platypus and the
trough was a channel for the large number of nerves that collect signals from the electrical and vibration
sensors in the bill (this is a signature feature of the platypi within monotremes), and therefore that the trough
is not evidence that Teinolophos had a pre-mammalian jaw joint and a pre-mammalian middle ear.[29]
Ironically Rich and Vickers-Rich were among the authors of the 2005 paper on which they later cast doubt.
A recently discovered intermediate form is the primitive mammal Yanoconodon, from 125 million
years ago in the Mesozoic, in which the ossicles have separated from the jaw and serve the hearing function
in the middle ear, yet maintain a slender connection to the jaw via the ossified Meckel's cartilage, which in
more advanced mammals dissolves during development.[30]

[edit] How these changes affected hearing

The frequency range and sensitivity of the ear is dependent upon the shape and arrangement of the
middle-ear bones. In early synapsids such as the pelycosaurs, the quadrate and articular had to function as
the jaw joint, and this severely limited how far these bones could be modified to alter the frequency range of
the ear. But once these bones were no longer involved in the jaw joint, variations which affected hearing
would not also affect jaw joint function, and this allowed unconstrained evolution of the mammalian hearing
apparatus.[31] By the Jurassic, the typical mammalian ear had evolved, in which the angular had become the
tympanic annula (a bony support for the tympanic membrane), while the articular and quadrate had become
the malleus and incus, respectively, connected in series with the stapes. This series of three bones acts as
an amplification system to allow enhanced hearing.[32]
The transition between these two states is one of the most well-documented[33] and supported in all
of evolution, and newly discovered fossils from this transitional period have recently improved our
understanding of this transition. But they also suggest that it was not a simple linear process from the early
therapsid jaw (quadrate-articular joint) and middle ear (with stapes as the only ossicles) to the modern
mammalian condition.[22]

[edit] Natural selection

It has been suggested that natural selection could be a factor in the preservation of the structure of
the middle ear in mammals.[1][22] Many of the earliest mammals were quite small, and the dentition
indicates that they were insectivorous. If they were "warm-blooded" (homeothermous), like modern
mammals, then they could have been nocturnal. This fits with the popular image of small, nocturnal
insectivorous mammals surviving in niches not accessible to the large, dominant contemporary dinosaurs.
The enhanced hearing, particularly in the higher frequencies, would be helpful for nocturnal animals, in
particular for detecting insects. This scenario is consistent with selective advantage being a contributory
factor to the transition.

[edit] Summary
While the stapes is present in many types of tetrapods, the addition of the incus and malleus (also
known as quadrate and articular) in the middle ear is a signature feature of mammals, distinguishing them
from reptiles and all other vertebrates. They therefore have the appearance of representing a discontinuity in
the tree of life. But in the early 19th century, it was hypothesized that these bones are not a total novelty, but
are the equivalents of two bones which non-mammals have in their jaws. This hypothesis made sense, not
only of the existence of these middle-ear bones, but also of certain other features of the anatomy, such as
the paths taken by nerves in the head.
As evolutionary biology began to be expanded upon, this relationship became treated as one of
common descent. For the evolutionary explanation to make sense, it seemed to demand that there would be
a transition in function between being part of the feeding mechanism in the joint of the jaw and serving only in
hearing; and this would mean that somehow there had to be an intermediate connecting these two quite
different functions. With the discovery of Morganucodon and other fossils, there were concrete examples of
this. There was a double jaw joint: the "older reptilian", as well as the "newer mammalian", in the same
animal. This meant a confirmation of the pattern of inference from comparative anatomy to evolutionary
biology.
The earliest mammals were generally small animals, probably nocturnal insectivores. This suggests
a plausible evolutionary mechanism driving the change, for with these small bones in the middle ear, a
mammal has extended its range of hearing for higher-pitched sounds which would improve the detection of
insects in the dark.[34] Natural selection would account for the success of this feature. And still one more
connection with another part of biology: genetics suggested a mechanism for this transition, the kind of major
change of function seen elsewhere in the world of life being studied by Evodevo.

[edit] See also

• Evolution of mammals
 • Morganucodon
• Maotherium asiaticus
• Oligokyphus
• Probainognathus
• Transitional fossil
• Yanoconodon

[edit] References
1. ^ a b Allin EF (December 1975). "Evolution of the mammalian middle ear". J. Morphol. 147
(4): 403–37. doi:10.1002/jmor.1051470404. PMID 1202224.
2. ^ Bowler, Peter J. (1996). "Chapter 6: The Origin of Birds and Mammals". Life's splendid
drama: evolutionary biology and the reconstruction of life's ancestry, 1860-1940 . Chicago: University
of Chicago Press. ISBN 0-226-06921-4.
3. ^ Janvier, Philippe (2002). Early vertebrates. Oxford Monographs on Geology and
Geophysics, 33. Oxford: Clarendon Press. pp. page 56. ISBN 978-0198526469.
4. ^ The Shoulder Bone’s Connected to the Ear Bone… Carl Zimmer blog The Loom at Discover
magazine for 2008 October 15.
5. ^ Gilbert, Scott F. (2003). Developmental biology (7th ed.). Sunderland, Mass: Sinauer
Associates. pp. page 435. ISBN 0-87893-258-5.
6. ^ Novacek MJ (1993). Hall BK, Hanken J. ed. The Skull. Chicago: University of Chicago
Press. pp. pages 438–545. ISBN 0-226-31568-1. "Novacek references these early works: Johann
Friedrich Meckel Handbuch der Menschlichen Anatomie, Halle, 1820; Reichert "Ueber die
Visceralbogen der Wirbelthiere im Allegemeinen und deren Metamorphosen bei den Vögln und
Säugethieren", Archiv für Anatomie, Physiologie, und wissenschaftliche Medizin , Leipzig 1837,
pages 120-122; and Gaupp "Die Reichertsche Theorie (Hammer-, Amboss- und Kieferfrage)" Archiv
für Anatomie und Entwicklungsgeschichte 1913, pages 1-416."
7. ^ Goodrich ES (1958). Studies on the Structure and Development of Vertebrates . Dover.
pp. page 474.
8. ^ Mallo M (March 2001). "Formation of the middle ear: recent progress on the developmental
and molecular mechanisms". Dev. Biol. 231 (2): 410–9. doi:10.1006/dbio.2001.0154.
PMID 11237469.
9. ^ Raff RA (December 2007). "Written in stone: fossils, genes and evo-devo". Nat. Rev.
Genet. 8 (12): 911–20. doi:10.1038/nrg2225. PMID 18007648.
10.^ Wilson J, Tucker AS (February 2004). "Fgf and Bmp signals repress the expression of
Bapx1 in the mandibular mesenchyme and control the position of the developing jaw joint". Dev. Biol.
266 (1): 138–50. doi:10.1016/j.ydbio.2003.10.012. PMID 14729484.
11.^ Tucker AS, Watson RP, Lettice LA, Yamada G, Hill RE (March 2004). "Bapx1 regulates
patterning in the middle ear: altered regulatory role in the transition from the proximal jaw during
vertebrate evolution". Development 131 (6): 1235–45. doi:10.1242/dev.01017. PMID 14973294.
12.^ a b AW Crompton and FA Jenkins, Jr (1973). "Mammals from Reptiles: A Review of
Mammalian Origins". Annual Review of Earth and Planetary Sciences 1: 131–155.
doi:10.1146/annurev.ea.01.050173.001023.
13.^ Masali M (October 1992). "The ear ossicles and the evolution of the primate ear: A
biomechanical approach". Human Evolution (Springer Netherlands) 7 (4): 1–5.
doi:10.1007/BF02436407.
 14.^ White T. "Unit 430: Mammalia: Overview". PALAEOS: The Trace of Life on Earth.
palaeos.com. http://www.palaeos.com/Vertebrates/Units/430Mammalia/430.000.html. Retrieved
2008-07-21.
15.^ a b c Cowen, Richard (2000). History of life. Oxford: Blackwell Science. pp. page 432.
ISBN 0-632-04444-6.
16.^ White T. "Amniota". PALAEOS: The Trace of Life on Earth. palaeos.com.
http://www.palaeos.org/Amniota. Retrieved 2008-07-21.
17.^ White T. "Synapsida: Varanopseidae". PALAEOS: The Trace of Life on Earth.
palaeos.com. http://www.palaeos.com/Vertebrates/Units/Unit390/200.html. Retrieved 2008-07-21.
18.^ Laurin M (January-March 1998). "The importance of global parsimony and historical bias in
understanding tetrapod evolution. Part I. Systematics, middle ear evolution and jaw suspension".
Annales des Sciences Naturelles - Zoologie et Biologie Animale 19 (1): 1–42. doi:10.1016/S0003-
4339(98)80132-9.
19.^ Laurin M. "Hearing in Stegocephalians". Tree of Life. Tree of Life Project.
http://www.tolweb.org/articles/?article_id=470. Retrieved 2008-07-21.
20.^ Müller J, Tsuji LA (2007). "Impedance-matching hearing in Paleozoic reptiles: evidence of
advanced sensory perception at an early stage of amniote evolution". PLoS ONE 2 (9): e889.
doi:10.1371/journal.pone.0000889. PMID 17849018.
21.^ Fay, Richard R.; Manley, Geoffrey A.; Popper, Arthur N. (2004). Evolution of the vertebrate
auditory system. Berlin: Springer. ISBN 0-387-21089-X.
22.^ a b c Luo ZX (December 2007). "Transformation and diversification in early mammal
evolution". Nature 450 (7172): 1011–9. doi:10.1038/nature06277. PMID 18075580.
23.^ a b Sidor CA (July 2001). "Simplification as a trend in synapsid cranial evolution". Evolution
55 (7): 1419–42. PMID 11525465.
 24.^ Page 229, Michael J. Benton, Vertebrate Palaeontology: Biology and evolution , Unwin
Hyman, 1990 ISBN 0-04-566001-8
25.^ Page 228 of Edward H. Colbert and Michael Morales, Evolution of the Vertebrates: A
History of the Backboned Animals Through Time, Wiley-Liss, 4th edition, 1991 ISBN 0-471-85074-8
26.^ Kermack KA, Mussett F, Rigney HW (January 1981). "The skull of Morganucodon".
Zoological Journal of the Linnean Society 71 (1): page 147 of pages 1–158. doi:10.1111/j.1096-
3642.1981.tb01127.x.
27.^ a b White T. "Mammaliformes". PALAEOS: The Trace of Life on Earth. palaeos.com.
http://www.palaeos.com/Vertebrates/Units/Unit420/420.100.html. Retrieved 2008-07-21.
28.^ Rich TH, Hopson JA, Musser AM, Flannery TF, Vickers-Rich P (February 2005).
"Independent origins of middle ear bones in monotremes and therians". Science (journal) 307 (5711):
910–4. doi:10.1126/science.1105717. PMID 15705848.
29.^ Rowe T, Rich TH, Vickers-Rich P, Springer M, Woodburne MO (January 2008). "The oldest
platypus and its bearing on divergence timing of the platypus and echidna clades". Proc. Natl. Acad.
Sci. U.S.A. 105 (4): 1238–42. doi:10.1073/pnas.0706385105. PMID 18216270.
30.^ PZ Myers (March 16, 2007). "Yanoconodon, a transitional fossil". Pharyngula: Evolution,
development, and random biological ejaculations from a godless liberal .
http://scienceblogs.com/pharyngula/2007/03/yanoconodon_a_transitional_fos.php.
31.^ Lombard RE, Hetherington TE (1993). "Structural Basis of Hearing and Sound
Transmission". in Hall BK, Hanken J. The Skull (volume 3 ed.). Chicago: University of Chicago Press.
pp. pages 241–302. ISBN 0-226-31571-1.
32.^ Köppl C (11 August 2009). "Evolution of sound localization in land vertebrates". Current
Biology 19 (15): R635–R639. doi:10.1016/j.cub.2009.05.035. PMID 19674542.
 33.^ Cuffey CA. "References". The Fossil Record: Evolution or "Scientific Creation" . GCSSEPM
Foundation. http://www.gcssepm.org/special/cuffey_14.htm. Retrieved 2008-07-21.
34.^ Biello D (2007-03-14). "From Jaw to Ear: Transition Fossil Reveals Ear Evolution in Action".
Scientific American. http://www.sciam.com/article.cfm?id=fossil-reveals-ear-evolution-in-action.
Retrieved 2009-06-17. "Now hear this: early mammal fossil shows how sensitive ear bones evolved"

[edit] Further reading

• Allin EF, Hopson JA (1992). "Chapter 28: Evolution of the Auditory System in Synapsida
("Mammal-Like Reptiles" and Primitive Mammals) as Seen in the Fossil Record". in Popper AN,
Webster DB, Fay RR. The Evolutionary biology of hearing. Berlin: Springer-Verlag. pp. 587–614.
ISBN 0-387-97588-8.
• Rosowski JJ (1992). "Chapter 29: Hearing in Transitional Mammals: Predictions from the
Middle-Ear Anatomy and Hearing Capabilities of Extant Mammals". in Popper AN, Webster DB, Fay
RR. The Evolutionary biology of hearing. Berlin: Springer-Verlag. pp. 615–632. ISBN 0-387-97588-8.
• Gould SJ (1993). "Chapter 6: An Earful of Jaw". Eight Little Piggies: reflections in natural
history. New York: Norton. ISBN 0-393-03416-X.
• Hopson JA (January 1987). "The mammal-like reptiles: a study of transitional fossils". The
American Biology Teacher 49 (1): 16–26. http://www.jstor.org/stable/4448410.
• Zhe-XI L, Kielan-Jaworowska Z, Cifelli RL (2004). "Chapter 3: Origin of mammals".
Mammals from the age of dinosaurs: origins, evolution, and structure . New York: Columbia University
Press. ISBN 0-231-11918-6.
• Rougier GW, White JR (2006). "Chapter 6: Major Changes in the Ear Region and
Basicranium of Early Mammals". in Carrano MT, Gaudin TJ, Blob RW, Wible JR. Amniote
 paleobiology: perspectives on the evolution of mammals, birds, and reptiles: a volume honoring
James Allen Hopson. Chicago: University of Chicago Press. pp. 269–311. ISBN 0-226-09477-4.
• Shubin N (2008). "Chapter 10: Ears". Your inner fish: a journey into the 3. 5-billion-year
history of the human body. New York: Pantheon Books. ISBN 0-375-42447-4.

[edit] External links

• Theobald D (2004). "29+ Evidences for Macroevolution: Part 1, Example 2: reptile-
mammals". TalkOrigins.
http://www.talkorigins.org/faqs/comdesc/section1.html#morphological_intermediates_ex2. Retrieved
2009-06-17.
• Cuffey CA (2001). "The Fossil Record: Evolution or "Scientific Creation": Mammal-Like
Reptiles". GCSSEPM Foundation. http://www.gcssepm.org/special/cuffey_05.htm. Retrieved 2009-
06-17.
• Matzke N (2005). "The testimony of Kevin Padian in Kitzmiller v. Dover". sciohost.org.
http://www.sciohost.org/ncse/kvd/Padian/Padian_transcript.html#ear. Retrieved 2009-06-17. "Based
on testimony by Kevin Padian in the case of Kitzmiller v. Dover"

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 W000

Evolution of sexual reproduction

From Wikipedia, the free encyclopedia
(Redirected from Evolution of sex)
Jump to: navigation, search
Scientists currently have developed several competing hypotheses to explain the evolution of sexual
reproduction. Many groups of organisms, notably the majority of animals and plants, reproduce sexually. The
evolution of sex contains two related, yet distinct, themes: its origin and its maintenance. However, since the
hypotheses for the origins of sex are difficult to test experimentally, most current work has been focused on
the maintenance of sexual reproduction.
It seems that a sexual cycle is maintained because it improves the quality of progeny (fitness),
despite reducing the overall number of offspring (the two-fold cost of sex). In order for sex to be evolutionarily
advantageous, it must be associated with a significant increase in the fitness of offspring. One of the most
widely accepted explanations for the advantage of sex lies in the creation of genetic variation. Another
explanation is based on two molecular advantages. First is the advantage of recombinational DNA repair
(promoted during meiosis because homologous chromosomes pair at that time), and second is the
advantage of complementation (also known as hybrid vigor, heterosis or masking of mutations).
For the advantage due to creation of genetic variation, there are three possible reasons this might
happen. First, sexual reproduction can bring together mutations that are beneficial into the same individual
(sex aids in the spread of advantageous traits). Second, sex acts to bring together currently deleterious
mutations to create severely unfit individuals that are then eliminated from the population ( sex aids in the
removal of deleterious genes). Last, sex creates new gene combinations that may be more fit than previously
existing ones, or may simply lead to reduced competition among relatives.
For the advantage due to DNA repair, there is an immediate large benefit to removal of DNA damage
by recombinational DNA repair during meiosis, since this removal allows greater survival of progeny with
undamaged DNA. The advantage of complementation to each sexual partner is avoidance of the bad effects
of their deleterious recessive genes in progeny by the masking effect of normal dominant genes contributed
by the other partner.
The classes of hypotheses based on the creation of variation are further broken down below. It is
important to realise that any number of these hypotheses may be true in any given species (they are not
mutually exclusive), and that different hypotheses may apply in different species. However, a research
framework based on creation of variation has yet to be found that allows one to determine whether the
reason for sex is universal for all sexual species, and, if not, which mechanism is acting in each species.
On the other hand, the maintenance of sex based on DNA repair and complementation applies
widely to all sexual species. This explanation for the maintenance of sex is explored further in Section 6.2.
Contents
[hide]
• 1 Historical perspective
• 2 Two-fold cost of sex
• 3 Promotion of genetic variation
• 4 Spread of advantageous traits
• 4.1 Novel genotypes
• 4.2 Increased resistance to parasites
• 5 Deleterious mutation clearance
• 5.1 Maintenance of mutation-free individuals
• 5.2 Removal of deleterious genes
• 6 Other explanations
• 6.1 Speed of evolution
• 6.2 DNA repair and complementation
• 7 Origin of sexual reproduction
• 8 Mechanistic origin of Sexual Reproduction
• 8.1 Viral Eukaryogenesis
• 8.2 Neomuran Revolution
• 9 See also
• 10 References
• 11 Further reading
[edit] Historical perspective
Modern philosophical-scientific thinking on the problem can be traced back to Erasmus Darwin in the
18th century; it also features in Aristotle's writings. The thread was later picked up by August Weismann in
1889, who understood that the purpose of sex was to generate genetic variation, as is detailed in the majority
of the explanations below. On the other hand, Charles Darwin, also in 1889, concluded that the effects of
hybrid vigor (complementation) “is amply sufficient to account for the ... genesis of the two sexes.” This is
consistent with the repair and complementation hypothesis, given below under “Other explanations.”
Several explanations have been suggested by biologists including W. D. Hamilton, Alexey
Kondrashov, George C. Williams, Harris Bernstein, Carol Bernstein, Michael M. Cox, Frederic A. Hopf and
Richard E. Michod to explain how sexual reproduction is maintained in a vast array of different living
organisms.

[edit] Two-fold cost of sex

 This diagram illustrates the two-fold cost of sex. If each individual were to contribute to the same
number of offspring (two), (a) the sexual population remains the same size each generation, where the (b)
asexual population doubles in size each generation.
In all sexual species, the population consists of two sexes, only one of which is capable of bearing
young (with the exception of simultaneous hermaphrodites). In an asexual species, each member of the
population is capable of bearing young. This implies that an asexual population has an intrinsic capacity to
grow more rapidly each generation. The cost was first described in mathematical terms by John Maynard
Smith.[1] He imagined an asexual mutant arising in a sexual population, half of which comprises males that
cannot themselves produce offspring. With female-only offspring, the asexual lineage doubles its
representation in the population each generation, all else being equal. Often all else is not equal, however, in
which case the realised fitness cost to sex may be much less than this intrinsic twofold cost of producing
males. For example, an asexual mutant arising in a sexual population occupies a niche frozen to that of its
parental genotype because the asexual descendants are genetically self-identical. Analysis of competitive
Lotka-Volterra equations suggests that the asexual lineage may never realise its full twofold advantage in
population growth capacity, if the broader niche of the sexual population confers even a small competitive
edge.[2]
An additional cost is that males and females must search for each other in order to mate, and sexual
selection often favours traits that reduce the survival of individuals.[1]
Evidence that the cost is not insurmountable comes from George C. Williams, who noted the
existence of species which are capable of both asexual and sexual reproduction. These species time their
sexual reproduction with periods of environmental uncertainty, and reproduce asexually when conditions are
more favourable. The important point is that these species are observed to reproduce sexually when they
could choose not to, implying that there is a selective advantage to sexual reproduction.[3]
Another disadvantage of sexual reproduction is that a sexually reproducing organism will only be
able to pass on 50% of its genes to each offspring. This is a consequence of the fact that gametes from
sexually reproducing species are haploid.[4]

[edit] Promotion of genetic variation

August Weismann proposed in 1889[5] an explanation for the evolution of sex, where the advantage
of sex is the creation of variation among siblings. It was then subsequently explained in genetics terms by
Fisher[6] and Muller[7] and has been recently summarised by Burt in 2000.[8]
George C. Williams gave an example based around the elm tree. In the forest of this example, empty
patches between trees can support one individual each. When a patch becomes available because of the
death of a tree, other trees' seeds will compete to fill the patch. Since the chance of a seed's success in
occupying the patch depends upon its genotype, and a parent cannot anticipate which genotype is most
successful, each parent will send many seeds, creating competition between siblings. Natural selection
therefore favours parents which can produce a variety of offspring.
A similar hypothesis is named the tangled bank hypothesis after a passage in Charles Darwin's The
Origin of Species:
"It is interesting to contemplate an entangled bank, clothed with many plants of many kinds,
with birds singing on the bushes, with various insects flitting about, and with worms crawling through
the damp earth, and to reflect that these elaborately constructed forms, so different from each other,
 and dependent on each other in so complex a manner, have all been produced by laws acting around
us."
The hypothesis, proposed by Michael Ghiselin in his 1974 book, The Economy of Nature and the
Evolution of Sex, suggests that a diverse set of siblings may be able to extract more food from its
environment than a clone, because each sibling uses a slightly different niche. One of the main proponents of
this hypothesis is Graham Bell of McGill University. The hypothesis has been criticised for failing to explain
how asexual species developed sexes. In his book, Evolution and Human Behavior (MIT Press, 2000), John
Cartwright comments:
"Although once popular, the tangled bank hypothesis now seems to face many problems, and
former adherents are falling away. The theory would predict a greater interest in sex among animals
that produce lots of small offspring that compete with each other. In fact, sex is invariably associated
with organisms that produce a few large offspring, whereas organisms producing small offspring
frequently engage in parthenogenesis [asexual reproduction]. In addition, the evidence from fossils
suggests that species go for vast periods of [geologic] time without changing much."
[edit] Spread of advantageous traits
[edit] Novel genotypes

This diagram illustrates how sex might create novel genotypes more rapidly. Two advantageous
alleles A and B occur at random. The two alleles are recombined rapidly in a sexual population (top), but in
an asexual population (bottom) the two alleles must independently arise because of clonal interference.
Sex could be a method by which novel genotypes are created. Since sex combines genes from two
individuals, sexually reproducing populations can more easily combine advantageous genes than can
asexual populations. If, in a sexual population, two different advantageous alleles arise at different loci on a
chromosome in different members of the population, a chromosome containing the two advantageous alleles
can be produced within a few generations by recombination. However, should the same two alleles arise in
different members of an asexual population, the only way that one chromosome can develop the other allele
is to independently gain the same mutation, which would take much longer.
Ronald Fisher also suggested that sex might facilitate the spread of advantageous genes by allowing
them to escape their genetic surroundings, if they should arise on a chromosome with deleterious genes.
But these explanations depend upon the rate of mutation. If favourable mutations are so rare that
each will become fixed in the population before the next arises (bearing in mind that mutation is a Poisson
process), then sexual and asexual populations would evolve at the same rate.
Additionally, these explanations depend upon group selection, which most theorists maintain is a
weak selective force relative to individual selection – sex is still disadvantageous to the individual due to the
twofold cost of sex. Therefore, these explanations do not explain why heterogonic species choose to adopt
sexual reproduction, as George C. Williams pointed out in his balance argument, and hence are insufficient
to explain the evolution of sex.
Supporters of these theories respond to the balance argument that the individuals produced by
sexual and asexual reproduction may differ in other respects too – which may influence the persistence of
sexuality. For example, in water fleas of the genus Cladocera, sexual offspring form eggs which are better
able to survive the winter.

[edit] Increased resistance to parasites

One of the most widely accepted theories to explain the evolution of sex is that it evolved as an
adaptation to assist sexual individuals in resisting parasites, also known as the Red Queen's Hypothesis.[4]
[9][10]
 When an environment changes, previously neutral or deleterious alleles can become favourable. If
the environment changed sufficiently rapidly (i.e. between generations), these changes in the environment
can make sex advantageous for the individual. Such rapid changes in environment are caused by the co-
evolution between hosts and parasites.
Imagine, for example that there is one gene in parasites with two alleles p and P conferring two types
of parasitic ability, and one gene in hosts with two alleles h and H, conferring two types of parasite
resistance, such that parasites with allele p can attach themselves to hosts with the allele h, and P to H. Such
a situation will lead to cyclic changes in allele frequency - as p increases in frequency, h will be disfavoured.
In reality, there will be several genes involved in the relationship between hosts and parasites. In an
asexual population of hosts, offspring will only have the different parasitic resistance if a mutation arises. In a
sexual population of hosts, however, offspring will have a new combination of parasitic resistance alleles.
In other words, like Lewis Carroll's Red Queen, sexual hosts are continually adapting in order to stay
ahead of their parasites.
Evidence for this explanation for the evolution of sex is provided by comparison of the rate of
molecular evolution of genes for kinases and immunoglobulins in the immune system with genes coding
other proteins. The genes coding for immune system proteins evolve considerably faster.[11][12]
Critics of the Red Queen hypothesis question whether the constantly-changing environment of hosts
and parasites is sufficiently common to explain the evolution of sex.
[edit] Deleterious mutation clearance
Mutations can have many different effects upon an organism. It is generally believed that the majority
of non-neutral mutations are deleterious, which means that they will cause a decrease in the organism's
overall fitness.[13] If a mutation has a deleterious effect, it will then usually be removed from the population
by the process of natural selection. Sexual reproduction is believed to be more efficient than asexual
reproduction in removing those mutations from the genome.[14]
There are two main hypotheses which explain how sex may act to remove deleterious genes from
the genome.

[edit] Maintenance of mutation-free individuals

Main article: Muller's ratchet
In a finite asexual population under the pressure of deleterious mutations, the random loss of
individuals without such mutations is inevitable. This is known as Muller's ratchet. In a sexual population,
however, mutation-free genotypes can be restored by recombination of genotypes containing deleterious
mutations.
This comparison will only work for a small population - in a large population, random loss of the most
fit genotype becomes unlikely even in an asexual population.
[edit] Removal of deleterious genes

Diagram illustrating different relationships between numbers of mutations and fitness. Kondrashov's
model requires synergistic epistasis, which is represented by the blue line - each mutation has a
disproproportionately large effect on the organism's fitness.
This hypothesis was proposed by Alexey Kondrashov, and is sometimes known as the deterministic
mutation hypothesis.[14] It assumes that the majority of deleterious mutations are only slightly deleterious,
and affect the individual such that the introduction of each additional mutation has an increasingly large effect
on the fitness of the organism. This relationship between number of mutations and fitness is known as
synergistic epistasis.
By way of analogy, think of a car with several minor faults. Each is not sufficient alone to prevent the
car from running, but in combination, the faults combine to prevent the car from functioning.
 Similarly, an organism may be able to cope with a few defects, but the presence of many mutations
could overwhelm its backup mechanisms.
Kondrashov argues that the slightly deleterious nature of mutations means that the population will
tend to be composed of individuals with a small number of mutations. Sex will act to recombine these
genotypes, creating some individuals with fewer deleterious mutations, and some with more. Because there
is a major selective disadvantage to individuals with more mutations, these individuals die out. In essence,
sex compartmentalises the deleterious mutations.
There has been much criticism of Kondrashov's theory, since it relies on two key restrictive
conditions. The first requires that the rate of deleterious mutation should exceed one per genome per
generation in order to provide a substantial advantage for sex. While there is some empirical evidence for it
(for example in Drosophila[15] and E. coli[16]), there is also strong evidence against it. Secondly, there
should be strong interactions among loci (synergistic epistasis), a mutation-fitness relation for which there is
only limited evidence. Conversely, there is also the same amount of evidence that mutations show no
epistasis (purely additive model) or antagonistic interactions (each additional mutation has a disproportionally
small effect).

[edit] Other explanations

[edit] Speed of evolution
Ilan Eshel suggested that sex prevents rapid evolution. He suggests that recombination breaks up
favourable gene combinations more often than it creates them, and sex is maintained because it ensures
selection is longer-term than in asexual populations - so the population is less affected by short-term
changes. This explanation is not widely accepted, as its assumptions are very restrictive.
Nevertheless, it has recently been shown in experiments with Chlamydomonas algae that sex can
remove the speed limit on evolution.[17]

[edit] DNA repair and complementation

As discussed in the earlier part of this article, conventionally, sexual reproduction is explained as an
adaptation for producing genetic variation through allelic recombination. As acknowledged above, however,
serious problems with this explanation have led many biologists to conclude that the benefit of sex is a major
unsolved problem in evolutionary biology.
An alternative “informational” approach to this problem led to the view that the two fundamental
aspects of sex, genetic recombination and outcrossing, are adaptive responses to the two major sources of
“noise” in transmitting genetic information. Genetic noise can occur as either physical damage to the genome
(e.g. chemically altered bases of DNA or breaks in the chromosome) or replication errors (mutations)[18][19]
[20] This alternative view is referred to as the repair and complementation hypothesis to distinguish it from
the traditional variation hypothesis.
The repair and complementation hypothesis assumes that genetic recombination is fundamentally a
DNA repair process, and that when it occurs during meiosis it is an adaptation for repairing the genomic DNA
that is passed on to progeny. Recombinational repair is the only repair process known which can accurately
remove double-strand damages in DNA, and such damages are both common in nature and ordinarily lethal
if not repaired. Recombinational repair is prevalent from the simplest viruses to the most complex
multicellular eukaryotes. It is effective against many different types of genomic damage, and, in particular is
highly efficient at overcoming double-strand damages. Studies of the mechanism of meiotic recombination
indicate that meiosis is an adaptation for repairing DNA.[21] These considerations form the basis for the first
part of the repair and complementation hypothesis.
In some lines of descent from the earliest organisms, the diploid stage of the sexual cycle, which was
at first transient, became the predominant stage because it allowed complementation, the masking of
deleterious recessive mutations (hybrid vigor or heterosis). Outcrossing, the second fundamental aspect of
sex, is maintained by the advantage of masking mutations and the disadvantage of inbreeding (mating with a
close relative) which allows expression of recessive mutations (commonly observed as inbreeding
depression). This is in accord with Charles Darwin[22] who concluded that the adaptive advantage of sex is
hybrid vigor, or as he put it, “the offspring of two individuals, especially if their progenitors have been
subjected to very different conditions, have a great advantage in height, weight, constitutional vigor and
fertility over the self fertilised offspring from either one of the same parents.”
However, outcrossing may be abandoned in favor of parthogenesis or selfing (which retain the
advantage of meiotic recombinational repair) under conditions in which the costs of mating are very high. For
instance, costs of mating are high when individuals are rare in a geographic area, such as when there has
been a forest fire and the individuals entering the burned area are the initial ones to arrive. At such times
mates are hard to find, and this favors parthenogenic species.
In the view of the repair and complementation hypothesis, the removal of DNA damage by
recombinational repair produces a new, less deleterious form of informational noise, allelic recombination, as
a by-product. This lesser informational noise generates genetic variation, viewed by some as the major effect
of sex, as discussed in the earlier parts of this article.
[edit] Origin of sexual reproduction
This section requires expansion.

All sexually reproducing organisms derive from a common ancestor which was a single celled
eukaryotic species.[citation needed] Many protists reproduce sexually, as do the multicellular plants, animals,
and fungi. There are a few species which have secondarily lost this feature, such as Bdelloidea and some
parthenocarpic plants.
Organisms need to replicate their genetic material in an efficient and reliable manner. The necessity
to repair genetic damage is one of the leading theories explaining the origin of sexual reproduction. Diploid
individuals can repair a mutated section of its DNA via homologous recombination, since there are two
copies of the gene in the cell and one copy is presumed to be undamaged. A mutation in an haploid
individual, on the other hand, is more likely to become resident, as the DNA repair machinery has no way of
knowing what the original undamaged sequence was.[18] The most primitive form of sex may have been one
organism with damaged DNA replicating an undamaged strand from a similar organism in order to repair
itself.[23]
Another theory is that sexual reproduction originated from selfish parasitic genetic elements that
exchange genetic material (that is: copies of their own genome) for their transmission and propagation. In
some organisms, sexual reproduction has been shown to enhance the spread of parasitic genetic elements
(e.g.: yeast, filamentous fungi).[24] Bacterial conjugation, a form of genetic exchange that some sources
describe as sex, is not a form of reproduction, but rather an example of horizontal gene transfer. However, it
does support the selfish genetic element theory, as it is propagated through such a "selfish gene", the F-
plasmid.[23] Similarly, it has been proposed that sexual reproduction evolved from ancient haloarchaea
through a combination of jumping genes, and swapping plasmids.[25]
A third theory is that sex evolved as a form of cannibalism. One primitive organism ate another one,
but rather than completely digesting it, some of the 'eaten' organism's DNA was incorporated into the 'eater'
organism.[23]
Sex may also be derived from prokaryotic processes. A comprehensive 'origin of sex as vaccination'
theory proposes that eukaryan sex-as-syngamy (fusion sex) arose from prokaryan unilateral sex-as-infection
when infected hosts began swapping nuclearised genomes containing coevolved, vertically transmitted
symbionts that provided protection against horizontal superinfection by more virulent symbionts. Sex-as-
meiosis (fission sex) then evolved as a host strategy to uncouple (and thereby emasculate) the acquired
symbiont genomes.[26]

[edit] Mechanistic origin of Sexual Reproduction

Though theories positing benefits that lead to the origin of sex are often problematic, several credible
theories on the evolution of the mechanisms of sexual reproduction have been proposed.

[edit] Viral Eukaryogenesis

The Viral Eukaryogenesis(VE) theory proposes that eukaryotic cells arose from a combination of a
lysogenic virus, an archaeon and a bacterium. This model suggests that the nucleus originated when the
lysogenic virus incorporated genetic material from the archaeon and the bacterium and took over the role of
information storage for the amalgam. The archaeal host transferred much of its functional genome to the
virus during the evolution of cytoplasm but retained the function of gene translation and general metabolism.
The bacterium transferred most of its functional genome to the virus as it transitioned into a mitochondrion.
[27]
For these transformations to lead to the eukaryotic cell cycle, the VE hypothesis specifies a pox-like
virus as the lysogenic virus. A pox-like virus is a likely ancestor because of its fundamental similarities with
eukaryotic nuclei. These include a double stranded DNA genome, a linear chromosome with short telomeric
repeats, a complex membrane bound capsid, the ability to produce capped mRNA, and the ability to export
the capped mRNA across the viral membrane into the cytoplasm. The presence of a lysogenic pox-like virus
ancestor explains the development of meiotic division, an essential component of sexual reproduction.[28]
Meiotic division in the VE hypothesis arose because of the evolutionary pressures placed on the
lysogenic virus as a result of its inability to enter into the lytic cycle. This selective pressure resulted in the
development of processes allowing the viruses to spread horizontally throughout the population. The
outcome of this selection was cell-to-cell fusion. (This is distinct from the conjugation methods used by
bacterial plasmids under evolutionary pressure, with important consequences.)[27] The possibility of this kind
of fusion is supported by the presence of fusion proteins in the envelopes of the pox viruses that allow them
to fuse with host membranes. These proteins could have been transferred to the cell membrane during viral
reproduction, enabling cell-to-cell fusion between the virus host and an uninfected cell. The theory proposes
meiosis originated from the fusion between two cells infected with related but different viruses which
recognised each other as uninfected. After the fusion of the two cells, incompatibilities between the two
viruses result in a meiotic-like cell division.[28]
The two viruses established in the cell would initiate replication in response to signals from the host
cell. A mitosis-like cell cycle would proceed until the viral membranes dissolved, at which point linear
chromosomes would be bound together with centromeres. The homologous nature of the two viral
centromeres would incite the grouping of both sets into tetrads. It is speculated that this grouping may be the
origin of crossing over, characteristic of the first division in modern meiosis. The partitioning apparatus of the
mitotic-like cell cycle the cells used to replicate independently would then pull each set of chromosomes to
one side of the cell, still bound by centromeres. These centromeres would prevent their replication in
subsequent division, resulting in four daughter cells with one copy of one of the two original pox-like viruses.
The process resulting from combination of two similar pox viruses within the same host closely mimics
meiosis.[28]

[edit] Neomuran Revolution

An alternative theory, proposed by Thomas Cavalier-Smith, was labeled the Neomuran Revolution.
The designation “Neomuran Revolution” refers to the appearances of the common ancestors of eukaryotes
and archaea. Cavalier-Smith proposes that the first neomurans emerged 850 million years ago. Other
molecular biologists assume that this group appeared much earlier, but Cavalier-Smith dismisses these
claims because they are based on the “theoretically and empirically” unsound model of molecular clocks.
Cavalier-Smith's theory of the Neomuran revolution has implications for the evolutionary history of the cellular
machinery for recombination and sex. It suggests that this machinery evolved in two distinct bouts separated
by a long period of stasis; first the appearance of recombination machinery in a bacterial ancestor which was
maintained for 3 Gy, until the neomuran revolution when the mechanics were adapted to the presence of
nucleosomes. The archaeal products of the revolution maintained recombination machinery that was
essentially bacterial, whereas the eukaryotic products broke with this bacterial continuity. They introduced
cell fusion and ploidy cycles into cell life histories. Cavalier-Smith argues that both bouts of mechanical
evolution were motivated by similar selective forces: the need for accurate DNA replication without loss of
viability.[29]
[edit] See also
Book:Sex

Books are collections of articles that can be downloaded or ordered in print.

• Asexual reproduction
• Biological reproduction
• Epistasis
• Genetic recombination
• Mutation
• Sexual reproduction

[edit] References
1. ^ a b John Maynard Smith The Evolution of Sex 1978.
2. ^ Doncaster, C. P. et al. 2000. The ecological cost of sex. Nature 404: 281-285
3. ^ George C. Williams Sex and Evolution 1975, Princeton University Press, ISBN 0-691-
08152-2
4. ^ a b Matt Ridley 1995 The Red Queen: Sex and the Evolution of Human Nature 1995
Penguin.
 5. ^ Weismann, A. 1889. Essays on heredity and kindred biological subjects. Oxford Univ.
Press, Oxford, UK
6. ^ Fisher, R. A. 1930. The genetical theory of natural selection. Clarendon Press, Oxford, UK
7. ^ Muller, H. J. 1932. Some genetic aspects of sex. Am. Nat. 8: 118–138.
8. ^ Burt, A. 2000. Perspective: sex, recombination, and the efficacy of selection—was
Weismann right? Evolution 54: 337–351.
9. ^ Van Valen, L. (1973) "A New Evolutionary Law." Evolutionary Theory 1, 1-30.
10.^ Hamilton, W. D. et al. "Sexual reproduction as an adaptation to resist parasites" 1990.
Proceedings of the National Academy of Sciences (PNAS) 87:3566-3573.
11.^ Kuma, K., Iwabe, N. & Miyata, T. 1995. Functional constraints against variations on
molecules from the tissue-level - slowly evolving brain-specific genes demonstrated by protein-
kinase and immunoglobulin supergene families. Molecular Biology and Evolution 12 (1): 123-130.
12.^ Wolfe KH & Sharp PM. 1993. Mammalian gene evolution - nucleotide-sequence divergence
between mouse and rat. Journal of molecular evolution 37 (4): 441-456 OCT 1993
13.^ Griffiths et al. 1999. Gene mutations, p197-234, in Modern Genetic Analysis, New York,
W.H. Freeman and Company.
14.^ a b Kondrashov, A. S. "Deleterious mutations and the evolution of sexual reproduction"
1988. Nature 336:435-440.
15.^ Whitlock, M. C., & Bourget, D. 2000. Factors affecting the genetic load in Drosophila:
synergistic epistasis and correlations among fitness components. Evolution 54:1654–1660.
16.^ Elena, S. F., & Lenski, R. E. 1997. Test of synergistic interactions among deleterious
mutations in bacteria. Nature 390:395–398.
17.^ Colegrave, N. 2002. Sex releases the speed limit on evolution. Nature 420: 664-666.
 18.^ a b Bernstein H, Byerly HC, Hopf FA, Michod RE (1984). "Origin of sex". J. Theor. Biol. 110
(3): 323–51. doi:10.1016/S0022-5193(84)80178-2. PMID 6209512.
19.^ Bernstein H, Byerly HC, Hopf FA, Michod RE (1985). "Genetic damage, mutation, and the
evolution of sex". Science 229 (4719): 1277–81. doi:10.1126/science.3898363. PMID 3898363.
20.^ Bernstein H, Hopf FA, Michod RE (1987). "The molecular basis of the evolution of sex".
Adv. Genet. 24: 323–70. doi:10.1016/S0065-2660(08)60012-7. PMID 3324702.
21.^ Cox MM (2001). "Historical overview: searching for replication help in all of the rec places".
Proc. Natl. Acad. Sci. U.S.A. 98 (15): 8173–80. doi:10.1073/pnas.131004998. PMID 11459950.
22.^ Darwin C. 1889. The effects of cross and self fertilisation in the vegetable kingdom.
Chapter XII. General Results pp. 436-463. D. Appleton and Company, New York
23.^ a b c Olivia Judson (2002). Dr. Tatiana's sex advice to all creation. New York: Metropolitan
Books. pp. 233–4. ISBN 0-8050-6331-5.
24.^ Hickey D (1982). "Selfish DNA: a sexually-transmitted nuclear parasite". Genetics 101 (3-
4): 519–31. PMID 6293914.
25.^ DasSarma, Shiladitya (2007). "Extreme Microbes". American Scientist. 95. pp. 224–231
26.^ Sterrer W (2002). "On the origin of sex as vaccination". Journal of Theoretical Biology 216
(4): 387–396. doi:10.1006/jtbi.2002.3008. PMID 12151256.
27.^ a b PJ Bell (2001). "Viral eukaryogenesis: was the ancestor of the nucleus a complex DNA
virus?". J Molec Biol 53 (3): 251–6.
28.^ a b c PJ Bell (2006). "Sex and the eukaryotic cell cycle is consistent with a viral ancestry for
the eukaryotic nucleus". J Theor Biol 243 (1): 54–63. doi:10.1016/j.jtbi.2006.05.015. PMID 16846615.
29.^ Cavalier-Smith, Thomas (2006). "Cell evolution and Earth history: stasis and revolution".
Royal Society of Biol Sci 361 (1470): 969–1006. doi:10.1098/rstb.2006.1842. PMID 16754610.
[edit] Further reading
• Bell, Graham (1982). The masterpiece of nature: the evolution and genetics of sexuality .
Berkeley: University of California Press. ISBN 0-520-04583-1.
• Bernstein, Harris; Carol Bernstein (1991). Aging, sex, and DNA repair. Boston: Academic
Press. ISBN 0-12-092860-4.
• Hurst, L.D.; J.R. Peck (1996). "Recent advances in the understanding of the evolution and
maintenance of sex". Trends in Ecology and Evolution 11: 46–52. doi:10.1016/0169-5347(96)81041-
X.
• Levin, Bruce R.; Richard E. Michod (1988). The Evolution of sex: an examination of current
ideas. Sunderland, Mass: Sinauer Associates. ISBN 0-87893-459-6.
• Maynard Smith, John (1978). The evolution of sex. Cambridge, UK: Cambridge University
Press. ISBN 0-521-21887-X.
• Michod, Richard E. (1995). Eros and evolution: a natural philosophy of sex. Reading, Mass:
Addison-Wesley Pub. Co. ISBN 0-201-40754-X.
• "Scientists put sex origin mystery to bed, Wild strawberry research provides evidence on
when gender emerges". MSNBC. http://www.msnbc.msn.com/id/27927661/. Retrieved 2008-11-25.
• Ridley, Mark (1993). Evolution. Oxford: Blackwell Scientific. ISBN 0-632-03481-5.
• Ridley, Mark (2000). Mendel's demon: gene justice and the complexity of life . London:
Weidenfeld & Nicolson. ISBN 0-297-64634-6.
• Ridley, Matt (1995). The Red Queen: sex and the evolution of human nature . New York:
Penguin Books. ISBN 0-14-024548-0.
• Szathmáry, Eörs; John Maynard Smith (1995). The Major Transitions in Evolution. Oxford:
W.H. Freeman Spektrum. ISBN 0-7167-4525-9.
 • Taylor, Timothy (1996). The prehistory of sex: four million years of human sexual culture .
New York: Bantam Books. ISBN 0-553-09694-X.
• Williams, George (1975). Sex and evolution. Princeton, N.J: Princeton University Press.
ISBN 0-691-08147-6.

[edit] External links

• The Origin of Sex: Cosmic Solution to an Ancient Mystery
• Why Sex is Good
• An essay summarising the different theories, dating from around 2001

[show]
v•d•e
The development of phenotype

Key concepts Genotype-phenotype distinction · Norms of reaction · Gene-

environment interaction · Heritability · Quantitative genetics

Dominance relationship · Epistasis · Polygenic inheritance ·

Genetic architecture Pleiotropy · Plasticity · Canalisation · Fitness landscape · Transgressive
phenotype
 Non-genetic
influences Epigenetics · Maternal effect · Dual inheritance theory

Developmental
architecture Segmentation · Modularity

Evolution of genetic
systems Evolvability · Mutational robustness · Evolution of sex

Influential figures C. H. Waddington · Richard Lewontin

Debates Nature versus nurture

List of evolutionary biology topics

[show]
v•d•e
Basic topics in evolutionary biology
 Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex

Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·

Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders
 Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_sexual_reproduction"

Categories: Evolutionary biology | Sex

[hide]
v•d•e
The development of phenotype

Key concepts Genotype-phenotype distinction · Norms of reaction · Gene-

 environment interaction · Heritability · Quantitative genetics

Dominance relationship · Epistasis · Polygenic inheritance ·

Genetic architecture Pleiotropy · Plasticity · Canalisation · Fitness landscape · Transgressive
phenotype

Non-genetic
influences Epigenetics · Maternal effect · Dual inheritance theory

Developmental
architecture Segmentation · Modularity

Evolution of genetic
systems Evolvability · Mutational robustness · Evolution of sex

Influential figures C. H. Waddington · Richard Lewontin

Debates Nature versus nurture

 W000

Evolution of the eye

From Wikipedia, the free encyclopedia

Jump to: navigation, search

 Major stages in the evolution of the eye.
The evolution of the eye has been a subject of significant study, as a distinctive example of a
homologous organ present in a wide variety of taxa. Certain components of the eye, such as the visual
pigments, appear to have a common ancestry – that is, they evolved once, before the animals radiated.
However, complex, image-forming eyes evolved some 50 to 100 times[1] – using many of the same proteins
and genetic toolkits in their construction. [2][3]
Complex eyes appear to have first evolved within a few million years, in the rapid burst of evolution
known as the Cambrian explosion. There is no evidence of eyes before the Cambrian, but a wide range of
diversity is evident in the Middle Cambrian Burgess shale.
Eyes show a wide range of adaptations to meet the requirements of the organisms which bear them.
Eyes may vary in their acuity, the range of wavelengths they can detect, their sensitivity in low light levels,
their ability to detect motion or resolve objects, and whether they can discriminate colours.
Contents
[hide]
• 1 History of research
• 2 Rate of evolution
• 3 One origin or many?
• 4 Stages of eye evolution
• 4.1 Early eyes
• 4.2 Lens formation and diversification
• 4.3 Other developments
• 5 Evolutionary baggage
• 6 Footnotes
• 7 References
• 8 External links
• 9 Further reading
[edit] History of research

The human eye, demonstrating the iris.

Since 1802, the evolution of a structure as complex as the projecting eye by natural selection has
been said to be difficult to explain.[4] Charles Darwin himself wrote, in his Origin of Species, that the
evolution of the eye by natural selection at first glance seemed "absurd in the highest possible degree".
However, he went on to explain that despite the difficulty in imagining it, it was perfectly feasible:
...if numerous gradations from a perfect and complex eye to one very imperfect and simple,
each grade being useful to its possessor, can be shown to exist; if further, the eye does vary
ever so slightly, and the variations be inherited, which is certainly the case; and if any variation
or modification in the organ be ever useful to an animal under changing conditions of life, then
 the difficulty of believing that a perfect and complex eye could be formed by natural selection,
though insuperable by our imagination, can hardly be considered real.[5]

He suggested a gradation from "an optic nerve merely coated with pigment, and without any other
mechanism" to "a moderately high stage of perfection", giving examples of extant intermediate grades of
evolution.[5]
Darwin's suggestions were soon shown to be correct, and current research is investigating the
genetic mechanisms responsible for eye development and evolution[6].

[edit] Rate of evolution

The first fossils of eyes appeared during the lower Cambrian period (about 540 million years ago).[7]
This period saw a burst of apparently rapid evolution, dubbed the "Cambrian explosion". One of the many
hypotheses for "causes" of this diversification, the "Light Switch" theory of Andrew Parker holds that the
evolution of eyes initiated an arms race that led to a rapid spate of evolution.[8] Earlier than this, organisms
may have had use for light sensitivity, but not for fast locomotion and navigation by vision.
Since the fossil record, particularly of the Early Cambrian, is so poor, it is difficult to estimate the rate
of eye evolution. Simple modelling, invoking small mutations exposed to natural selection, demonstrates that
a primitive optical sense organ based upon efficient photopigments could evolve into a complex human-like
eye in approximately 400,000 years.[9][note 1]
[edit] One origin or many?
Whether one considers the eye to have evolved once or multiple times depends somewhat on the
definition of an eye. Much of the genetic machinery employed in eye development is common to all eyed
organisms, which may suggest that their ancestor utilized some form of light-sensitive machinery – even if it
lacked a dedicated optical organ. However, even photoreceptor cells may have evolved more than once from
molecularly similar chemoreceptors, and photosensitive cells probably existed long before the Cambrian
explosion.[10] Higher-level similarities – such as the use of the protein crystalin in the independently derived
cephalopod and vertebrate lenses[11] – reflect the co-option of a protein from a more fundamental role to a
new function within the eye.[12]
Shared traits common to all light-sensitive organs include the family of photo-receptive proteins
called opsins. All seven sub-families of opsin were already present in the last common ancestor of animals.
In addition, the genetic toolkit for positioning eyes is common to all animals: the PAX6 gene controls where
the eye develops in organisms ranging from mice to humans to fruit flies.[13][14][15] These high-level genes
are, by implication, much older than many of the structures that they are today seen to control; they must
originally have served a different purpose, before being co-opted for a new role in eye development.[12]
Sensory organs probably evolved before the brain did—there is no need for an information-processing
organ (brain) before there is information to process.[16]
[edit] Stages of eye evolution

The stigma (2) of the euglena hides a light-sensitive spot.

The earliest predecessors of the eye were photoreceptor proteins that sense light, found even in
unicellular organisms, called "eyespots". Eyespots can only sense ambient brightness: they can distinguish
light from dark, sufficient for photoperiodism and daily synchronization of circadian rhythms. They are
insufficient for vision, as they can not distinguish shapes or determine the direction light is coming from.
Eyespots are found in nearly all major animal groups, and are common among unicellular organisms,
including euglena. The euglena's eyespot, called a stigma, is located at its anterior end. It is a small splotch
of red pigment which shades a collection of light sensitive crystals. Together with the leading flagellum, the
eyespot allows the organism to move in response to light, often toward the light to assist in photosynthesis,
[17] and to predict day and night, the primary function of circadian rhythms. Visual pigments are located in
the brains of more complex organisms, and are thought to have a role in synchronising spawning with lunar
cycles. By detecting the subtle changes in night-time illumination, organisms could synchronise the release
of sperm and eggs to maximise the probability of fertilisation.
Vision itself relies on a basic biochemistry which is common to all eyes. However, how this
biochemical toolkit is used to interpret an organism's environment varies widely: eyes have a wide range of
structures and forms, all of which have evolved quite late relative to the underlying proteins and molecules.
[17]
At a cellular level, there appear to be two main "designs" of eyes, one possessed by the protostomes
(molluscs, annelid worms and arthropods), the other by the deuterostomes (chordates and echinoderms).[17]
The functional unit of the eye is the receptor cell, which contains the opsin proteins and responds to
light by initiating a nerve impulse. The light sensitive opsins are borne on a hairy layer, to maximise the
surface area. The nature of these "hairs" differs, with two basic forms underlying photoreceptor structure:
microvilli and cilia.[18] In the protostomes, they are microvilli: extensions or protrusions of the cellular
membrane. But in the deuterostomes, they are derived from cilia, which are separate structures.[17] This
now looks like something of a simplification, as some microvilli contain traces of cilia – but other observations
appear to support a fundamental difference between protostomes and deuterostomes.[17] These
considerations centre on the response of the cells to light – some use sodium to cause the electric signal that
will form a nerve impulse, and others use potassium; further, protostomes on the whole construct a signal by
allowing more sodium to pass through their cell walls, whereas deuterostomes allow less through.[17]
This suggests that when the two lineages diverged in the Precambrian, they had only very primitive
light receptors, which developed into more complex eyes independently.
[edit] Early eyes
The basic light-processing unit of eyes is the photoreceptor cell, a specialized cell consisting of two
molecules in a membrane: the opsin, a light-sensitive protein, surrounding the chromophore, a pigment that
distinguishes colors. Groups of such cells are termed "eyespots", and have evolved independently
somewhere between 40 and 65 times. These eyespots permit animals to gain only a very basic sense of the
direction and intensity of light – enough to know when they are safely in a cave, for example, but not enough
to discriminate an object from its surroundings.[17]
Developing an optical system that can discriminate the direction of light to within a few degrees is
apparently much more difficult, and only six of the thirty-something phyla[note 2] possess such a system.
However, these phyla account for 96% of living species.[17]

The planarian has "cup" eyespots that can slightly distinguish light direction.
These complex optical systems started out as the multicellular eyepatch gradually depressed into a
cup, which first granted the ability to discriminate brightness in directions, then in finer and finer directions as
the pit deepened. While flat eyepatches were ineffective at determining the direction of light, as a beam of
light would activate the exact same patch of photo-sensitive cells regardless of its direction, the "cup" shape
of the pit eyes allowed limited directional differentiation by changing which cells the lights would hit
depending upon the light's angle. Pit eyes, which had arisen by the Cambrian period, were seen in ancient
snails,[clarification needed] and are found in some snails and other invertebrates living today, such as
planaria. Planaria can slightly differentiate the direction and intensity of light because of their cup-shaped,
heavily-pigmented retina cells, which shield the light-sensitive cells from exposure in all directions except for
the single opening for the light. However, this proto-eye is still much more useful for detecting the absence or
presence of light than its direction; this gradually changes as the eye's pit deepens and the number of
photoreceptive cells grows, allowing for increasingly precise visual information.[19]
When a photon is absorbed by the chromophore, a chemical reaction causes the photon's energy to
be transduced into electrical energy and relayed, in higher animals, to the nervous system. These
photoreceptor cells form part of the retina, a thin layer of cells that relays visual information,[20] as well as
the light and daylength information needed by the circadian rhythm system, to the brain. However, some
jellyfish, such as Cladonema, have elaborate eyes but no brain. Their eyes transmit a message directly to the
muscles without the intermediate processing provided by a brain.[16]
During the Cambrian explosion, the development of the eye accelerated rapidly, with radical
improvements in image-processing and detection of light direction.[21]
 The primitive nautilus eye functions similarly to a pinhole camera.
The "pinhole camera" eye was developed as the pit deepened into a cup, then a chamber. By
reducing the size of the opening, the organism achieved true imaging, allowing for fine directional sensing
and even some shape-sensing. Eyes of this nature are currently found in the nautilus. Lacking a cornea or
lens, they provide poor resolution and dim imaging, but are still, for the purpose of vision, a major
improvement over the early eyepatches.[22]
Overgrowths of transparent cells prevented contamination and parasitic infestation. The chamber
contents, now segregated, could slowly specialize into a transparent humour, for optimizations such as
colour filtering, higher refractive index, blocking of ultraviolet radiation, or the ability to operate in and out of
water. The layer may, in certain classes, be related to the moulting of the organism's shell or skin.
It is likely that a key reason eyes specialize in detecting a specific, narrow range of wavelengths on
the electromagnetic spectrum—the visible spectrum—is because the earliest species to develop
photosensitivity were aquatic, and only two specific wavelength ranges of electromagnetic radiation, blue and
green visible light, can travel through water. This same light-filtering property of water also influenced the
photosensitivity of plants.[23][24][25]

[edit] Lens formation and diversification

Light from a distant object and a near object being focused by changing the curvature of the lens.
Lenses evolved independently in a number of lineages. Simple 'pit-eyes' probably developed lenses
to improve the amount of light that reached the retina; the focal length of an early lobopod with lens-
containing simple eyes focussed the image behind the retina, so while no part of the image could be brought
into focus, the intensity of light allowed the organism to inhabit deeper (and therefore darker) waters.[26] A
subsequent increase of the lens's refractive index probably resulted in an in-focus image being formed.[26]
The development of the lens in camera-type eyes probably followed a different trajectory. The
transparent cells over a pinhole eye's aperture split into two layers, with liquid in between.[ citation needed]
The liquid originally served as a circulatory fluid for oxygen, nutrients, wastes, and immune functions,
allowing greater total thickness and higher mechanical protection. In addition, multiple interfaces between
solids and liquids increase optical power, allowing wider viewing angles and greater imaging resolution.
Again, the division of layers may have originated with the shedding of skin; intracellular fluid may infill
naturally depending on layer depth.[citation needed]
Note that this optical layout has not been found, nor is it expected to be found.[ citation
needed]Fossilization rarely preserves soft tissues, and even if it did, the new humour would almost certainly
close as the remains desiccated, or as sediment overburden forced the layers together, making the fossilized
eye resemble the previous layout.[citation needed]

Compound eye of Antarctic krill.

Vertebrate lenses are composed of adapted epithelial cells which have high concentrations of the
protein crystallin. In the embryo, the lens is living tissue, but the cellular machinery is not transparent so must
be removed before the organism can see. Removing the machinery means the lens is composed of dead
cells, packed with crystalins which must last the life of the organism. The refractive index gradient which
makes the lens useful is caused by the radial shift in crystallin concentration in different parts of the lens,
rather than by the specific type of protein: it is not the presence of crystallin, but the relative distribution of it,
that renders the lens useful.[27]
It is biologically difficult to maintain a transparent layer of cells. Deposition of transparent, nonliving,
material eased the need for nutrient supply and waste removal. Trilobites used calcite, a mineral which has
not been used by any other organism; in other compound eyes[ verification needed] and camera eyes, the
material is crystallin. A gap between tissue layers naturally forms a biconvex shape, which is optically and
mechanically ideal for substances of normal refractive index. A biconvex lens confers not only optical
resolution, but aperture and low-light ability, as resolution is now decoupled from hole size—which slowly
increases again, free from the circulatory constraints.
Independently, a transparent layer and a nontransparent layer may split forward from the lens: a
separate cornea and iris. (These may happen before or after crystal deposition, or not at all.) Separation of
the forward layer again forms a humour, the aqueous humour. This increases refractive power and again
eases circulatory problems. Formation of a nontransparent ring allows more blood vessels, more circulation,
and larger eye sizes. This flap around the perimeter of the lens also masks optical imperfections, which are
more common at lens edges. The need to mask lens imperfections gradually increases with lens curvature
and power, overall lens and eye size, and the resolution and aperture needs of the organism, driven by
hunting or survival requirements. This type is now functionally identical to the eye of most vertebrates,
including humans. Indeed, "the basic pattern of all vertebrate eyes is similar."[28]
[edit] Other developments
Color vision
The ability to see colors presents distinct selective advantages for species, such as being better able
to recognize predators, food and mates. Indeed, it is thought that simple sensory-neural mechanisms may
selectively control general behaviour patterns, such as escape, foraging, and hiding. Many examples of
wavelength-specific behaviour patterns have been identified, in two primary groups: less than 450nm,
associated with natural light sources, and greater than 450nm, associated with reflected light sources.[29] As
opsin molecules were subtly fine-tuned to detect different wavelengths of light, at some point color vision
developed when photoreceptor cells developed multiple pigments.[20] As a chemical adaption rather than a
mechanical one, this may have occurred at any of the early stages of the eye's evolution, and the capability
may have disappeared and reappeared as organisms became predator or prey. Similarly, night and day
vision emerged when receptors differentiated into rods and cones, respectively.
Further information: Evolution of color vision
Focusing mechanism
Some species move the lens back and forth, some stretch the lens flatter. Another mechanism
regulates focusing chemically and independently of these two, by controlling growth of the eye and
maintaining focal length. Note that a focusing method is not a requirement. As photographers know, focal
errors increase as aperture increases. Thus, countless organisms with small eyes are active in direct sunlight
and survive with no focus mechanism at all. As a species grows larger, or transitions to dimmer
environments, a means of focusing need only appear gradually.
Location
 Prey generally have eyes on the sides of their head so to have a larger field of view, from which to
avoid predators. Predators, however, have eyes in front of their head in order to have better depth
perception.[30][31] Flatfish are predators which lie on their side on the bottom, and have eyes placed
asymmetrically on the same side of
the head. A transitional fossil from the
common symmetric position is
Amphistium.

[edit] Evolutionary
baggage
The eyes of many taxa record
their evolutionary history in their
imperfect design. The vertebrate eye,
for instance, is built "backwards and
upside down", requiring "photons of
light to travel through the cornea, lens,
aquaeous fluid, blood vessels,
ganglion cells, amacrine cells,
horizontal cells, and bipolar cells
before they reach the light-sensitive
rods and cones that transduce the light
signal into neural impulses – which are
then sent to the visual cortex at the
Vertebrates and octopuses developed the camera eye
independently. In the vertebrate version the nerve fibers pass in
front of the retina, and there is a blind spot where the nerves pass
through the retina. In the vertebrate example, 4 represents the blind
spot, which is notably absent from the octopus eye. In vertebrates, 1
represents the retina and 2 is the nerve fibers, including the optic
nerve (3), whereas in the octopus eye, 1 and 2 represent the nerve
fibers and retina respectively.
back of the brain for processing into meaningful patterns." [32] This reduction in efficiency may be countered
by the formation of a reflective layer, the tapetum, behind the retina. Light which is not absorbed by the retina
on the first pass may bounce back and be detected.
The camera eyes of cephalopods, in contrast, are constructed the 'right way out', with the nerves
attached to the rear of the retina. This means that they do not have a blind spot. This difference may be
accounted for by the origins of eyes; in cephalopods they develop as an invagination of the head surface
whereas in vertebrates they originate as an extension of the brain.

[edit] Footnotes
1. ^ David Berlinski, an intelligent design proponent, questioned the basis of the calculations,
and the author of the original paper refuted Berlinski's criticism.
• Berlinski, David (April 2001). Commentary magazine.
• Nilsson, Dan-E.. Beware of Pseudo-science: a response to David Berlinski's attack
on my calculation of how long it takes for an eye to evolve .
http://www.talkreason.org/articles/blurred.cfm#lund.
• "Evolution of the Eye" on PBS
2. ^ The precise number varies from author to author.

[edit] References
1. ^ Haszprunar (1995). "The mollusca: Coelomate turbellarians or mesenchymate annelids?".
in Taylor. Origin and evolutionary radiation of the Mollusca : centenary symposium of the
Malacological Society of London. Oxford: Oxford Univ. Press. ISBN 0-19-854980-6.
 2. ^ Kozmik, Z; Daube, Michael; Frei, Erich; Norman, Barbara; Kos, Lidia; Dishaw, Larry J.;
Noll, Markus; Piatigorsky, Joram (2003). "Role of Pax Genes in Eye Evolution A Cnidarian PaxB
Gene Uniting Pax2 and Pax6 Functions". Developmental Cell 5: 773–785. doi:10.1016/S1534-
5807(03)00325-3.
3. ^ Land, M.F. and Nilsson, D.-E., Animal Eyes, Oxford University Press, Oxford (2002).
4. ^ In 1802, William Paley claimed that the eye was a miracle of design.
5. ^ a b Darwin, Charles (1859). On the Origin of Species. London: John Murray.
6. ^ Gehring WJ (2005). "New perspectives on eye development and the evolution of eyes and
photoreceptors". J. Hered. 96 (3): 171–84. doi:10.1093/jhered/esi027. PMID 15653558.
7. ^ Parker, A. R. (2009). "On the origin of optics". Optics & Laser Technology.
doi:10.1016/j.optlastec.2008.12.020. edit
8. ^ Parker, Andrew (2003). In the Blink of an Eye: How Vision Sparked the Big Bang of
Evolution. Cambridge, MA: Perseus Pub.. ISBN 0738206075.
9. ^ Nilsson, D-E; Pelger S (1994). "A pessimistic estimate of the time required for an eye to
evolve". Proc R Soc Lond B 256 (1345): 53–58. doi:10.1098/rspb.1994.0048. PMID 8008757.
10.^ Nilsson, DE (1996). "Eye ancestry: old genes for new eyes". Current biology : CB 6 (1): 39–
42. PMID 8805210. edit
11.^ Zinovieva, R. (1999). "O-Crystallin, arginine kinase and ferritin from the octopus lens".
Biochimica et Biophysica Acta (BBA) - Protein Structure and Molecular Enzymology 1431: 512–517.
doi:10.1016/S0167-4838(99)00066-7. edit
12.^ a b Scotland, R. (2010). "Deep homology: A view from systematics". BioEssays : news and
reviews in molecular, cellular and developmental biology 32 (5): NA–ME.
doi:10.1002/bies.200900175. PMID 20394064. edit
 13.^ Halder, G; Callaerts, P; Gehring, WJ (1995). "New perspectives on eye evolution". Current
opinion in genetics & development 5 (5): 602–9. doi:10.1016/0959-437X(95)80029-8.
PMID 8664548. edit
14.^ Halder, G.; Callaerts, P.; Gehring, W. (1995). "Induction of ectopic eyes by targeted
expression of the eyeless gene in Drosophila". Science 267 (5205): 1788.
doi:10.1126/science.7892602. PMID 7892602. edit
15.^ Tomarev, SI; Callaerts, P; Kos, L; Zinovieva, R; Halder, G; Gehring, W; Piatigorsky, J
(1997). "Squid Pax-6 and eye development". Proceedings of the National Academy of Sciences of
the United States of America 94 (6): 2421–6. doi:10.1073/pnas.94.6.2421. PMID 9122210. edit
16.^ a b Gehring, W. J. (13 January 2005). "New Perspectives on Eye Development and the
Evolution of Eyes and Photoreceptors" (Full text). Journal of Heredity (Oxford Journals) 96 (3): 171–
184. doi:10.1093/jhered/esi027. PMID 15653558.
http://jhered.oxfordjournals.org/cgi/content/full/96/3/171. Retrieved 2008-04-26.
17.^ a b c d e f g h M F Land; R D Fernald (1992). "The Evolution of Eyes". Annual Review of
Neuroscience 15: 1–29. doi:10.1146/annurev.ne.15.030192.000245. PMID 1575438.
18.^ Autrum, H (1979). "Introduction". in H. Autrum (editor). Comparative Physiology and
Evolution of Vision in Invertebrates- A: Invertebrate Photoreceptors . Handbook of Sensory
Physiology. VII/6A. New York: Springer-Verlag. pp. 6–9. ISBN 3540088377
19.^ Eye-Evolution?
20.^ a b Fernald, Russell D. (2001). The Evolution of Eyes: How Do Eyes Capture Photons?
Karger Gazette 64: "The Eye in Focus".
21.^ Conway-Morris, S. (1998). The Crucible of Creation. Oxford: Oxford University Press.
22.^ Dawkins, Richard (1986). The Blind Watchmaker.
 23.^ Fernald, Russell D. (2001). The Evolution of Eyes: Why Do We See What We See? Karger
Gazette 64: "The Eye in Focus".
24.^ Fernald, Russell D. (1998). Aquatic Adaptations in Fish Eyes. New York, Springer.
25.^ Fernald RD (1997). "The evolution of eyes". Brain Behav. Evol. 50 (4): 253–9.
doi:10.1159/000113339. PMID 9310200.
26.^ a b Schoenemann, B.; Liu, J. N.; Shu, D. G.; Han, J.; Zhang, Z. F. (2008). "A miniscule
optimized visual system in the Lower Cambrian". Lethaia 42: 265. doi:10.1111/j.1502-
3931.2008.00138.x. edit
27.^ Fernald, Russell D. (2001). The Evolution of Eyes: Where Do Lenses Come From? Karger
Gazette 64: "The Eye in Focus".
28.^ Ali, Mohamed Ather; Klyne, M.A. (1985). Vision in Vertebrates. New York: Plenum Press.
p. 1. ISBN 0-306-42065-1.
29.^ Menzel, Randolf (1979). "Spectral Sensitivity and Color Vision in Invertebrates". in H.
Autrum (editor). Comparative Physiology and Evolution of Vision in Invertebrates- A: Invertebrate
Photoreceptors. Handbook of Sensory Physiology. VII/6A. New York: Springer-Verlag. pp. 504–506;
551–558. ISBN 3540088377
30.^ http://www.blm.gov/id/st/en/prog/wildlife/carnivores.html
31.^ Boroditsky, Lera (6/24/99). "Light & Eyes: Lecture Notes". Lecture Notes. Stanford.
http://www-psych.stanford.edu/~lera/psych115s/notes/lecture2/. Retrieved 11 May 2010.
32.^ Dr. Michael Shermer, as quoted by Christopher Hitchens in his book "God is Not Great"
(pg.82)
[edit] External links
• "Evolution of the Eye". WGBH Educational Foundation and Clear Blue Sky Productions .
PBS. 2001. http://www.pbs.org/wgbh/evolution/library/01/1/l_011_01.html.
• Creationism Disproved? Video from the National Center for Science Education on the
evolution of the eye
• Evolution: Education and Outreach Special Issue: Evolution and Eyes volume 1, number 4,
October, 2008, pages 351-559. ISSN 1936-6426 (Print) 1936-6434 (Online)
• Myers, PZ (2007-12-21). "Evolution of vertebrate eyes". Pharyngula. ScienceBlogs.
http://scienceblogs.com/pharyngula/2007/12/evolution_of_vertebrate_eyes.php. Retrieved 2007-12-
23. "a review of Lamb TD, Collin SP, Pugh EN Jr. (2007) Evolution of the vertebrate eye: opsins,
photoreceptors, retina and eye cup. Nat Rev Neurosci 8(12):960-76."

[edit] Further reading

• Lamb TD, Collin SP, Pugh EN (December 2007). "Evolution of the vertebrate eye: opsins,
photoreceptors, retina and eye cup". Nat. Rev. Neurosci. 8 (12): 960–76. doi:10.1038/nrn2283.
PMID 18026166. illustration
• Land, M. F., & Nilsson, D-E, Animal Eyes, Oxford: Oxford University Press, 2002 ISBN 0-19-
8509685 "The origin of vision", Chapter 1, pages 1–15
• Journal Evolution: Education and Outreach Volume 1, Number 4 / October, 2008. Special
Issue: The Evolution of Eyes. 26 articles, free access.
 [hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
Taxa evolution Humans · Influenza · Insects · Lemur · Life · Molluscs · Plants · Sirenians (sea
cows) · Spiders

Modes of speciation Anagenesis · Catagenesis · Cladogenesis

History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_the_eye"

Categories: Evolution by phenotype | Eye

 W000

Histoire évolutive de l'œil

Un article de Wikipédia, l'encyclopédie libre.
(Redirigé depuis Histoire évolutive de l'oeil)
Aller à : Navigation, rechercher
L'œil est un système sensoriel utilisé dans l'ensemble du règne animal. En effet, la lumière donnant
un grand nombre d'informations sur l'environnement, elle est, et a été, une force sélective majeure, qui a
conduit à la formation d'une grande variété de systèmes capables de percevoir la lumière.
La définition de l'œil varie selon les biologistes, certains considérant qu'un œil est un organe capable
de produire une image à partir de différences d'intensités entre photorécepteurs[1]. Dans cet article, l'œil est
défini dans un sens plus large, comme étant l'association fonctionnelle d'au moins une cellule
photoréceptrice et une cellule pigmentée[2].
L'évolution de l'œil commence pendant la période pré-cambrienne, alors que se forme une opsine
primitive, à l'origine des opsines des yeux actuels, responsables de la photoréception. La radiation évolutive
du Cambrien est ensuite marquée par l'évolution convergente d'yeux complexes chez de nombreux
embranchements formés à cette époque. Les yeux ainsi formés présentent une grande diversité de
structures, notamment sur la manière dont la lumière est captée et concentrée. Par la suite, les yeux
évoluent sous l'influence des pressions sélectives de leur environnement, et les changement d'habitats ou de
mode de vie sont souvent associés à des changements importants dans la structure et la fonction de l'œil.
Sommaire
[masquer]
• 1 Diversité des yeux
• 1.1 Diversité fonctionnelle
• 1.2 Diversité structurale
• 2 Pressions sélectives contraignant l'évolution de l'œil
• 3 Formation du proto-œil
• 4 Diversification de l'œil
• 4.1 Évolution des pigments photosensibles
• 4.1.1 Chez les Vertébrés
• 4.1.1.1 Méthode d'étude
• 4.1.1.2 La vision en faible
luminosité
• 4.1.1.3 Perception des couleurs
rouges et vertes
• 4.1.1.4 Perception du violet et des
ultraviolets
• 4.1.2 Chez les Insectes
• 4.1.2.1 Évolution des
chromophores
• 4.1.2.2 Phylogénie des opsines
Diversité des yeux [modifier]
Environ 1/3 des 33 embranchements animaux existants possède des organes photosensibles, et un
autre tiers possède des yeux, le dernier tiers n'ayant a priori pas d'organes spécialisés dans la détection de
la lumière[1]. On constate aujourd'hui une importante diversité, aussi bien dans la structure morphologique,
anatomique ou moléculaire de l'œil, que dans la type d'information prélevé à partir de la lumière[3].

Diversité fonctionnelle [modifier]

Les yeux, s'ils permettent tous de percevoir et d'analyser la lumière, n'en retirent pas tous les mêmes
informations. En effet, peuvent-être analysées :
• La direction, ce qui nécessite différentes orientation des pigments.
• L'intensité. Tous les yeux sont capables de percevoir l'intensité lumineuse. Cependant, il
existe une grande variabilité dans la sensibilité (intensité minimale perçue) et la qualité de contraste
(différence minimale d'intensité perçue) au sein du règne animal.
• La longueur d'onde. Alors que certains yeux ne détectent pas du tout la longueur d'onde de
la lumière incidente, de nombreux yeux sont capables de percevoir les couleurs. La perception des
couleurs et des contrastes varie grandement selon le embranchement (parfois même le genre)
considéré.
• La formation d'une image. La concentration de pigments dans une zone de la rétine, la
fovéa, permet la formation d'une image chez certains yeux. Seuls 6 embranchements animaux (les
Cnidaires, les Mollusques, les Arthropodes, les Chordés, les Annélides et les Onychophores) sur 33
possèdent des yeux capables de former des images, mais ces embranchements représentent, à eux
seuls, 96% des espèces animales connues[4]. Là encore, la résolution de l'image formée varie
énormément d'une espèce à l'autre.

Diversité structurale [modifier]

Toutes les structures de l'œil présentent une variabilité, parfois associée à différentes fonctions :
• Il existe de nombreux pigments différents, composés d'une opsine (5 classes différentes
chez les Vertébrés) et d'un chromophore (2 différents). Le nombre de pigments différents, et le type
de pigments présents, diffèrent entre les espèces, et même entre les différentes cellules
photoréceptrices d'une même espèce. De plus, il peut exister un polymorphisme spécifique. au
niveau du nombre et du type de pigment présents.
• Les cellules photoréceptrices peuvent être ciliées ou rhabdomérique (et possèdent alors des
microvillosités).
• La vision chromatique peut-être obtenue par une composition variée de pigments (chez
l'homme, par exemple), mais aussi par la présence de gouttelettes lipidiques dans la rétine, et
formant un filtre coloré.
• Les yeux peuvent ne contenir qu'une seule chambre oculaire (œil camérulaire, chez les
Vertébrés et les Céphalopodes) ou être composés de multiples yeux élémentaires appelés
ommatidies (œil composé, chez la majorité des Arthropodes).
• S'il y a création d'une image, celle-ci peut-être formée par ombrage, par réfraction ou par
réflexion, ces trois types d'images étant trouvé dans la plupart des embranchements pouvant former
des images.
• Les crystallines, responsables de la réfraction dans la lentille, sont généralement taxon-
spécifique.
 • La position et le nombre d'yeux sont très variables selon les espèces.

Pressions sélectives contraignant l'évolution de l'œil [modifier]

La lumière a été la principale force sélective sur Terre[5], résultant en l'apparition de nombreux
systèmes photorécepteurs dans l'ensemble du vivant, permettant le développement de rythmes circadiens et
d'horloges biologiques réglés sur la lumière du Soleil, ainsi que d'yeux permettant l'orientation par rapport à
la lumière environnante.
L'évolution de l'œil est, en conséquence, dépendante de la quantité et du type de lumière présent
dans son environnement, et on observe de très nombreuses adaptations liées à l'environnement lumineux
des animaux.
Par ailleurs, la perception de la lumière est limitée par des contraintes physiques, notamment les lois
de l'optique, qui limitent le nombre de possibilités pour collecter la lumière. Ceci explique la convergence de
structure d'un grand nombre d'yeux. C'est ainsi que les poissons et les céphalopodes primitifs ont développé
indépendamment une lentille sphérique, réponse adaptée à la collecte de lumière en milieu aquatique[6].

Formation du proto-œil [modifier]

Les premières perceptions de la lumière ont probablement évolué au début du Cambrien, grâce à la
formation d'une proto-opsine, ancêtre de toutes les opsines actuelles[7]. Cette molécule aurait intégré la
membrane de la première cellule photoréceptrice, grâce à un domaine protéique constitué de 7 hélices
transmembranaires, lequel aurait été réutilisé à partir des protéines impliquées dans la chémoréception,
c'est-à-dire la perception de molécules chimiques.
 Cette proto-opsine s'est ensuite associée au rétinal, formant un groupe de photopigments (les
opsines se liant au rétinal, ou retinal-binding opsins) présents aussi bien chez les animaux que chez les
Eubactéries et certaines algues unicellulaires.
Par ailleurs, le gène Pax6, jouant un rôle important dans le développement de l'œil chez les
Métazoaires, est associé très tôt à ce photopigment au cours de l'évolution. Ainsi, la forme ancestrale de
Pax6 était probablement impliquée dans le développement des premiers organes ou organites
photosenseurs.
Enfin, on retrouve des cellules ciliées et rhabdomériques dans la plupart des phyla dotés d'yeux, ce
qui suggère une différenciation de ces deux types de cellules antérieure à l'explosion cambrienne et
l'apparition convergente des nombreuses formes d'yeux.

Diversification de l'œil [modifier]

Lors de la radiation évolutive du Cambrien (ou Big Bang de l'évolution des animaux), des yeux plus
complexes ("pit eye") sont formés à de très nombreuses reprises (au moins 40 fois[8]), de manière
indépendante, à partir de l'œil primitif. Ces yeux ont une structure en forme de puits, dans lequel les cellules
réceptrices tapissent le fond de la cavité.
Il y a alors évolution de l'opsine dans les différents phyla aboutissant à la formation de différentes
familles d'opsines. L'évolution des opsines est alors souvent influencée par l'environnement lumineux des
différents animaux. De plus, le mécanisme de collecte de la lumière évolue en parallèle, et aboutit dans de
nombreux cas à la créations de nouveaux systèmes de collecte de la lumière et de formation d'images,
parfois de manière convergente.
Évolution des pigments photosensibles [modifier]
L'ensemble des organismes animaux dotés d'yeux perçoivent la lumière par des photopigments.
Ceux-ci sont tous issues d'une même photopigment primitif ayant évolué avant la radiation cambrienne. Les
photopigments présentent donc de nombreuses caractéristiques communes. Ils sont constituées d'une partie
protéique membranaire (apoprotéine), l'opsine, et d'une partie lipidique, le chromophore. L'apoprotéine est
constituée de 7 hélices transmembranaires séparées par des boucles hydrophiles reliant ces domaines,
alternativement cytoplasmique et extracellulaires. Le chromophore est un dérivé de la vitamine A (ou retinal),
qui présente un grand nombre de doubles liaisons conjuguées permettant l'absorption de photons à une
longueur d'onde particulière par passage d'une configuration 11-cis à une configuration all-trans. Il s'agit
toujours du 11-cis-retinal ou du 11-cis-3,4-dehydroretinal. La réponse cellulaire est déclenchée suite à
l'activation du photopigment dont l'opsine déclenche, via un second messager, une cascade d'activation.
On remarque que la plupart des opsines actuelles ont un maximum d'absorption dans le bleu, aux
alentours de 500nm. Ceci est dû aux origines des pigments, qui ont tout d'abord évolué dans un milieu
aquatique. L'eau absorbant la plupart des radiations lumineuses, seule persiste en profondeur la lumière
bleue de longueur d'onde d'environ 480nm. Ainsi, les pigments ancestraux préférentiellement sélectionnés
étaient ceux capables d'absorber ces longueurs d'onde. La spécificité des pigments pour certaines longueurs
d'onde préférentielles a ensuite évolué, souvent en adéquation avec l'environnement lumineux.
Deux parties de l'opsine sont extrêmement conservées au cours de l'évolution des différentes
lignées animales : la boucle cytoplasmique reliant les hélices 1 et 2, ainsi que le site d'attachement covalent
au chromophore situé au sein de la septième hélice transmembranaire. Cette conservation montre
l'importance de ces deux sites dans la fonction de l'opsine.
 Les autres régions, moins conservées, permettent de suivre l'évolution des opsines dans les
différents embranchements animaux. Si l'évolution des opsines est bien étudiée dans certains taxa, tels les
Insectes et les Vertébrés, on connait moins la manière dont elles se sont séparées avant la formation des
grands embranchements. Cependant, on sait que l'ensemble des opsines des Vertébrés s'individualisent
nettement des opsines des autres Métazoaires[9], ce qui montre qu'une seule opsine a été à l'origine des
diverses opsines chez les Vertébrés. Ainsi, la suite se concentre sur l'évolution des opsines dans certains
embranchements particuliers.

Chez les Vertébrés [modifier]

On distingue chez les Vertébrés 6 grandes familles d'opsines :
• Les Rhodopsines (ou RH1) exprimées préférentiellement dans les cellules en bâtonnets.
• Les pigments proches de la Rhodopsine (Rhodopsin-like pigments ou RH2).
• Les pigments sensibles aux courtes longueurs d'ondes de type 1 et 2 ( Short wavelength
sensitive pigments, ou SWS1 et SWS2).
• Les pigments sensibles aux longueurs d'ondes moyennes à longues ( Middle and long
wavelength sensitive pigments, ou M/LWS).
• Les pigments pinéaux (ou pigments P) ne se trouvent pas dans la rétine, mais exclusivement
dans la glande pinéale.
Les opsines de ces 5 familles s'avèrent être plus proches les unes des autres que de n'importe
quelle autre opsine animale, et la proximité évolutive est plus proche au sein de chaque famille qu'entre les
familles. Ainsi, ces 6 familles de pigments se sont différenciées après la séparation des Vertébrés des autres
embranchements animaux, et avant la séparation des Vertébrés en ordres distincts. Ces familles se sont
formées par 5 duplications de gènes différentes, puis spécialisation de chacun des deux gènes formés dans
la perception de certaines longueur d'ondes. Ainsi, les 4 familles de pigments des cellules en cônes se sont
différenciées, d'abord, et les pigments des cellules en bâtonnets se sont ensuite individualisés à partir des
pigments de type RH2.
└─o Pigment ancestral des Vertébrés
├─o
│ ├─o Pigments P
│ │
│ └─o Pigments M/LWS
└─o
├─o Pigments SWS1
└─o
├─o Pigments SWS2
└─o
├─o Pigments RH2
│
└─o Pigments RH1

Par la suite, certaines opsines ont été perdues par divers taxons. L'homme, par exemple, ne
possède ni pigment SWS2, ni pigment RH2. Ces opsines ont été perdues au cours de l'évolution des
Mammifères. Aucun Mammifère ne possède de pigment RH2, et seul l'Ornithorynque possède des pigments
SWS2. Ainsi, les pigments RH2 ont régressé chez l'ancêtre des mammifères, alors que les pigments SWS2
ont probablement régressé chez l'ancêtre des Mammifères placentaires, après la séparation des
Monotrèmes.
À l'heure actuelle, l'évolution des opsines RH1 (responsable de la vision en faible luminosité), SWS1
(responsable de la vision ultraviolette) et M/LWS (responsable de la vision chromatique en rouge/vert) est
mieux connue que celle des autres opsines des Vertébrés.

Méthode d'étude [modifier]

Une méthode apportant un grand nombre d'informations sur l'évolution des opsines est celle utilisée
par S. Yokoyama et al[10]. Elle consiste à construire une phylogénie des différents pigments, puis à prédire,
par inférence, la séquence peptidique des opsines ancestrales correspondantes aux différents nœuds. Enfin,
les pigments ancestraux inférés sont synthétisés, et la longueur d'onde d'absorption maximale des différents
pigments, actuels et synthétiques, est mesurée. On peut ainsi avoir une idée du spectre d'absorption des
différents pigments ancestraux et de la vision de l'ancêtre, et retracer plus finement les transitions évolutives.
De plus, certaines mutations (sélectionnées après comparaison avec les pigments actuels) sont introduites
dans des pigments ancestraux. Leur effet sur la longueur d'onde d'absorption maximale est alors évalué. On
peut ainsi chercher à retrouver les mutations ayant des modifications importantes dans les caractéristiques
d'absorption des pigments, et qui auraient pu être sélectionnées.
Cette méthode est néanmoins soumise à plusieurs biais. Tout d'abord, l'effet des mutations
ponctuelles dépend fortement de la séquence peptidique du reste de la molécule, la même mutation pouvant
avoir des modifications de longueur d'onde opposées dans deux pigments différents. Ainsi, on ne peut
prédire les changements survenus par le passé à partir des mutations observées sur les pigments
contemporains. De plus, la plupart des mutations prédites et étudiées ne se retrouvent jamais dans la nature,
et il faudrait étudier l'influence des mutations réellement observées.
La vision en faible luminosité [modifier]
Les pigments RH1 se trouvent essentiellement dans les cellules en bâtonnets des Vertébrés, et sont
par conséquent responsables de la vision en faible luminosité et ne servent pas à la discrimination des
couleurs. L'évolution de ces pigments est marquée par de nombreuses mutations responsables de
modification de la longueur d'onde d'absorption maximale du photopigment. Si le pigment RH1 de l'ancêtre
des Vertébrés (vivant en eau de surface) absorbait le mieux autour de 500nm, on observe aujourd'hui un
décalage de l'absorption du pigment RH1 vers le bleu chez une grand nombre de Vertébrés vivant en milieu
aquatique profond. Ceci traduit probablement une adaptation à l'absorption des longueurs d'ondes
supérieures par l'eau, imposant la sélection convergente dans plusieurs ordres des individus capables de
détecter la lumière atteignant les profondeurs.

Perception des couleurs rouges et vertes [modifier]

Il existe au moins quatre manières de percevoir les couleurs à partir du système dichromatique vert-
rouge chez les Vertébrés :
• Grâce aux pigments M/LWS ;
• Grâce aux pigments RH2 (ces derniers ayant perdu leur fonction chez les Mammifères) ;
• Grâce au couplage de l'un ou l'autre des deux chromatophores à la molécule d'opsine. La
perception des couleurs peut alors être régulée par l'individu. C'est le cas de nombreux poissons,
oiseaux et reptiles qui ne possèdent pas de pigment MWS[11] ;
• Grâce à la présence de différents types de gouttelettes lipidiques dans des cellules
photoréceptrices spécialisées. Ces gouttelettes jouent alors le rôle de filtre pour certaines longueurs
 d'onde et permettent une discrimination des couleurs. On retrouve notamment ce système chez
certaines espèces d'oiseaux[12].
L'évolution du système de perception des couleurs par les pigments M/LWS s'est faite de manière
convergente chez plusieurs ordres de Vertébrés comme les Cervidés et les Primates. Le pigment ancestral
est un pigment LWS dont la longueur d'onde d'absorption maximale avoisinait probablement les 560nm. Il y
a ensuite eu de manière répétée une duplication du gène codant le pigment LWS, et une évolution de l'une
de ces copies vers une absorption des longueurs d'ondes plus faibles, correspondant au vert. Cette transition
vers les faibles longueurs d'ondes s'est faite par un nombre limité de mutations, et est principalement due à
trois mutations qui ont été sélectionnées de manière répétée dans des familles distinctes de Vertébrés. Il y a
donc dans ce cas convergence fonctionnelle et génétique de l'évolution de la vision en rouge/vert.
L'avantage évolutif conféré par la perception des couleurs reste controversé. En effet, si la
perception des couleurs permet d'obtenir une quantité plus importante d'informations sur son environnement
(notamment sur ses prédateurs, ses proies ou ses partenaires potentiels), elle réussit plus difficilement à
déceler les camouflages par la couleur. On a aujourd'hui du mal à prendre en compte tous les avantages
sélectifs qui peuvent être impliqués dans la perception ou non des couleurs, et il est probable que cet
avantage dépende fortement du contexte écologique de l'espèce.

Perception du violet et des ultraviolets [modifier]

Chez les Vertébrés, le violet et les ultraviolets sont détectés grâce aux pigments SWS1 et SWS2. Ce
dernier pigment ayant perdu sa fonction chez les Mammifères placentaires, il a été beaucoup moins étudié et
on connait encore mal son évolution. Nous nous concentrerons donc sur la vision violette et ultraviolette
permise par le pigment SWS1 par la suite.
 L'ancêtre des Vertébrés possédait un pigment SWS1 dont la longueur d'absorption maximale
approchait probablement les 360nm, ce qui lui permettait probablement de voir les ultraviolets. Cependant ce
pigment a évolué plusieurs fois de manière convergente vers une absorption du bleu, notamment chez
l'ancêtre des Oiseaux et chez plusieurs ordres de Mammifères. Cette perte répétée de la vision ultraviolette a
été sélectionnée pour deux raisons principales : Tout d'abord, les ultraviolets étant très énergétiques, ils
peuvent endommager les tissus rétiniens et anéantir la fonction de l'œil. La disposition de pigments jaunes
dans la lentille et la cornée, absorbant les ultraviolets et protégeant la rétine, a donc été sélectionnée au
cours de l'évolution, ce qui est défavorable à la vision ultraviolette. De plus, le passage de l'absorption
maximale des pigments SWS1 vers le bleu permet une discrimination plus fine des tons de bleu et de violet.
Cependant, on observe un maintien de la vision ultraviolette chez plusieurs groupes de Vertébrés. La
souris marque par exemple son trajet d'excréments reflétant les ultraviolets, et s'en sert ainsi comme moyen
de repère et de communication. Certains oiseaux migrateurs sont même retournés à une vision ultraviolette.
Cette réversion aurait été sélectionnée en raison des capacités d'orientation par rapport au Soleil conférées
par la vision ultraviolette.

Chez les Insectes [modifier]

Évolution des chromophores [modifier]

L'ancêtre des Insectes n'utilisait que le 11-cis retinal (A1) comme chromophore, et la capacité à
utiliser le 11-cis-3-hydroxy retinal (A3) n'apparait qu'à la fin du Crétacé[13]. On retrouve des pigments A3
chez 9 ordres d'Insectes, dans lesquels la plupart des espèces utilisent soit le chromophore A1, soit le
chromophore A3, et quelques espèces sont capables d'utiliser les deux chromophores.
 Le changement de chromophore associé à une même opsine modifie légèrement le spectre
d'absorption du pigment. L'utilisation des deux chromophores chez une seule espèce pourrait ainsi permettre
une adaptation à des conditions visuelles différentes, notamment chez les Odonates où 44 des 46 espèces
étudiées possèdent les deux chromophores A1 et A3.
L'apparition convergente du chromophore A3 pendant le Crétacé pourrait être expliquée par
l'augmentation de la concentration en oxygène atmosphérique durant le Carbonifère. En effet, les
précurseurs des pigments A1 et A3, respectivement le carotène et les xanthophylles, sont issus des plantes.
Or la synthèse de xantophylle par les plantes à partir de carotène nécessite du dioxygène. Ainsi,
l'augmentation de la concentration de dioxygène pendant le Carbonifère aurait favorisé la synthèse de
xanthophylles chez les plantes et rendu ce précurseur plus accessible pour les Insectes, permettant
l'utilisation du chromophore A3.

Phylogénie des opsines [modifier]

L'étude phylogénétique des molécules d'opsines présentes dans différentes espèces d'insectes
permet de délimiter nettement trois groupes monopylétiques distincts regroupant des pigments excités par
des longueur d'onde proche[14]. On découpe ainsi les opsines des Insectes en trois classes, selon qu'elles
absorbent dans l'ultra-violet, le bleu ou le vert. Ces trois classes d'opsines sont retrouvées dans la plupart
des espèces étudiées, ce qui suggère que la différenciation de ces pigments a eu lieu chez l'ancêtre
commun des Insectes. Par ailleurs, des opsines sensibles au rouge sont apparues plusieurs fois de manière
convergente chez plusieurs espèces d'insectes. Enfin, on a pu observer une duplication du gène codant une
des opsines chez plusieurs espèces, dont la Drosophile.
Évolution des cellules photoréceptrices [modifier]
 Évolution des cellules photoréceptrices de l'œil chez les Métazoaires[15].
Une manière d'optimiser la perception de la lumière est d'augmenter la surface membranaire des
cellules réceptrices. On distingue ainsi trois types de cellules réceptrices, en fonction de la stratégie utilisée
pour augmenter la surface cellulaire :
• Les récepteurs ciliés possèdent une expansion cytoplasmique en forme de disque. On les
trouve majoritairement chez les Deutérostomiens. L'activation du pigment provoque, dans ces
cellules, une fermeture des canaux membranaires sodiques, et par conséquent une
hyperpolarisation de la membrane, enclenchant la réponse cellulaire[16].
• Les récepteurs rhabdomériques possèdent des microvillosités au niveau de la membrane
porteuse du pigment photosensible. Ces récepteurs, présents surtout chez les Protostomiens,
ouvrent les canaux sodiques en réponse à l'activation du pigment, ce qui dépolarise la membrane et
déclenche la réponse cellulaire[17].
• Les récepteurs multiciliés possédant de nombreux cils, sont plus rares et trouvés
uniquement chez certains Mollusques et Annélides. Ils répondent à l'activation du pigment par une
augmentation de la conductance au K+, et ne semblent être impliqués que dans les réponses
défensives lors de la détection de mouvement ou d'ombre[18].
On voit tout d'abord qu'il existe un nombre relativement faible de types cellulaires de récepteurs
photosensibles chez les animaux. Ceci est probablement dû au faible nombre de possibilités pour utiliser les
membranes dans la photoréception.
Par ailleurs, tous les photorécepteurs présents dans les yeux animaux semblent être des cellules
homologues, et dérivent donc toutes du même type cellulaire ancestral[15]. En effet, tous les
photorécepteurs étudiés utilisent des facteurs de différenciation homologues, tel pax6, et des protéines
effectrices homologues, telles que l'opsine, la rhodopsine kinase ou l'arrestine. Par ailleurs, si les deux
principaux types de récepteurs sont associés préférentiellement à certains taxons, on trouve des récepteurs
ciliés et rhabdomériques chez les Protostomiens comme chez les Deutérostomiens. Ceci suggère une
différenciation de ces deux types de cellules antérieure à la séparation Deutérostomiens/Protostomiens.
Enfin, même si la plupart des Vertébrés n'ont pas de cellules rhabdomériques, certaines cellules des
Deutérostomes semblent être homologues aux cellules rhabdomériques des Protostomes. Ainsi les cellules
des ganglions de la rétine, les cellules amacrines et les cellules horizontales seraient trois types cellulaires
frères, dérivés du récepteur rhabdomérique ancestral des Deutérostomiens.

Évolution des mécanismes de formation de l'image [modifier]

Après l'explosion cambrienne, un œil-puits se développe de manière indépendante dans de
nombreux taxons animaux ; en effet, on retrouve des yeux-puits dans 28 des 33 embranchements de
Métazoaires. Nombre de ces yeux ont alors évolué en structures plus complexes capable de former de
réelles images. Cette évolution convergente de fonction est due à l'important avantage sélectif conférée par
la formation d'images précises, riches en information. L'évolution convergente a pu être accélérée par la
concurrence entre les différents taxons : si un prédateur développe un œil efficace, la pression de sélection
augmente sur les proies, et l'avantage sélectif conféré par la formation d'image par la proie n'en devient que
plus grand.
L'évolution de l'œil a créé indépendamment différents mécanismes de formation des images. On
recense aujourd'hui 10 mécanismes permettant la formation d'images, et la plupart utilisent un système
optique (lentille ou miroir). De plus, même si certains systèmes optiques ont évolué de manière unique,
certains autres ont évolué de nombreuses fois indépendamment. Ceci s'explique par le faible nombre de
possibilités permettant de former une image d'une part, et par les contraintes imposés par le milieu sur le
système optique.
On distingue par leur structure (et non par leur origine évolutive) deux grands types d'yeux :
• L'œil camérulaire, ou œil simple, ou encore œil à une seule chambre. Il ne contient qu'une
seule cavité tapissée de cellules réceptrices, issue de l'œil-puits ancestral.
• L'œil composé, constitué d'un grand nombre d'yeux élémentaires appelés ommatidies.
Chaque ommatidie correspond à un œil-puits ancestral.
On trouve des yeux camérulaires aussi bien que des yeux composés dans la plupart des 6
embranchements animaux ayant développé un œil complexe.

Les yeux camérulaires [modifier]

L'œil-puits, présent dans la plupart des embranchements, a subi 3 types de transformations au cours
de l'évolution, sélectionnées pour l'augmentation de la qualité visuelle qu'elles procurent. Une image peut
ainsi être formée soit par filtrage des rayons incidents (diaphragme), soit par réfraction et concentration de la
lumière incidente en certains points (lentille), soit par réflexion des rayons après leur traversée des cellule
réceptrices (miroir).
L'augmentation de la taille de la coupe formée par les cellules photo-réceptrices couplée à la
réduction de l'ouverture à la lumière permet de concentrer les rayons venant d'une même direction sur un
nombre restreint de cellules réceptrices, et permet ainsi à l'œil d'analyser la direction de la lumière et de
former une image. Un tel œil, appelé œil en trou d'épingle (pinhole eye), a évolué au moins une fois chez les
mollusques, et on le retrouve par exemple chez les Haliotis. Chez le Nautile, une forme panchronique de
mollusque céphalopode, l'évolution de muscles permettant de stabiliser l'œil pendant la nage et donc
stabiliser l'image a évolué en parallèle de l'œil en trou d'épingle. Cependant, ces yeux possèdent des
limitations physiques.
En effet, l'augmentation de la résolution nécessite une réduction de l'ouverture donc une diminution
de quantité de lumière perçue (sensibilité). Réciproquement, une augmentation de la sensibilité passe par
l'augmentation de l'ouverture, et l'image devient alors plus floue. Il semblerait que le plus simple système
optique placé au niveau de l'ouverture ait suffit à augmenter les qualités optiques d'un tel œil. Compte tenu
du grand nombre d'apparitions convergentes de lentilles dans les yeux animaux, on ne sait toujours pas
pourquoi une telle solution n'a pas évolué dans le cas de cet œil.
L'ajout d'un système réfractif (lentille) à un œil-puits ancestral permet de concentrer les rayons issus
d'une même direction vers une zone limitée de la couche réceptrice, ce qui permet la formation d'une image.
Les premières lentilles apparues sont homogènes et de forme sphérique. De tels systèmes réfractifs ont
évolué au moins 8 fois de manière indépendante : chez les Céphalopodes, chez les Vertébrés, chez les
Copépodes, et 4 fois indépendantes chez les Gastéropodes[4]. Il existe aujourd'hui une grande diversité de
structure et de composition des lentilles chez les animaux, qui sont toutes issues de lentilles sphériques
homogènes.
Chez les animaux aquatiques, des lentilles possédant un gradient d'indice de réfraction se sont
développées à plusieurs reprises, dans plusieurs embranchements distincts. Ces lentilles sphériques,
appelées lentilles de Matthiessen, ont une densité qui décroît en s'éloignant du centre, ce qui a pour
conséquence la formation d'un gradient décroissant d'indice de réfraction du centre vers la périphérie. La
lumière est ainsi déviée continument en traversant la lentille, et non pas seulement à l'entrée de la lentille
comme c'est le cas lorsque la lentille est homogène. Dans le cas d'un gradient de densité particulier, proche
d'un gradient parabolique, l'image ainsi formée est corrigée de toute aberration sphérique. Le fait que de
telles lentilles aient évolué presque systématiquement chez les animaux aquatiques suggère qu'il n'existe
qu'une solution pour corriger l'aberration sphérique en milieu marin, et que cette solution a été sélectionnée à
chaque fois par l'évolution.
D'autres solutions, bien moins répandues, ont néanmoins évolué chez certains animaux marins. Elle
impliquent en général plusieurs lentilles issues de lentilles sphériques homogènes. Le Crustacé Pontella, par
exemple, a développé un système de 3 lentilles superposées, de formes différentes, qui permettent la
correction de l'aberration sphérique[19].
L'œil de certains animaux terrestres, tels les Mammifères et les Araignées, possèdent un système
réfractif supplémentaire placé devant la lentille : la cornée. Celle-ci possède alors l'essentiel du pouvoir de
réfraction de l'œil (les 2/3 chez l'homme), la lentille jouant essentiellement une rôle dans l'ajustement du
point focal (accommodation). L'évolution d'un œil à cornée à partir d'un œil ne possédant qu'une lentille a
probablement nécessité une diminution du pouvoir de réfraction de la lentille alors que celui de la cornée
augmentait parallèlement. En effet, une telle transition évolutive ne passe par aucune phase contre
sélectionnée, qui empêcherait la formation d'un œil à lentille.
La troisième solution évolutive permettant la formation d'une image consiste à placer derrière la
couche de cellules réceptrices une couche cellulaire réfléchissante, agissant comme un miroir. La forme
sphérique de ce miroir permet là encore de concentrer les rayons lumineux issus d'une direction sur une
zone réceptrice limitée, et donc de former une image. Ce miroir est apparu de manière convergente sous
forme d'une multi-couche réfléchissante, chez les Rotifères, les Plathelminthes, les Copépodes et certains
Mollusques.
 Les yeux composés [modifier]
Les yeux composés sont répandus dans le règne animal, et présentent une importante diversité. On
distingue deux grands types de vision. La vision par apposition ... (focale et afocale). La vision par
superposition ... (neurale, réfractive, réflective ou parabolique).
Les yeux composés ont probablement évolué plusieurs fois de manière indépendante chez les
Annélides, les Arthropodes et chez les Mollusques bivalves de la famille des Arcacae. Ces yeux sont à
chaque fois issus d'un œil-puits, à partir duquel se forme un œil composé peu complexe. Cet œil ancestral
est formé de cellules réceptrices placées au fond d'un certain nombre de tubes constitués de cellules
pigmentées. On retrouve encore cette forme ancestrale chez certaines espèces contemporaines
d'Annélides. Cet œil composé aurait alors évolué à chaque fois en un œil capable d'une vision par apposition
focale.
L'évolution à partir de cet œil composé ancestral est encore mal connue. Chez les Arthropodes, on
distingue trois groupes principaux possédant des yeux à vision par apposition focale, et l'origine évolutive de
la vision chez ces groupes reste discutée[20] ' [21]. La vision par apposition afocale aurait évolué à partir de
la vision par apposition focale. Il semblerait par ailleurs que la vision par superposition neurale ait évolué
plusieurs fois à partir de cette dernière. Enfin, la vision par superposition parabolique n'a été découverte
récemment et reste mal comprise. Son étude pourrait cependant apporter d'importantes informations
évolutives, puisque l'œil impliqué combine des structures trouvées dans les yeux à visions par apposition et
par superposition.
Un modèle d'évolution de l'œil [modifier]
 Les différentes étapes de la formation d'un œil selon le modèle de Nilsson et Pelger. Le nombre de
modifications élémentaires de 1% nécessaire pour passer d'une structure à l'autre est indiqué à droite des
flèches. Selon cette séquence évolutive, un œil de structure semblable à celui de l'homme peut être formé en
moins de 400 000 générations. En bleu clair: couche protectrice transparente. En bleu foncé: lentille à
gradient d'index réfractif formé par une augmentation locale de la concentration en protéines dans la couche
protectrice. En violet: couche photoréceptrice. En noir: couche pigmentée noire.
La formation de structures complexes par le seul moyen de la sélection naturelle a souvent été mise
en doute par les opposants de la théorie de l'évolution, et l'évolution de l'œil est souvent l'exemple
emblématique de ce problème. Charles Darwin lui-même reconnaissait la difficulté que posait l'évolution de
l'œil pour sa théorie. La difficulté pour expliquer l'évolution de l'œil par la sélection naturelle consiste à
trouver une voie évolutive plausible constituée d'une longue succession de légères modifications, chacune
d'entre elle devant être avantageuse et donc sélectionnée.
Afin de montrer qu'une telle évolution était possible, et d'estimer le nombre de générations
nécessaires à l'évolution d'un œil, Dan Nilsson et Susanne Pelger ont développé en 1994 un modèle
d'évolution de l'œil[22]. Pour cela, ils ont imaginé une séquence possible de 8 structures d'œil allant d'une
simple couche de cellules photoréceptrices à un œil camérulaire tel qu'on l'observe chez les céphalopodes
ou les vertébrés, à partir des lois fondamentales de l'optique, des lois de la sélection naturelle et des
exemples réellement observés dans la nature. Ils ont ensuite calculé le nombre de modifications
élémentaires nécessaires au passage d'une structure à l'autre. Enfin, ils déduisent de ce calcul le nombre de
générations nécessaire à l'évolution d'un œil selon cette séquence et d'après la théorie de la sélection
naturelle. En construisant ce modèle pessimiste (la séquence évolutive et toutes les hypothèses de calcul
sont choisies de manière à maximiser le nombre de générations), ils estiment qu'il ne faut pas plus de
quelques centaines de milliers de générations pour former un œil camérulaire.

Séquence évolutive [modifier]

La principale difficulté pour estimer le nombre de générations nécessaires à l'évolution d'un œil est le
choix de la séquence évolutive suivie. Pour cela, les auteurs ont choisi une séquence de structures
cohérentes avec les observations réelles et les apports de l'anatomie comparative, mais ont cherché des
structures les plus générales possibles, en essayant d'éviter des structures spécifiques à un taxon particulier.
Les auteurs se sont ensuite fondés sur les lois de la sélection naturelle, et ont cherché une fonction sur
laquelle la sélection pourrait s'exercer tout au long de la séquence évolutive. Ils ont ainsi choisi de se
concentrer sur la résolution spatiale de l'œil. Ce choix permet d'évaluer de manière objective la qualité de
chaque structure oculaire, simplement par l'application des lois de l'optique.
La première structure a été choisie comme une lame circulaire de cellules photoréceptrices, coincée
en sandwich entre une lame de cellules pigmentées noires au-dessous et une couche transparente
protectrice au-dessus. Cette première structure, plausible compte tenu de ce que l'on observer dans la
nature, permet de s'affranchir du problème de l'évolution des cellules photoréceptrices, et de ne se consacrer
qu'à l'évolution de la structure de l'œil.
Deux types de transformations permettent alors d'améliorer la résolution de cette structure
trilaminaire : la courbure de la structure, orientant chaque cellule dans une direction particulière (les
différentes cellules sont alors sensibles à des directions différentes), et l'obstruction de l'ouverture par des
cellules pigmentées noires. Afin de déterminer dans quel ordre ces transformations ont pu intervenir au cours
de l'évolution, Nilsson et Pelger ont étudié l'influence de transformations élémentaires successives des deux
types (courbure ou obstruction de l'ouverture) sur la résolution optique. Ils montrent ainsi que la courbure
apporte une meilleure amélioration tant que l'œil est moins profond que large.
Par ailleurs, l'obstruction de l'ouverture de l'œil s'accompagne aussi d'une diminution de la quantité
de lumière admise. Les cellules photoréceptrices ne réagissant qu'à partir d'une intensité lumineuse donnée,
une ouverture trop petite diminue aussi les qualités optiques de la structure. Ainsi, à partir de la théorie de
Snyder et Warrant et McIntyre[23], les auteurs montrent qu'il existe une obstruction optimale de l'ouverture
de l'œil pour laquelle la résolution obtenue est maximale (pour une intensité extérieure et un diamètre d'œil
donné).
Enfin, ils montrent que même une petite augmentation locale d'indice de réfraction du matériau
protecteur peut jouer le rôle d'une lentille et améliorer la résolution, et que l'augmentation du pouvoir
réfractoire est toujours avantageuse jusqu'à l'obtention d'une lentille optimale d'indice de réfraction de 1,52.
De plus, la position optimale de la lentille change en fonction de l'évolution de la forme de l'œil, et la position
de la lentille est amenée à évoluer en conséquence.
À partir de ces propriétés sur les mécanismes de la sélection naturelle et de la formation d'une
image, Nilsson et Pelger proposent une séquence évolutive de 8 étapes présentée ci-dessous. Lors des
étapes 2 et 3, la structure trilaminaire est courbée jusqu'à prendre la forme d'un hémisphère, rempli par la
couche protectrice. Les couches photosensible et pigmentée s'agrandissent ensuite lors des étapes 4 et 5
sans que le rayon de courbure de la structure ne change, entraînant ainsi une diminution de la taille du
diaphragme. Une lentille est ensuite formée par une augmentation locale de la concentration en protéines
dans la couche protectrice (causant une augmentation de l'indice de réfraction). Enfin, la lentille continue de
se former lors des étapes 7 et 8 alors que la taille de l'ouverture diminue grâce à une croissance de la
couche pigmentée et que la forme de l'œil change. Cette séquence aboutit à un œil camérulaire comme on
peut l'observer chez les Vertébrés et les Céphalopodes.
 Nombre d'étapes élémentaires nécessaires [modifier]
Les auteurs calculent ensuite le nombre de modifications élémentaires de 1% nécessaires à passer
d'une structure à l'autre. Ces modifications de 1% peuvent affecter n'importe quel caractère quantitatif,
comme la longueur d'une structure ou la concentration en protéines responsable du changement d'indice de

réfraction. Ainsi, il faut 70 étapes de 1% pour qu'une structure double de taille, puisque . Ainsi,
la formation de l'œil par cette séquence évolutive nécessite au total 1829 modifications de 1%. Par ailleurs,
Nilsson et Pelger montrent que la qualité optique de chaque structure est proportionnelle au nombre d'étapes
de 1% nécessaire pour former la structure. Ainsi, il n'existe pas d'étape limitante (c'est-à-dire d'étape
nécessitant beaucoup de modifications élémentaires pour une faible augmentation de la résolution spatiale)
au cours de leur séquence.

Nombre de générations nécessaires [modifier]

Nilsson et Pelger utilisent alors la théorie de la sélection naturelle pour calculer le nombre de
générations nécessaires à l'évolution d'un œil par cette séquence évolutive. Pour cela, ils se servent de
l'équation développée par Falconer en 1989, donnant la relation entre la valeur d'un caractère à une
génération donnée, et celle de ce même caractère à la génération suivante: R = h2iVm, où m est la valeur
moyenne du caractère à la génération n, R la valeur moyenne du caractère à la génération n+1, h
l'héritabilité du caractère, i l'intensité de la sélection sur ce caractère et V le coefficient de variation du
caractère.
 Les auteurs ont choisi h2 = 0,50, et des valeurs particulièrement faibles pour le coefficient de
variation et l'intensité de sélection afin de maximiser le nombre de générations nécessaires (i=0,01 et V =
0,01). Ainsi, selon leur modèle, R = 0,00005m, c'est-à-dire que le caractère change de 0,005% par
génération. Ainsi, si n est le nombre de générations nécessaires à former l'œil de l'étape 8, il faut que:
1,011829 = 1,00005n, soit n = 363992.

Critiques [modifier]

Conclusion [modifier]
Ainsi, selon leur modèle volontairement pessimiste, Nilsson et Pelger estiment qu'il faut moins de
400 000 générations pour former un œil camérulaire comme celui des Vertébrés. Les premiers fossiles
possédant des yeux datant de 550 millions d'années, l'œil camérulaire a largement eu le temps d'évoluer
depuis cette époque. Cependant, l'œil ne peut évoluer comme une structure autonome, et nécessite la
présence d'autres structures pour être fonctionnel et apporter un avantage sélectif. Ainsi, le développement
du traitement neural de l'information visuelle a pu être un facteur plus limitant que l'évolution de la structure
oculaire elle-même. Cette estimation montre ainsi que la structure de l'œil nécessite relativement peu de
temps pour évoluer et qu'elle peut répondre rapidement à des modifications des pressions sélectives
exercées par le milieu.

Controverses sur l'évolution de l'œil [modifier]

Plusieurs opposants à la théorie de l'évolution darwinienne, notamment Henri Bergson dans
L'évolution créatrice, arguent que certains organes, comme l'œil humain (ou celui du homard), exigent un
agencement très précis et concourant de différents éléments pour fonctionner correctement. Ils ne pourraient
donc être le résultat d'une évolution progressive par sélection naturelle : une ébauche d'œil ne fonctionnerait
pas et ne donnerait pas un avantage sélectif significatif. Arthur Koestler estime même, dans Janus, que cette
mutation inutile aurait de fortes chances de se diluer et de disparaitre bien avant que les suivantes n'arrivent
pour la compléter.
Bergson s'oppose à Darwin sur un point : la réductibilité du phénomène vivant à une explication
mécaniste, surtout pour des raisons épistémologiques liées à la fonction de notre intelligence ; celle-ci, dans
une perspective pragmatique, vise l'action et constitue elle-même un outil d'adaptation. Le modèle établi par
Darwin, selon lui conséquence de notre faculté de connaître, resterait encore insuffisant et exigerait une
explication philosophique et intuitive de l'évolution. L'existence même d'une évolution n'est jamais remise en
cause par Bergson ni par Koestler, mais tous deux insistent sur le fait que si son rôle d'élimination est
indiscutable, son apport explicatif à des successions d'innovations en cascade reste problématique même
sur les durées considérées, essentiellement pour des raisons de dilution de mutations qui, isolées,
resteraient sans effet.
En fait, un organe photosensible même élémentaire, capable par exemple de simplement distinguer
le jour de la nuit ou le passage d'une ombre, d'un prédateur ou d'un proie potentielle, procure un avantage
sélectif suffisant[24] pour qu'on puisse envisager qu'il s'impose rapidement. De là à obtenir un organe
beaucoup plus complexe tel que l'œil des vertébrés, le nombre d'étapes est certes considérable et demande
une coordination des modifications de plusieurs organes adjacents, mais chaque amélioration accentuera
l'avantage sélectif et s'imposera.
Des travaux montrent que l'apparition d'un œil complexe peut être en fait très rapide. En effet, Nilssion et
Pelger montrent en 1994 que, sous des hypothèses particulièrement pessimistes, 400 000 générations
suffisent à former un œil camérulaire (comme celui des Vertébrés) selon les modèles prédits par la théorie
de l'évolution[25].
Le fait que la rétine des céphalopodes et celle des vertébrés ne soit pas « montée » dans le même
sens montre que les yeux de ces deux groupes ne sont pas homologues et se sont développés par des voies
différentes. Ce montage à l'envers, considéré comme une imperfection chez les vertébrés, est présenté par
Richard Dawkins[26] comme une preuve de l'imperfection de l'œil des vertébrés, et donc du fait que ce
dernier ne résulte pas de l'application d'un plan préétabli mais au contraire de d'adaptations successives,
comme un argument contre le créationnisme et l'Intelligent Design.
Certaines considérations pourraient montrer ce montage inverse de la rétine des vertébrés comme une
imperfection : existence d'un point aveugle, rétine moins attachée au fond de l'œil (?), lumière filtrée par la
rétine avant les récepteurs...
Néanmoins, certaines de ces caractéristiques « néfastes » ont été minimisés : comme le point aveugle dont
l'étendue a été réduite, une liaison plus forte avec le fond de l'œil (?) Ou bien même, peut être utilisée(s)
comme avantage(s) dans certaines circonstances : comme le pouvoir filtrant de la couche rétinienne.

Notes et références [modifier]

1. ↑ a et b M.F. Land & D.E. Nilsson, Animal Eyes, Oxford University Press, 2002
2. ↑ W.J. Gehring & K. Ikeo, « Pax6: Mastering eye morphogenesis and eye evolution », dans
Trends in Genetics, vol. 15, 1999, p. 371-377
3. ↑ (en) J.N. Lythgoe, The Ecology of Vision, Oxford: Clarendon Press, 1979
4. ↑ a et b M.F. Land et R.D. Fernald, « The Evolution of Eyes », dans Annual Review of
Neurosciences, vol. 15, 1992, p. 1-29
 5. ↑ R.D. Fernald, « Evolution of Eyes », dans Current Opinion in Neurobiology, vol. 10, 2000,
p. 444-450
6. ↑ A. Packard, « Cephalopods and fish: the limits of convergence », dans Bio. Rev., vol. 47,
1972, p. 241-307
7. ↑ R.D. Fernald, « Evolving eyes », dans Int. J. Dev. Biol., vol. 48, 2004, p. 701-705
8. ↑ L.V. Salvini-Plawen & E. Mayr, « Evolution of photoreceptors and eyes », dans Evol. Biol,
vol. 10, 1977, p. 207-263
9. ↑ S. Yokoyama, « Molecular Évolution Of Vertebrate Visual Pigments », dans Progress in
Retinal and Eye Research, vol. 19, no 4, 2000, p. 385-419
10.↑ S. Yokoyama, « Evolution of Dim-Ligth and Color Vision Pigments », dans Annual Review
of Genomics and Human Genetics, vol. 9, 2 Juin 2008, p. 259-282
11.↑ A.G. Palacios, F.J. Parela, R. Srivastava & T.J. Goldsmith, « Spectral sensitivity of cones
in the goldfish, Carassius auratus », dans Vision Research, vol. 38, 1998, p. 2135-2146
12.↑ J.K. Bowmaker & A. Knowles, « The visual pigments and oil droplets of the chicken
retina », dans Vision Research, vol. 17, 1977, p. 755-764
13.↑ Seki T. & Vogt K., « Evolutionnary aspects of of the diversity of visual pigment
chromophores in the class Insecta », dans Comp. Biochem. Physiol. B, vol. 119, 1998, p. 53-64
14.↑ (en) Adriana D. Briscoe et Chittka, « The evolution of color vision in inects », dans
Annual Review of Entomology, vol. 46, 2001, p. 471-510 (ISSN 0066-4170) [ résumé [archive] ]
15.↑ a et b D. Arendt, « Evolution of eyes and photoreceptor cell types », dans International
Journal of Developmental Biology, vol. 47, 2003, p. 563-571
16.↑ K.W. Yau & D.A. Baylor, « Cyclic GMP-activated conductance of retinal photoreceptor
cells », dans Annu. Rev. Neurosci., vol. 12, 1989, p. 289-327
 17.↑ M.G.F. Fuortes œ P.M. O'Bryan, « Generators potentials in invertebrate photoreceptors »,
dans Handbook of Sensory Physiology, 1972, p. 321-338
18.↑ M.F. Land, « Functional aspects of the optical and retinal organization of the mollusc
eye », dans Symp. Zool. Soc. London, vol. 23, 1968, p. 75-96
19.↑ (en) M.A. Ali, Photoreception and Vision in Invertebrates, New York: Plenum, 1984,
p. 401-438
20.↑ (en) S.M. Manton & D.T. Anderson, The Origin of Major Invertebrate Groups, London:
Academic, 1979, « Polyphyly and the evolution of arthropods », p. 269-321
21.↑ (en) H.F. Paulus, Arthropod phylogeny, New York: Van Nostrand Reinhold, 1979, « Eye
structure and the monophyly of arthropoda », p. 299-383
22.↑ (en) D.E. Nilsson et S. Pelger, « A pessimistic estimate of the time required for an eye to
evolve », dans Proc. R. Soc. Lond. B, vol. 256, 1994, p. 53-58
23.↑ (en) E.J. Warrant et P.D. McIntyre, « Arthropod eye design and the pysical limits to
spatial resolving power », dans Progresss in Neurobiology, vol. 40, no 4, 1993, p. 413-461
(ISSN 0301-0082)
24.↑ Dans certaines circonstances, comme celles du Cambrien.
25.↑ Nilsson & Pelger, A pessimistic estimate of the time required for an eye to evolve.Proc.
Biol. Sci .1994 Apr 22;256(1345):53-8
26.↑ L'Horloger Aveugle, 1986.
 Voir aussi [modifier]
Articles connexes [modifier]
• Théorie synthétique de l'évolution
• Œil
• Vision des oiseaux
• Vision de l'insecte

Lien externe [modifier]

• Sur l'évolution de la vision chez les Vertébrés.

[Dérouler]
v·d·m
Œil
Rétine · Cornée · Iris · Pupille · Cristallin · Macula · Sclère · Fovéa · Point aveugle · Corps vitré - ·
Humeur aqueuse · Liquide lacrymal · Choroïde · Corps ciliaire · Conjonctive · Structure d'angle · Tapetum
lucidum
Système sensoriel · Système visuel · Vision · Œil humain · Ophtalmologie · Histoire évolutive de l'œil
 [Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

 Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_de_l%27%C5%93il
».
Catégorie : Histoire évolutive | [+]
 W000

Histoire évolutive des dents

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Les dents sont des organes minéralisés extrêmement durs situés à l'entrée de l'appareil digestif de
nombreux vertébrés. Elles sont apparues chez l'ancêtre des Gnathostomes (Vertébrés à mâchoire) il y a
environ 450 millions d'années. Elles sont probablement issues des odontodes, structures dermo-
épidermiques en forme de tubercules coniques qui étaient disposés régulièrement sur toute la surface des
Placodermes, groupe frère des Gnathostomes. Les dents ont ensuite évolué de manière très différente dans
les différentes lignées de Vertébrés. Ainsi, on peut observer une différenciation des dents des Mammifères
en types dentaires très distincts, ou même une perte complète des dents comme chez les Oiseaux.
Sommaire
[masquer]
• 1 L'apparition des dents chez les Gnathostomes
• 2 Évolution des dents chez les Chondrichtyens
• 3 Évolution des dents chez les Sarcoptérygiens
• 4 Évolution des dents chez les Vertébrés Tétrapodes
• 4.1 Évolution des dents chez les Sauropsides
• 4.1.1 La perte des dents chez les Oiseaux
• 4.2 Évolution des dents chez les Mammifères
• 4.2.1 Régionalisation des dents
• 4.2.2 Évolution de la forme des dents
• 4.2.3 Évolution de la formule dentaire
• 5 Notes et références
• 6 Voir aussi
• 6.1 Articles connexes
• 6.2 Liens externes
• 6.3 Bibliographie
L'apparition des dents chez les Gnathostomes [modifier]
Cette section est vide, pas assez détaillée ou incomplète. Votre aide est la bienvenue !

Évolution des dents chez les Chondrichtyens [modifier]

Cette section est vide, pas assez détaillée ou incomplète. Votre aide est la bienvenue !

Évolution des dents chez les Sarcoptérygiens [modifier]

Cette section est vide, pas assez détaillée ou incomplète. Votre aide est la bienvenue !

Évolution des dents chez les Vertébrés Tétrapodes [modifier]

Cette section est vide, pas assez détaillée ou incomplète. Votre aide est la bienvenue !

Évolution des dents chez les Sauropsides [modifier]

Cette section est vide, pas assez détaillée ou incomplète. Votre aide est la bienvenue !

La perte des dents chez les Oiseaux [modifier]

Cette section est vide, pas assez détaillée ou incomplète. Votre aide est la bienvenue !
Évolution des dents chez les Mammifères [modifier]

Régionalisation des dents [modifier]

Les premiers Mammifères Synapsides possédaient une denture isomorphe, c'est-à-dire que toutes
les dents de la mâchoire avaient la même forme, ainsi que le montrent les fossiles de Pélycosaures
(Synapsides non Thérapsides) comme ceux de Dimétrodon. On observe déjà un début de régionalisation
chez les premiers Thérapsides, chez lesquels on distingue des incisives, des canines et des post-canines. Il
existe de plus une dent particulière située en avant de la canine, appelée pré-canine. L'apparition de
l'hétérodontie, c'est-à-dire de la différenciation de plusieurs types dentaires, a donc eu lieu chez l'ancêtre des
Thérapsides. Chez les Mammaliaformes, la régionalisation continue, et les molaires et les prémolaires se
différencient à partir des post-canines. Par la suite, cette régionalisation s'accentue, et une différenciation
des dents d'un même type dentaire se développe souvent, mais aucun nouveau type dentaire n'apparaît. On
observe cependant plusieurs fois la perte d'un ou plusieurs types dentaires, comme chez les Monotrèmes,
dépourvus d'incisives et de canines.
Évolution de la forme des dents [modifier]

Évolution de la formule dentaire [modifier]

 Arbre phylogénétique des Mammifères Cynodontes[1] montrant l'évolution de la formule dentaire au
cours du Mésozoïque. Légende des noeuds (losanges): 1-Cynodontia; 2-Mammaliforms; 3-Crown
Mammalia; 4-Monotrèmes; 5-Eutrichonodontids; 6-Trechnotherians; 7-Sphalacotheroids; 8-Crown Therians.
On observe une tendance générale à la réduction du nombre de dent chez les Mammifères. Les
dents jugales (molaires et prémolaires) sont le plus souvent perdues, mais le nombre d'incisive est souvent
réduit lui-aussi, et la perte des canines a eu lieu plusieurs fois indépendamment. Cette réduction ne
correspond pourtant pas à une même pression de sélection exercée sur l'ensemble des Mammifères, et
aucun mécanisme global de cette réduction n'a été mis en évidence. Cette réduction semble être, dans
chaque cas, une adaptation au régime alimentaire. Les mécanismes développementaux à l'origine de ces
pertes de dents ne sont pas élucidés.

Notes et références [modifier]

1. ↑ (en) Z.X. Luo, P. Chen, G. Li et M. Chen, « A new eutriconodont mammal and
evolutionary development in early mammals », dans Nature, vol. 446, 2007, p. 288-293 [ lien
DOI [archive] ]

Voir aussi [modifier]

Articles connexes [modifier]
• Dent animale
• Dent
 • Dentition
• Évolution

Liens externes [modifier]

Bibliographie [modifier]
[Enrouler]
v·d·m
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive
 Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie
l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation
 • Portail de l’origine et de l’évolution du vivant
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_dents ».

Catégories : Histoire évolutive | Odonto-stomatologie | Dent | [+]

 W000

Evolutionary developmental biology

From Wikipedia, the free encyclopedia

Jump to: navigation, search

Part of the Biology series on

Evolution
 Mechanisms and processes

Adaptation
Genetic drift
Gene flow
Mutation
Natural selection
Speciation

Research and history

Introduction
Evidence
Evolutionary history of life
History
Level of support
Modern synthesis
Objections / Controversy
Social effect
Theory and fact

Evolutionary biology fields

Cladistics
Ecological genetics
Evolutionary development
Evolutionary psychology
Molecular evolution
Phylogenetics
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Biology portal · v • d • e
Evolutionary developmental biology (evolution of development or informally, evo-devo) is a field of
biology that compares the developmental processes of different organisms to determine the ancestral
relationship between them, and to discover how developmental processes evolved. It addresses the origin
and evolution of embryonic development; how modifications of development and developmental processes
lead to the production of novel features, such as the evolution of feathers;[1] the role of developmental
plasticity in evolution; how ecology impacts in development and evolutionary change; and the developmental
basis of homoplasy and homology.[2]
Although interest in the relationship between ontogeny and phylogeny extends back to the nineteenth
century, the contemporary field of evo-devo has gained impetus from the discovery of genes regulating
embryonic development in model organisms. General hypotheses remain hard to test because organisms
differ so much in shape and form.[3] Nevertheless, it now appears that just as evolution tends to create new
genes from parts of old genes (molecular economy), evo-devo demonstrates that evolution alters
developmental processes (genes and gene networks) to create new and novel structures from the old gene
networks (such as bone structures of the jaw deviating to the ossicles of the middle ear) or will conserve
(molecular economy) a similar program in a host of organisms such as eye development genes in molluscs,
insects, and vertebrates [4] [5]. Initially the major interest has been in the evidence of homology in the cellular
and molecular mechanisms that regulate body plan and organ development. However more modern
approaches include developmental changes associated with speciation.[6]
Contents
[hide]
• 1 Basic principles
• 2 History
• 3 The developmental-genetic toolkit
• 4 Development and the origin of novelty
• 5 See also
• 6 References
• 6.1 Sources
• 7 Further reading
• 8 External links

[edit] Basic principles

Charles Darwin's theory of evolution is based on three principles: natural selection, heredity, and
variation. At the time that Darwin wrote, the principles underlying heredity and variation were poorly
understood. In the 1940s, however, biologists incorporated Gregor Mendel's principles of genetics to explain
both, resulting in the modern synthesis. It was not until the 1980s and 1990s, however, when more
comparative molecular sequence data between different kinds of organisms was amassed and detailed, that
an understanding of the molecular basis of the developmental mechanisms has arisen. Currently, it is well
understood how genetic mutation occurs. However, developmental mechanisms are not understood
sufficiently to explain which kinds of phenotypic variation can arise in each generation from variation at the
genetic level. Evolutionary developmental biology studies how the dynamics of development determine the
phenotypic variation arising from genetic variation and how that affects phenotypic evolution (specially its
direction). At the same time evolutionary developmental biology also studies how development itself evolves.
Thus, the origins of evolutionary developmental biology come from both an improvement in molecular biology
techniques as applied to development, and the full appreciation of the limitations of classic neo-Darwinism as
applied to phenotypic evolution. Some evo-devo researchers see themselves as extending and enhancing
the modern synthesis by incorporating into it findings of molecular genetics and developmental biology.
Others, drawing on findings of discordances between genotype and phenotype and epigenetic mechanisms
of development, are mounting an explicit challenge to neo-Darwinism.
Evolutionary developmental biology is not yet a unified discipline, but can be distinguished from
earlier approaches to evolutionary theory by its focus on a few crucial ideas. One of these is modularity: as
has been long recognized, plants and animal bodies are modular: they are organized into developmentally
and anatomically distinct parts. Often these parts are repeated, such as fingers, ribs, and body segments.
Evo-devo seeks the genetic and evolutionary basis for the division of the embryo into distinct modules, and
for the partly independent development of such modules.
Another central idea is that some gene products function as switches whereas others act as diffusible
signals. Genes specify proteins, some of which act as structural components of cells and others as enzymes
that regulate various biochemical pathways within an organism. Most biologists working within the modern
synthesis assumed that an organism is a straightforward reflection of its component genes. The modification
of existing, or evolution of new, biochemical pathways (and, ultimately, the evolution of new species of
organisms) depended on specific genetic mutations. In 1961, however, Jacques Monod, Jean-Pierre
Changeux and François Jacob discovered within the bacterium Escherichia coli a gene that functioned only
when "switched on" by an environmental stimulus.[7] Later, scientists discovered specific genes in animals,
including a subgroup of the genes which contain the homeobox DNA motif, called Hox genes, that act as
switches for other genes, and could be induced by other gene products, morphogens, that act analogously to
the external stimuli in bacteria. These discoveries drew biologists' attention to the fact that genes can be
selectively turned on and off, rather than being always active, and that highly disparate organisms (for
example, fruit flies and human beings) may use the same genes for embryogenesis (e.g., the genes of the
"developmental-genetic toolkit", see below), just regulating them differently.
Similarly, organismal form can be influenced by mutations in promoter regions of genes, those DNA
sequences at which the products of some genes bind to and control the activity of the same or other genes,
not only protein-specifying sequences. In addition to providing new support for Darwin's assertion that all
organisms are descended from a common ancestor, this finding suggested that the crucial distinction
between different species (even different orders or phyla) may be due less to differences in their content of
gene products than to differences in spatial and temporal expression of conserved genes. The implication
that large evolutionary changes in body morphology are associated with changes in gene regulation, rather
than the evolution of new genes, suggested that the action of natural selection on promoters responsive to
Hox and other "switch" genes may play a major role in evolution.
Another focus of evo-devo is developmental plasticity, the basis of the recognition that organismal
phenotypes are not uniquely determined by their genotypes. If generation of phenotypes is conditional, and
dependent on external or environmental inputs, evolution can proceed by a "phenotype-first" route,[3][8] with
genetic change following, rather than initiating, the formation of morphological and other phenotypic
novelties. The case for this was argued for by Mary Jane West-Eberhard in her 2003 book Developmental
plasticity and evolution.[8]
[edit] History
An early version of recapitulation theory, also called the biogenetic law or embryological parallelism,
was put forward by Étienne Serres in 1824–26 as what became known as the "Meckel-Serres Law" which
attempted to provide a link between comparative embryology and a "pattern of unification" in the organic
world. It was supported by Étienne Geoffroy Saint-Hilaire as part of his ideas of idealism, and became a
prominent part of his version of Lamarckism leading to disagreements with Georges Cuvier. It was widely
supported in the Edinburgh and London schools of higher anatomy around 1830, notably by Robert Edmond
Grant, but was opposed by Karl Ernst von Baer's embryology of divergence in which embryonic parallels only
applied to early stages where the embryo took a general form, after which more specialised forms diverged
from this shared unity in a branching pattern. The anatomist Richard Owen used this to support his idealist
concept of species as showing the unrolling of a divine plan from an archetype, and in the 1830s attacked the
transmutation of species proposed by Lamarck, Geoffroy and Grant.[9] In the 1850s Owen began to support
an evolutionary view that the history of life was the gradual unfolding of a teleological divine plan,[10] in a
continuous "ordained becoming", with new species appearing by natural birth.[11]
In On the Origin of Species (1859), Charles Darwin proposed evolution through natural selection, a
theory central to modern biology. Darwin recognised the importance of embryonic development in the
understanding of evolution, and the way in which von Baer's branching pattern matched his own idea of
descent with modification[12]:
“ We can see why characters derived from the embryo should be of equal
importance with those derived from the adult, for a natural classification of course
includes all ages.[13] ”
 Ernst Haeckel (1866), in his endeavour to produce a synthesis of Darwin's theory with Lamarckism
and Naturphilosophie, proposed that "ontogeny recapitulates phylogeny," that is, the development of the
embryo of every species (ontogeny) fully repeats the evolutionary development of that species (phylogeny),
in Geoffroy's linear model rather than Darwin's idea of branching evolution.[12] Haeckel's concept explained,
for example, why humans, and indeed all vertebrates, have gill slits and tails early in embryonic
development. His theory has since been discredited. However, it served as a backdrop for a renewed interest
in the evolution of development after the modern evolutionary synthesis was established (roughly 1936 to
1947).
Stephen Jay Gould called this approach to explaining evolution as terminal addition; as if every
evolutionary advance was added as new stage by reducing the duration of the older stages. The idea was
based on observations of neoteny.[14] This was extended by the more general idea of heterochrony
(changes in timing of development) as a mechanism for evolutionary change.[15]
D'Arcy Thompson postulated that differential growth rates could produce variations in form in his
1917 book On Growth and Form. He showed the underlying similarities in body plans and how geometric
transformations could be used to explain the variations.
Edward B. Lewis discovered homeotic genes, rooting the emerging discipline of evo-devo in
molecular genetics. In 2000, a special section of the Proceedings of the National Academy of Sciences
(PNAS) was devoted to "evo-devo",[16] and an entire 2005 issue of the Journal of Experimental Zoology Part
B: Molecular and Developmental Evolution was devoted to the key evo-devo topics of evolutionary innovation
and morphological novelty.[17]
[edit] The developmental-genetic toolkit
The developmental-genetic toolkit consists of a small fraction of the genes in an organism's genome
whose products control its development. These genes are highly conserved among Phyla. Differences in
deployment of toolkit genes affect the body plan and the number, identity, and pattern of body parts. The
majority of toolkit genes are components of signaling pathways, and encode for the production of
transcription factors, cell adhesion proteins, cell surface receptor proteins, and secreted morphogens, all of
these participate in defining the fate of undifferentiated cells, generating spatial and temporal patterns, which
in turn form the body plan of the organism. Among the most important of the toolkit genes are those of the
Hox gene cluster, or complex. Hox genes, transcription factors containing the more broadly distributed
homeobox protein-binding DNA motif, function in patterning the body axis. Thus, by combinatorial specifying
the identity of particular body regions, Hox genes determine where limbs and other body segments will grow
in a developing embryo or larva. A paragon of a toolbox gene is Pax6/eyeless, which controls eye formation
in all animals. It has been found to produce eyes in mice and Drosophila, even if mouse Pax6/eyeless was
expressed in Drosophila.[18]
This means that a big part of the morphological evolution undergone by organisms is a product of
variation in the genetic toolkit, either by the genes changing their expression pattern or acquiring new
functions. A good example of the first is the enlargement of the beak in Darwin's Large Ground-finch
(Geospiza magnirostris), in which the gene BMP is responsible for the larger beak of this bird, relative to the
other finches.[19]
The loss of legs in snakes and other squamates is another good example of genes changing their
expression pattern. In this case the gene Distal-less is very under-expressed, or not expressed at all, in the
regions where limbs would form in other tetrapods.[20] This same gene determines the spot pattern in
butterfly wings,[21] which shows that the toolbox genes can change their function.
 Toolbox genes, as well as being highly conserved, also tend to evolve the same function
convergently or in parallel. Classic examples of this are the already mentioned Distal-less gene, which is
responsible for appendage formation in both tetrapods and insects, or, at a finer scale, the generation of wing
patterns in the butterflies Heliconius erato and Heliconius melpomene. These butterflies are Müllerian mimics
whose coloration pattern arose in different evolutionary events, but is controlled by the same genes.[22] The
previous supports Kirschner and Gerhardt's theory of Facilitated Variation, which states that morphological
evolutionary novelty is generated by regulatory changes in various members of a large set of conserved
mechanisms of development and physiology.[23]

[edit] Development and the origin of novelty

Among the more surprising and, perhaps, counterintuitive (from a neo-Darwinian viewpoint) results of
recent research in evolutionary developmental biology is that the diversity of body plans and morphology in
organisms across many phyla are not necessarily reflected in diversity at the level of the sequences of
genes, including those of the developmental genetic toolkit and other genes involved in development.
Indeed, as Gerhart and Kirschner have noted, there is an apparent paradox: "where we most expect to find
variation, we find conservation, a lack of change".[24]
Even within a species, the occurrence of novel forms within a population does not generally correlate
with levels of genetic variation sufficient to account for all morphological diversity. For example, there is
significant variation in limb morphologies amongst salamanders and in differences in segment number in
centipedes, even when the respective genetic variation is low.
A major question then, for evo-devo studies, is: If the morphological novelty we observe at the level
of different clades is not always reflected in the genome, where does it come from? Apart from neo-
Darwinian mechanisms such as mutation, translocation and duplication of genes, novelty may also arise by
mutation-driven changes in gene regulation. The finding that much biodiversity is not due to differences in
genes, but rather to alterations in gene regulation, has introduced an important new element into evolutionary
theory.[25] Diverse organisms may have highly conserved developmental genes, but highly divergent
regulatory mechanisms for these genes. Changes in gene regulation are "second-order" effects of genes,
resulting from the interaction and timing of activity of gene networks, as distinct from the functioning of the
individual genes in the network.
The discovery of the homeotic Hox gene family in vertebrates in the 1980s allowed researchers in
developmental biology to empirically assess the relative roles of gene duplication and gene regulation with
respect to their importance in the evolution of morphological diversity. Several biologists, including Sean B.
Carroll of the University of Wisconsin–Madison suggest that "changes in the cis-regulatory systems of genes"
are more significant than "changes in gene number or protein function".[26] These researchers argue that the
combinatorial nature of transcriptional regulation allows a rich substrate for morphological diversity, since
variations in the level, pattern, or timing of gene expression may provide more variation for natural selection
to act upon than changes in the gene product alone.
Epigenetic alterations of gene regulation or phenotype generation that are subsequently consolidated
by changes at the gene level constitute another class of mechanisms for evolutionary innovation. Epigenetic
changes include modification of the genetic material due to methylation and other reversible chemical
alteration [27], as well as nonprogrammed remolding of the organism by physical and other environmental
effects due to the inherent plasticity of developmental mechanisms.[8] The biologists Stuart A. Newman and
Gerd B. Müller have suggested that organisms early in the history of multicellular life were more susceptible
to this second category of epigenetic determination than are modern organisms, providing a basis for early
macroevolutionary changes.[28]
[edit] See also
Evolutionary biology portal

• Developmental biology
• Animal evolution
• Plant Evolutionary Developmental Biology
• Baldwin effect
• Developmental systems theory
• Evolution of multicellularity
• Genetic assimilation
• Ontogeny
• Ontogeny recapitulates phylogeny
• List of gene families
• Important publications in evolutionary developmental biology
• Evolution and Development Leading journal
• Body plan
• Cell signaling
• Signal transduction
• Cell signaling networks
• Transcription factor
• Enhancer
 • Enhanceosome
• promoter (biology)
• Gene regulatory network

[edit] References
1. ^ Prum, R.O., Brush, A.H. (March 2003). "Which Came First, the Feather or the Bird?".
Scientific American 288 (3): 84–93. doi:10.1038/scientificamerican0303-84. PMID 12616863.
2. ^ Hall, Brian K. (2000). "Evo-devo or devo-evo—does it matter". Evolution & Development 2:
177–178. doi:10.1046/j.1525-142x.2000.00003e.x.
3. ^ a b Palmer, RA (2004). "Symmetry breaking and the evolution of development". Science
306 (5697): 828–833. doi:10.1126/science.1103707. PMID 15514148.
4. ^ Tomarev, Stanislav I.; Callaerts, Patrick; Kos, Lidia; Zinovieva, Rina; Halder, Georg;
Gehring, Walter; Piatigorsky, Joram (1997). "Squid Pax-6 and eye development". Proceedings of the
National Academy of Sciences 94 (6): 2421–2426. doi:10.1073/pnas.94.6.2421.
http://www.pnas.org/content/94/6/2421.full.
5. ^ Pichaud, Franck; Desplan, Claude (August 2002). "Pax genes and eye organogenesis".
Current opinion in genetics and development 12 (4): 430–434. doi:10.1016/S0959-437X(02)00321-0.
PMID 12100888.
6. ^ Pennisi, E (2002). "EVOLUTIONARY BIOLOGY:Evo-Devo Enthusiasts Get Down to
Details". Science 298 (5595): 953–955.. doi:10.1126/science.298.5595.953. PMID 12411686.
7. ^ Monod, J; Changeux, JP; Jacob, F (1963). "Allosteric proteins and cellular control
systems". Journal of Molecular Biology 6: 306–329. doi:10.1016/S0022-2836(63)80091-1.
PMID 13936070.
 8. ^ a b c West-Eberhard, M-J. (2003). Developmental plasticity and evolution. New York:
Oxford University Press. ISBN 978-0-19-512235-0.
9. ^ Desmond 1989, pp. 53–53, 86–88, 337–340
Secord 2003, pp. 252–253
10.^ Bowler 2003, pp. 120–128, 208
Secord 2003, pp. 424, 512
11.^ Desmond & Moore 1991, pp. 490–491
12.^ a b Bowler 2003, p. 170, 190-191
13.^ Darwin, Charles (1859). On the Origin of Species. London: John Murray. pp. 439–430.
ISBN 0801413192.
14.^ Ridley, Mark (2003). Evolution. Wiley-Blackwell. ISBN 978-1-4051-0345-9.
http://www.blackwellpublishing.com/ridley/.
15.^ Gould, Stephen Jay (1977). Ontogeny and Phylogeny. Cambridge, Massachusetts:
Harvard University Press. ISBN 0-674-63940-5.
16.^ Goodman CS and Coughlin BS (Eds). (2000). "Special feature: The evolution of evo-devo
biology". Proceedings of the National Academy of Sciences 97 (9): 4424–4456.
doi:10.1073/pnas.97.9.4424. PMID 10781035. PMC 18255.
http://www.pnas.org/cgi/content/full/97/9/4424.
17.^ Müller GB and Newman SA (Eds.) (2005). "Special issue: Evolutionary Innovation and
Morphological Novelty". Journal of Exp. Zool. Part B: Molecular and Developmental Evolution 304B:
485–631. http://www3.interscience.wiley.com/cgi-bin/jissue/112149101.
18.^ Xu, P.X., Woo, I., Her, H., Beier, D.R., Maas, R.L. (1997). "Mouse Eya homologues of the
Drosophila eyes absent gene require Pax6 for expression in lens and nasal placode". Development
124 (1): 219–231. PMID 9006082.
 19.^ Abzhanov, A.; Protas, M.; Grant, B.R.; Grant, P.R.; Tabin, C.J. (2004). "Bmp4 and
Morphological Variation of Beaks in Darwin's Finches". Science 305 (5689): 1462–1465.
doi:10.1126/science.1098095. PMID 15353802.
20.^ Cohn, M.J.; Tickle, C. (1999). "Developmental basis of limblessness and axial patterning in
snakes.". Nature 399 (6735): 474–479. doi:10.1038/20944. PMID 10365960.
21.^ Beldade, P.; Brakefield, P.M.; Long, A.D. (2002). "Contribution of Distal-less to quantitative
variation in butterfly eyespots". Nature 415 (6869): 315–318. doi:10.1038/415315a. PMID 11797007.
22.^ Baxter, S.W.; Papa, R.; Chamberlain, N.; Humphray, S.J.; Joron, M.; Morrison, C.; ffrench-
Constant, R.H.; McMillan, W.O.; Jiggins, C.D. (2008). "Convergent Evolution in the Genetic Basis of
Mullerian Mimicry in Heliconius Butterflies". Genetics 180 (3): 1567–1577.
doi:10.1534/genetics.107.082982. PMID 18791259.
23.^ Gerhart, John; Kirschner, Marc (2007). "The theory of facilitated variation". Proceedings of
the National Academy of Sciences 104 (suppl1): 8582–8589. doi:10.1073/pnas.0701035104.
PMID 17494755.
24.^ Gerhart, John; Kirschner, Marc (1997). Cells, Embryos and Evolution. Blackwell Science.
ISBN 978-0865425743.
25.^ Carroll, Sean B.; Grenier, Jennifer K.; Weatherbee, Scott D. (2005). From DNA to Diversity:
Molecular Genetics and the Evolution of Animal Design — Second Edition. Blackwell Publishing.
ISBN 1405119500.
26.^ Carroll, Sean B. (2000). "Endless forms: the evolution of gene regulation and morphological
diversity". Cell 101: 577–80. doi:10.1016/S0092-8674(00)80868-5.
27.^ Jablonka, Eva; Lamb, Marion (1995). Epigenetic Inheritance and Evolution: The
Lamarckian Dimension. Oxford, New York: Oxford University Press. ISBN 978-0198540632.
 28.^ Müller, Gerd B. and Newman, Stuart A., ed (2003). Origination of Organismal Form:
Beyond the Gene in Developmental and Evolutionary Biology . MIT Press.

[edit] Sources
• Bowler, Peter J. (2003). Evolution: the history of an idea. Berkeley: University of California
Press. ISBN 0-520-23693-9.
• Desmond, Adrian J. (1989). The politics of evolution: morphology, medicine, and reform in
radical London. Chicago: University of Chicago Press. ISBN 0-226-14374-0.
• Desmond, Adrian J.; Moore, James William (1991). Darwin. London: Michael Joseph.
ISBN 0-7181-3430-3.
• Secord, James A. (2003). Victorian sensation: the extraordinary publication, reception, and
secret authorship of Vestiges of the natural history of creation . Chicago: University of Chicago Press.
ISBN 0-226-74411-6.

[edit] Further reading

• Buss, Leo W. (1987). The Evolution of Individuality. Princeton University Press. ISBN 978-
0691084688.
• Carroll, Sean B. (2005). Endless Forms Most Beautiful: The New Science of Evo Devo and
the Making of the Animal Kingdom. Norton. ISBN 9780393060164.
• Goodwin, Brian (1994). How the Leopard Changed its Spots. Phoenix Giants.
ISBN 9780691088099.
 • Hall, Brian K. & Olsen, Wendy M., ed (2007). Keywords and Concepts in Evolutionary
Developmental Biology. New Delhi, India: Discovery Publishing House. ISBN 978-81-8356-256-0.
• Kirschner, Marc; Gerhart, John (2005). The Plausibility of Life: Resolving Darwin's Dilemma .
Yale University Press. ISBN 9780300108651.
• Laubichler, Manfred D. and Maienschein, Jane, ed (2007). From Embryology to Evo-Devo: A
History of Developmental Evolution. The MIT Press. ISBN 978-0-262-12283-2.
• Minelli, Alessandro (2003). The Development of Animal Form: Ontogeny, Morphology, and
Evolution. Cambridge University Press. ISBN 978-0521808514.
• Orr, H. Allen (2005-10-24). "Turned on: A revolution in the field of evolution?". The New
Yorker. http://www.newyorker.com/critics/books/articles/051024crbo_books1. Discussion of Carroll,
Endless Forms Most Beautiful
• Raff, Rudolf A. (1996). The Shape of Life: Genes, Development, and the Evolution of Animal
Form. The University of Chicago Press. ISBN 978-0226702667.
• Sommer, Ralf J. (2009). "The future of evo–devo: model systems and evolutionary theory".
Nature Reviews Genetics 10 (6): 416–422. doi:10.1038/nrg2567. PMID 19369972.

[edit] External links

• Scott F. Gilbert, The morphogenesis of evolutionary developmental biology
• Tardigrades (water bears) as evo-devo models, a short video from NPR's Science Friday
 [hide]
v•d•e
Basic topics in evolutionary biology

Evidence of common descent

Processes of
evolution Adaptation · Macroevolution · Microevolution · Speciation

Population genetic
mechanisms Genetic drift · Gene flow · Mutation · Natural selection

Evolutionary
developmental
biology (Evo-devo) Canalisation · Inversion · Modularity · Phenotypic plasticity
concepts

Evolution of organs Aging · Cellular · DNA · The Ear · The Eye · Flagella · Flight · Hair ·
and biological processes Human intelligence · Modular · Muticellular · Sex
 Birds · Butterflies · Dinosaurs · Dolphins and whales · Fungi · Horses ·
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History of Charles Darwin · On the Origin of Species ·

evolutionary thought Modern evolutionary synthesis · Gene-centered view of evolution · Life
(classification trees)

Other subfields Ecological genetics · Molecular evolution · Phylogenetics · Systematics

List of evolutionary biology topics · Timeline of evolution

Retrieved from "http://en.wikipedia.org/wiki/Evolutionary_developmental_biology"

Categories: Evolutionary biology | Developmental biology

 W000

Phylogenetic tree
From Wikipedia, the free encyclopedia
(Redirected from Evolutionary tree)
Jump to: navigation, search
"ptree" redirects here. For Patricia tree, see Radix tree.
 Fig. 1: A speculatively rooted tree for rRNA genes
A phylogenetic tree or evolutionary tree is a branching diagram or "tree" showing the inferred
evolutionary relationships among various biological species or other entities based upon similarities and
differences in their physical and/or genetic characteristics. The taxa joined together in the tree are implied to
have descended from a common ancestor. In a rooted phylogenetic tree, each node with descendants
represents the inferred most recent common ancestor of the descendants, and the edge lengths in some
trees may be interpreted as time estimates. Each node is called a taxonomic unit. Internal nodes are
generally called hypothetical taxonomic units (HTUs) as they cannot be directly observed. Trees are useful in
fields of biology such as systematics and comparative phylogenetics.

Contents
[hide]
• 1 History
• 2 Types
• 3 Construction
• 4 Limitations
• 5 See also
• 5.1 The "tree of life"
• 5.2 Fields of study
• 6 References
• 7 Further reading
• 8 External links
• 8.1 Images
• 8.2 General
[edit] History
The idea of a "tree of life" arose from ancient notions of a ladder-like progression from lower to higher
forms of life (such as in the Great Chain of Being). Early representations of branching phylogenetic trees
include a "Paleontological chart" showing the geological relationships among plants and animals in the book
Elementary Geology, by Edward Hitchcock (first edition: 1840).
Charles Darwin (1859) also produced one of the first illustrations and crucially popularized the notion
of an evolutionary "tree" in his seminal book The Origin of Species. Over a century later, evolutionary
biologists still use tree diagrams to depict evolution because such diagrams effectively convey the concept
that speciation occurs through the adaptive and random splitting of lineages. Over time, species classification
has become less static and more dynamic.
[edit] Types

Fig. 1: Unrooted tree of the myosin supergene family[1]

 Fig. 2: A highly resolved, automatically generated Tree Of Life, based on completely sequenced
genomes [2][3].

A phylogenetic tree, showing how Eukaryota and Archaea are more closely related to each other
than to Bacteria, based on Cavalier-Smith's theory of bacterial evolution. (Cf. LUCA, Neomura.)
A rooted phylogenetic tree is a directed tree with a unique node corresponding to the (usually
imputed) most recent common ancestor of all the entities at the leaves of the tree. The most common method
for rooting trees is the use of an uncontroversial outgroup — close enough to allow inference from sequence
or trait data, but far enough to be a clear outgroup.
 Unrooted trees illustrate the relatedness of the leaf nodes without making assumptions about
ancestry at all. While unrooted trees can always be generated from rooted ones by simply omitting the root, a
root cannot be inferred from an unrooted tree without some means of identifying ancestry; this is normally
done by including an outgroup in the input data or introducing additional assumptions about the relative rates
of evolution on each branch, such as an application of the molecular clock hypothesis. Figure 1 depicts an
unrooted phylogenetic tree for myosin, a superfamily of proteins.[4]
Both rooted and unrooted phylogenetic trees can be either bifurcating or multifurcating, and either
labeled or unlabeled. A rooted bifurcating tree has exactly two descendants arising from each interior node
(that is, it forms a binary tree), and an unrooted bifurcating tree takes the form of an unrooted binary tree, a
free tree with exactly three neighbors at each internal node. In contrast, a rooted multifurcating tree may have
more than two children at some nodes and an unrooted multifurcating tree may have more than three
neighbors at some nodes. A labeled tree has specific values assigned to its leaves, while an unlabeled tree,
sometimes called a tree shape, defines a topology only. The number of possible trees for a given number of
leaf nodes depends on the specific type of tree, but there are always more multifurcating than bifurcating
trees, more labeled than unlabeled trees, and more rooted than unrooted trees. The last distinction is the
most biologically relevant; it arises because there are many places on an unrooted tree to put the root. For
labeled bifurcating trees, there are

total rooted trees and

 total unrooted trees, where n represents the number of leaf nodes. Among labeled bifurcating trees,
the number of unrooted trees with n leaves is equal to the number of rooted trees with n − 1 leaves.[5]
A dendrogram is a broad term for the diagrammatic representation of a phylogenetic tree.
A cladogram is a tree formed using cladistic methods. This type of tree only represents a branching
pattern, i.e., its branch lengths do not represent time.
A phylogram is a phylogenetic tree that explicitly represents number of character changes through its
branch lengths.
A chronogram is a phylogenetic tree that explicitly represents evolutionary time through its branch
lengths.

[edit] Construction
Main article: Computational phylogenetics
Phylogenetic trees among a nontrivial number of input sequences are constructed using
computational phylogenetics methods. Distance-matrix methods such as neighbor-joining or UPGMA, which
calculate genetic distance from multiple sequence alignments, are simplest to implement, but do not invoke
an evolutionary model. Many sequence alignment methods such as ClustalW also create trees by using the
simpler algorithms (i.e. those based on distance) of tree construction. Maximum parsimony is another simple
method of estimating phylogenetic trees, but implies an implicit model of evolution (i.e. parsimony). More
advanced methods use the optimality criterion of maximum likelihood, often within a Bayesian Framework,
and apply an explicit model of evolution to phylogenetic tree estimation.[5] Identifying the optimal tree using
many of these techniques is NP-hard[5], so heuristic search and optimization methods are used in
combination with tree-scoring functions to identify a reasonably good tree that fits the data.
Tree-building methods can be assessed on the basis of several criteria:[6]
• efficiency (how long does it take to compute the answer, how much memory does it need?)
• power (does it make good use of the data, or is information being wasted?)
• consistency (will it converge on the same answer repeatedly, if each time given different data
for the same model problem?)
• robustness (does it cope well with violations of the assumptions of the underlying model?)
• falsifiability (does it alert us when it is not good to use, i.e. when assumptions are violated?)
Tree-building techniques have also gained the attention of mathematicians. Trees can also be built
using T-theory.[7]

[edit] Limitations
Although phylogenetic trees produced on the basis of sequenced genes or genomic data in different
species can provide evolutionary insight, they have important limitations. They do not necessarily accurately
represent the species evolutionary history. The data on which they are based is noisy; the analysis can be
confounded by horizontal gene transfer[8], hybridisation between species that were not nearest neighbors on
the tree before hybridisation takes place, convergent evolution, and conserved sequences.
 Also, there are problems in basing the analysis on a single type of character, such as a single gene
or protein or only on morphological analysis, because such trees constructed from another unrelated data
source often differ from the first, and therefore great care is needed in inferring phylogenetic relationships
among species. This is most true of genetic material that is subject to lateral gene transfer and
recombination, where different haplotype blocks can have different histories. In general, the output tree of a
phylogenetic analysis is an estimate of the character's phylogeny (i.e. a gene tree) and not the phylogeny of
the taxa (i.e. species tree) from which these characters were sampled, though ideally, both should be very
close. For this reason, serious phylogenetic studies generally use a combination of genes that come from
different genomic sources (e.g., from mitochondrial or plastid vs. nuclear genomes), or genes that would be
expected to evolve under different selective regimes, so that homoplasy (false homology) would be unlikely
to result from natural selection.
When extinct species are included in a tree, they are terminal nodes, as it is unlikely that they are
direct ancestors of any extant species. Scepticism must apply when extinct species are included in trees that
are wholly or partly based on DNA sequence data, due to the fact that little useful "ancient DNA" is preserved
for longer than 100,000 years, and except in the most unusual circumstances no DNA sequences long
enough for use in phylogenetic analyses have yet been recovered from material over 1 million years old.
In some organisms, endosymbionts have an independent genetic history from the host.[9]
Phylogenetic networks are used when bifurcating trees are not suitable, due to these complications
which suggest a more reticulate evolutionary history of the organisms sampled.
[edit] See also
Evolutionary biology portal

[edit] The "tree of life"

• Evolutionary history of life - An overview of the major time periods of life on earth
• Life - The top level for Wikipedia articles on living species, reflecting a diversity of
classification systems.
• Three-domain system (cell types)
• Wikispecies - An external Wikimedia Foundation project to construct a "tree of life"
appropriate for use by scientists

[edit] Fields of study

• Archaeopteryx
• Cladistics
• Comparative phylogenetics
• Computational phylogenetics
• Evolutionary biology
• Phylogenetics
[edit] References
1. ^ Hodge T, Cope M (1 October 2000). "A myosin family tree". J Cell Sci 113 Pt 19 (19): 3353–
4. PMID 10984423. http://jcs.biologists.org/cgi/content/full/113/19/3353.
2. ^ Letunic, I; Bork, P (2007). "Interactive Tree Of Life (iTOL): an online tool for phylogenetic
tree display and annotation." (Pubmed). Bioinformatics 23(1) (1): 127–8.
doi:10.1093/bioinformatics/btl529. PMID 17050570.
3. ^ Ciccarelli, FD; Doerks, T; Von Mering, C; Creevey, CJ; Snel, B; Bork, P (2006). "Toward
automatic reconstruction of a highly resolved tree of life." (Pubmed). Science 311(5765) (5765):
1283–7. doi:10.1126/science.1123061. PMID 16513982.
4. ^ Maher BA (2002). "Uprooting the Tree of Life". The Scientist 16: 18. http://www.the-
scientist.com/yr2002/sep/research1_020916.html.
5. ^ a b c Felsenstein J. (2004). Inferring Phylogenies Sinauer Associates: Sunderland, MA.
6. ^ Penny, D., Hendy, M. D. & M. A. Steel. 1992. Progress with methods for constructing
evolutionary trees. Trends in Ecology and Evolution 7: 73-79.
7. ^ A. Dress, K. T. Huber, and V. Moulton. 2001. Metric Spaces in Pure and Applied
Mathematics. Documenta Mathematica LSU 2001: 121-139
8. ^ Woese C (2002). "On the evolution of cells". Proc Natl Acad Sci USA 99 (13): 8742–7.
doi:10.1073/pnas.132266999. PMID 12077305.
9. ^ Villarreal, L.P., & Witzany, G. (2010). Viruses are Essential Agents within the Roots and
Stem of the Tree of Life. Journal of Theoretical Biology 262: 698-710.
 [edit] Further reading
• Schuh, R. T. and A. V. Z. Brower. 2009. Biological Systematics: principles and applications
(2nd edn.) ISBN 978-0-8014-4799-0

[edit] External links

Wikimedia Commons has media related to: Phylogenetic tree

[edit] Images
• Phylogenetic Trees Based on 16s rDNA
• Poster-sized tree of life illustration
• A 3D View
• Human Y-Chromosome 2002 Phylogenetic Tree
• In 2003, the Science journal dedicated a special issue to the tree of life, including an online
version of a tree of life.
• iTOL: Interactive Tree Of Life
• Phylogenetic Tree of Artificial Organisms Evolved on Computers
[edit] General
• Discover Life An interactive tree based on the U.S. National Science Foundation's
Assembling the Tree of Life Project
• PhyloCode
• A Multiple Alignment of 139 Myosin Sequences and a Phylogenetic Tree
• Tree of Life Web Project
• Aisee.com, detailed and comprehensive family tree of dinosaurs yet available
• SplitsTree
• Dendroscope
• Phylogenetic inferring on the T-REX server
• NCBI's Taxonomy Database[1]

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Relevant fields Computational phylogenetics · Molecular phylogeny · Cladistics

Basic concepts Phylogenetic tree · Phylogenetic network · Long branch attraction · Clade ·
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 Inference Maximum parsimony · Maximum likelihood · Neighbor-joining · UPGMA ·
methods Bayesian inference · Least squares

Current topics PhyloCode · DNA barcoding

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 w000

Evolution of mammals
From Wikipedia, the free encyclopedia

Jump to: navigation, search

 Restoration of Thrinaxodon, a member of the cynodont group, which includes the ancestors of
mammals.
Further information: Evolutionary history of life
The evolution of mammals within the synapsid lineage (mammal-like-reptiles) was a gradual process
that took approximately 70 million years, beginning in the mid-Permian. By the mid-Triassic, there were many
species that looked like mammals, and the first true mammals appeared in the early Jurassic. The earliest
known marsupial, Sinodelphys, appeared 125 million years ago in the early Cretaceous, around the same
time as Eomaia, the first known eutherian (member of placentals' "parent" group); and the earliest known
monotreme, Teinolophos, appeared two million years later. After the Cretaceous-Tertiary extinction wiped out
the non-avian dinosaurs (birds are generally regarded as the surviving dinosaurs) and several other
mammalian groups, placental and marsupial mammals diversified into many new forms and ecological
niches throughout the Tertiary, by the end of which all modern orders had appeared.
 From the point of view of phylogenetic nomenclature, mammals are the only surviving synapsids.
The synapsid lineage became distinct from the sauropsid ("reptile") lineage in the late Carboniferous period,
between 320 and 315 million years ago,[1] and were the most common and largest land vertebrates of the
Permian period.[2] But in the Triassic period a previously obscure group of sauropsids, the archosaurs,
became the dominant vertebrates and one archosaur group, the dinosaurs, dominated the rest of the
Mesozoic era. These changes forced the Mesozoic mammaliforms ("nearly mammals") into nocturnal niches,
and may have contributed greatly to the development of mammalian traits such as endothermy, hair and a
large brain. Later in the Mesozoic mammals spread into other ecological niches, for example aquatic, gliding
and even preying on dinosaurs.
Most of the evidence consists of fossils. For many years fossils of Mesozoic mammals and their
immediate ancestors were very rare and fragmentary, but since the mid 1990s there have been many
important new finds, especially in China. The relatively new techniques of molecular phylogenetics have also
shed light on some aspects of mammalian evolution by estimating the timing of important divergence points
for modern species. When used carefully, these techniques often, but not always, agree with the fossil
record.
Although mammary glands are the signature feature of modern mammals, little is known about the
evolution of lactation, and virtually nothing is known about the evolution of another distinctive feature, the
neocortex region of the brain. Most study of the evolution of mammals centers around the development of the
middle ear bones from components of the ancestral amniote jaw joint. Other much-studied aspects include
the evolution of erect limb posture, a bony secondary palate, fur and hair, and warm-bloodedness.
Contents
[hide]
• 1 Definition of "mammal"
• 2 The ancestry of mammals
• 2.1 Amniotes
• 2.2 Synapsids
• 3 Therapsids
• 3.1 Therapsid family tree
• 3.2 Biarmosuchia
• 3.3 Dinocephalians
• 3.4 Anomodonts
• 3.5 Theriodonts
• 3.6 Cynodonts
• 4 Triassic takeover
• 5 From cynodonts to true mammals
• 5.1 Many uncertainties
• 5.2 Mammals or mammaliformes?
• 5.3 Family tree — cynodonts to mammals
• 5.4 Multituberculates
• 5.5 Morganucodontidae
• 5.6 Docodonts
[edit] Definition of "mammal"
 Mammalian and non-mammalian jaws. In the mammal configuration, the quadrate and articular
bones are much smaller and form part of the middle ear. Note that in mammals the lower jaw consists of only
the dentary bone.
Living mammal species can be identified by the presence of milk producing mammary glands in
females. Other features are required when classifying fossils, since mammary glands and other soft-tissue
features are not visible in fossils.
Paleontologists therefore use a distinguishing feature that is shared by all living mammals (including
monotremes) but is not present in any of the early Triassic therapsids ("mammal-like reptiles"): Mammals use
two bones for hearing that all other amniotes use for eating. The earliest amniotes had a jaw joint composed
of the articular (a small bone at the back of the lower jaw) and the quadrate (a small bone at the back of the
upper jaw). All non-mammalian amniotes use this system including lizards, crocodilians, dinosaurs (and their
descendants the birds), and therapsids. But mammals have a different jaw joint, composed only of the
dentary (the lower jaw bone, which carries the teeth) and the squamosal (another small skull bone). In
mammals, the quadrate and articular bones have become the incus and malleus bones in the middle ear.[3]
[4]
Mammals also have a double occipital condyle; they have two knobs at the base of the skull that fit
into the topmost neck vertebra, and other vertebrates have a single occipital condyle.[3] But paleontologists
use only the jaw joint and middle ear as criteria for identifying fossil mammals, as it would be confusing if
they found a fossil that had one feature but not the other (e.g. a mammalian jaw and ear but a non-
mammalian single occipital condyle).
Due to the incremental changes in transitional fossils, it has been said
 We may again ask the question, What is a mammal? Where we draw the line between reptile
and mammal has no biological significance. It is purely a matter of convenience. There are two
obvious choices, both immediately following a period of rapid evolution that make as definite a
break as we can hope to find.[5]

[edit] The ancestry of mammals

Tetrapods

Amphibians

Amniotes
Sauropsids (including
dinosaurs)

Pelycosaurs

Therapsids

Mammals
[edit] Amniotes
The first fully terrestrial vertebrates were amniotes — their eggs had internal membranes that allowed
the developing embryo to breathe but kept water in. This allowed amniotes to lay eggs on dry land, while
amphibians generally need to lay their eggs in water (a few amphibians, such as the Surinam toad, have
evolved other ways of getting round this limitation). The first amniotes apparently arose in the late
Carboniferous from the ancestral reptiliomorphs.
Within a few million years two important amniote lineages became distinct: mammals' synapsid
ancestors and the sauropsids, from which lizards, snakes, crocodilians, dinosaurs and birds are descended.
[1] The earliest known fossils of synapsids and sauropsids (such as Archaeothyris and Hylonomus resp.)
date from about 320 to 315 million years ago. Unfortunately it is difficult to be sure about when each of them
evolved, since vertebrate fossils from the late Carboniferous are very rare, and therefore the actual first
occurrences of each of these types of animal might have been considerably earlier.[6]

[edit] Synapsids

The original synapsid skull structure has one hole behind each eye, in a fairly low position on the
skull (lower right in this image).
Synapsid skulls are identified by the distinctive pattern of the holes behind each eye, which served
the following purposes:
• made the skull lighter without sacrificing strength.
• saved energy by using less bone.
• probably provided attachment points for jaw muscles. Having attachment points further away
from the jaw made it possible for the muscles to be longer and therefore to exert a strong pull over a
wide range of jaw movement without being stretched or contracted beyond their optimum range.
Early Permian terrestrial fossils indicate that one synapsid group, the pelycosaurs, were the most
common land vertebrates of their time and included the largest land animals of the time.[2]
[edit] Therapsids
Therapsids descended from pelycosaurs in the middle Permian and took over their position as the
dominant land vertebrates. They differ from pelycosaurs in several features of the skull and jaws, including
larger temporal fenestrae and incisors that are equal in size.[7]
The therapsids went through a series of stages, beginning with animals that were very like their
pelycosaur ancestors and ending with some that could easily be mistaken for mammals:[8]
• gradual development of a bony secondary palate. Most books and articles interpret this as a
prequisite for the evolution of mammals' high metabolic rate, because it enabled these animals to eat
and breathe at the same time. But some scientists point out that some modern ectotherms use a
fleshy secondary palate to separate the mouth from the airway, and that a bony palate provides a
surface on which the tongue can manipulate food, facilitating chewing rather than breathing.[9] The
interpretation of the bony secondary palate as an aid to chewing also suggests the development of a
faster metabolism, since chewing makes it possible to digest food more quickly. In mammals the
palate is formed by two specific bones, but various Permian therapsids had other combinations of
bones in the right places to function as a palate.
• the dentary gradually becomes the main bone of the lower jaw.
• progress towards an erect limb posture, which would increase the animals' stamina by
avoiding Carrier's constraint. But this process was erratic and very slow — for example: all
herbivorous therapsids retained sprawling limbs (some late forms may have had semi-erect hind
limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones also
had semi-sprawling hindlimbs. In fact, modern monotremes still have semi-sprawling limbs.
• in the Triassic, progress towards the mammalian jaw and middle ear.
 • there is plausible evidence of hair in Triassic therapsids, but none for Permian therapsids
(see below).
• some scientists have argued that some Triassic therapsids show signs of lactation (see
below).

[edit] Therapsid family tree

(simplified from [7]; only those that are most relevant to the evolution of mammals are described
below)
Therapsids

Eutherapsida

Dinocephalia

Neotherapsida
Anomodonts

Dicynodonts
 Theriodontia

Gorgonopsia

Eutheriodontia

Therocephalia

Cynodontia
(Mammals,
eventually)
 Only the dicynodonts, therocephalians and cynodonts survived into the Triassic.

[edit] Biarmosuchia
The Biarmosuchia were the most primitive and pelycosaur-like of the therapsids.

[edit] Dinocephalians
Dinocephalians ("terrible heads") were large, some as large as a rhinoceros, and included both
carnivores and herbivores. Some of the carnivores had semi-erect hindlimbs, but all dinocephalians had
sprawling forelimbs. In many ways they were very primitive therapsids, for example they had no secondary
palate and their jaws were rather "reptilian".[10]

[edit] Anomodonts

Lystrosaurus, one of the few species of dicynodonts that survived the Permian-Triassic extinction
event
The anomodonts ("anomalous teeth") were the most successful of the herbivorous therapsids — one
sub-group, the dicynodonts, survived almost to the end of the Triassic. But anomodonts were very different
from modern herbivorous mammals, as their only teeth were a pair of fangs in the upper jaw and it is
generally agreed that they had beaks like those of birds or ceratopsians.[11]
[edit] Theriodonts
The theriodonts ("beast teeth") and their descendants had jaw joints in which the lower jaw's articular
bone tightly gripped the skull's very small quadrate bone. This allowed a much wider gape, and one group,
the carnivorous gorgonopsians ("gorgon faces"), took advantage of this to develop "sabre teeth". But the
theriodont's jaw hinge had a longer term significance — the much reduced size of the quadrate bone was an
important step in the development of the mammalian jaw joint and middle ear.
The gorgonopsians still had some primitive features: no bony secondary palate (but other bones in
the right places to perform the same functions); sprawling forelimbs; hindlimbs that could operate in both
sprawling and erect postures. But the therocephalians ("beast heads"), which appear to have arisen at about
the same time as the gorgonopsians, had additional mammal-like features, e.g. their finger and toe bones
had the same number of phalanges (segments) as in early mammals (and the same number that primates
have, including humans).[12]

[edit] Cynodonts
 Life restoraton of the cynodont Trirachodon within a burrow
The cynodonts, a theriodont group that also arose in the late Permian, include the ancestors of all
mammals — one sub-group, the trithelodonts, is widely regarded as the most likely to contain mammals'
ancestor. Cynodonts' mammal-like features include further reduction in the number of bones in the lower jaw;
a secondary bony palate; cheek teeth with a complex pattern in the crowns; the brain filled the endocranial
cavity.[13]
Multi-chambered burrows have been found, containing as many as 20 skeletons of the Early Triassic
cynodont Trirachodon; the animals are thought to have been drowned by a flash flood. The extensive shared
burrows indicate that these animals were capable of complex social behaviors.[14]

[edit] Triassic takeover

The catastrophic Permian-Triassic mass extinction killed off about 70 percent of terrestrial vertebrate
species, and the majority of land plants. As a result[15]
• Ecosystems and food chains collapsed, and the recovery took about 6 million years.
• The survivors had to re-start the struggle for dominance of their former ecological niches —
even the cynodonts, which had seemed on the way to dominance at the end of the Permian.
But the cynodonts lost out to a previously obscure group of sauropsids, the archosaurs (which
include the ancestors of crocodilians, dinosaurs and birds). This reversal of fortunes is often called the
"Triassic takeover". Several explanations have been offered for it, but the most likely is that the early Triassic
was predominantly arid and therefore archosaurs' superior water conservation gave them a decisive
advantage (all known sauropsids have glandless skins and excrete uric acid, which requires less water to
keep it sufficiently liquid than urea, which mammals excrete and presumably therapsids excreted).[8][16] The
Triassic takeover was gradual — in the earliest part of the Triassic cynodonts were the main predators and
lystrosaurs were the main herbivores, but by the mid-Triassic archosaurs dominated all the large carnivore
and herbivore niches.
But the Triassic takeover may have been a vital factor in the evolution of cynodonts into mammals.
The cynodonts' descendants were only able to survive as small, mainly nocturnal insectivores.[13] As a
result:
• The therapsid trend towards differentiated teeth with precise occlusion accelerated, because
of the need to hold captured arthropods and crush their exoskeletons.
• Nocturnal life required advances in thermal insulation and temperature regulation to enable
the ancestors of mammals to be active in the cool of the night.[17]
• Acute senses of hearing and smell became vital.
• This accelerated the development of the mammalian middle ear, and therefore of the
mammalian jaw since bones that had been part of the jaw joint became part of the middle
ear.
• The increase in the size of the olfactory and auditory lobes of the brain increased
brain weight as a total percentage of body weight. Brain tissue requires a disproportionate
amount of energy.[18][19] The need for more food to support the enlarged brains increased
the pressures for improvements in insulation, temperature regulation and feeding.
• As a side-effect of the nocturnal life, discerning colors became less important (they lost two
out of four opsins), and this is reflected in the fact that most mammals have poor color vision,
including the "lower primates" such as lemurs.[20]
 [edit] From cynodonts to true mammals
[edit] Many uncertainties
While the Triassic takeover probably accelerated the evolution of mammals, it made life more difficult
for paleontologists because good fossils of the nearly-mammals are extremely rare, mainly because they
were mostly smaller than rats:
• They were largely restricted to environments that are less likely to provide good fossils. The
best terrestrial environments for fossilization are floodplains, where seasonal floods quickly cover
dead animals in a protective layer of silt that is later compressed into sedimentary rock. But
floodplains are dominated by medium to large animals, and the Triassic therapsids and near-
mammals could not compete with archosaurs in the medium to large size range.
• Their delicate bones were vulnerable to being destroyed before they could be fossilized — by
scavengers (including fungi and bacteria) and by being trodden on.
• Small fossils are harder to spot and more vulnerable to being destroyed by weathering and
other natural stresses before they are discovered.
In fact it was said as recently as the 1980s that all the Mesozoic fossils of mammals and near-
mammals could be contained in a few shoeboxes — and they were mostly teeth, which are the most durable
of all tissues.[21] Since then, the number of Mesozoic fossil mammals has increased, from 116 genera
known in 1979 to about 310 in 2007, with an increase in quality such that "at least 18 Mesozoic mammals are
represented by nearly complete skeletons".[22]
As a result:
 • In many cases it is difficult to assign a Mesozoic mammal or near-mammal fossil to a genus.
• All the available fossils of a genus seldom add up to a complete skeleton, and hence it is
difficult to decide which genera are most like each other and therefore most likely to be closely-
related. In other words, it becomes very difficult to classify them by means of cladistics, which is the
most reliable and least subjective method currently available.
So the evolution of mammals in the Mesozoic is full of uncertainties, although there is no room for
doubt that true mammals did first appear in the Mesozoic.

[edit] Mammals or mammaliformes?

One result of these uncertainties has been a change in the paleontologists' definition of "mammal".
For a long time a fossil was considered a mammal if it met the jaw-ear criterion (the jaw joint consists only of
the squamosal and dentary; and the articular and the quadrate bones have become the middle ear's malleus
and incus). But more recently paleontologists have usually defined "mammal" as the last common ancestor
of monotremes, marsupials and placentals and all of its descendants. So they had to define another clade
mammaliformes to accommodate all the animals that are more mammal-like than cynodonts but less closely
related to monotremes, marsupials and placentals.[23] Although this now appears to be the majority
approach, some paleontologists have resisted it because it simply moves most of the problems into the new
clade without solving them; the clade mammaliformes includes some animals with "mammalian" jaw joints
and some with "reptilian" (articular-to-quadrate) jaw joints; and the newer definition of "mammal" and
"mammaliformes" depend on last common ancestors of both groups, which have not yet been found.[21]
Despite these objections, this article follows the majority approach and treats most of the cynodonts'
Mesozoic descendants as mammaliformes.
[edit] Family tree — cynodonts to mammals
(based on Mammaliformes - Palaeos)
Cynodontia

Dvinia

Procynosuchidae

Epicynodontia

Eucynodontia

Probainognathia
Trithelodontidae
Chiniquodontidae

Mammaliaformes
Allotheria
Multituberc

Morganucodontidae

Docodonta

Hadrocodium
 Symmetrodonta
Kuehneotheriidae

crown group
Mammals (all
descendants of the last
common ancestor of all
living mammals)
[edit] Multituberculates

Skull of the multituberculate Ptilodus

Multituberculates (named for the multiple tubercles on their "molars") are often called the "rodents of
the Mesozoic" but this is an example of convergent evolution rather than meaning that they are closely
related to the Rodentia. At first sight they look like mammals: their jaw joints consists of only the dentary and
squamosal bones, and the quadrate and articular bones are part of the middle ear; their teeth are
differentiated, occlude and have mammal-like cusps; they have a zygomatic arch; the structure of the pelvis
suggests that they gave birth to tiny helpless young, like modern marsupials. And they lived for over 120
million years (from mid Jurassic, about 160M years ago, to early Oligocene, about 35M years ago), which in
terms of clade longevity would make them the most successful mammaliformes ever. But a closer look
shows that they are very different from modern mammals:[23]
 • Their "molars" have two parallel rows of tubercles, unlike the tribosphenic (three-peaked)
molars of early mammals.
• The chewing action is completely different. Mammals chew with a side-to-side grinding
action, which means that usually the molars occlude on only one side at a time. Multituberculates'
jaws were incapable of side-to-side movement and chewed by dragging the lower teeth backwards
against the upper ones as the jaw closed.
• The anterior (forward) part of the zygomatic arch mostly consists of the maxilla (upper
jawbone) rather than the jugal, and the jugal is a small bone in a little slot in the maxillary process
(extension).
• The squamosal does not form part of the braincase.
• The rostrum (snout) is unlike that of mammals, in fact it looks more like that of a pelycosaur
such as Dimetrodon. The multituberculate rostrum is box-like, with the large flat maxillae forming the
sides, the nasal the top, and the tall premaxilla at the front.
 Morganucodontidae and other transitional forms had both types of jaw joint: dentary-squamosal
(front) and articular-quadrate (rear).

[edit] Morganucodontidae
The Morganucodontidae first appeared in the late Triassic, about 205M years ago. They are an
excellent example of transitional fossils, since they have both the dentary-squamosal and articular-quadrate
jaw joints.[24] They were also one of the first discovered and most thoroughly studied of the mammaliformes,
since an unusually large number of morganucodont fossils have been found.

[edit] Docodonts

Reconstruction of Castorocauda. Note the fur and the adaptations for swimming (broad, flat tail;
webbed feet) and for digging (robust limbs and claws).
The most notable member of the docodonts is Castorocauda ("beaver tail"), which lived in the mid
Jurassic about 164M years ago and was first discovered in 2004 and described in 2006. Castorocauda was
not a typical docodont (most were omnivores) and not a true mammal, but it is extremely important in the
study of the evolution of mammals because the first find was an almost complete skeleton (a real luxury in
paleontology) and it breaks the "small nocturnal insectivore" stereotype:[25]
• It was noticeably larger than most Mesozoic mammal-like fossils — about 17 in (43 cm) from
its nose to the tip of its 5-inch (130 mm) tail, and may have weighed 500–800 g (18–28 oz).
• It provides the earliest absolutely certain evidence of hair and fur. Previously the earliest was
Eomaia, a true mammal from about 125M years ago.
• It had aquatic adaptations including flattened tail bones and remnants of soft tissue between
the toes of the back feet, suggesting that they were webbed. Previously the earliest known semi-
aquatic mammal-like animals were from the Eocene, about 110M years later.
• Castorocauda's powerful forelimbs look adapted for digging. This feature and the spurs on its
ankles make it resemble the platypus, which also swims and digs.
• Its teeth look adapted for eating fish: the first two molars had cusps in a straight row, which
made them more suitable for gripping and slicing than for grinding; and these molars are curved
backwards, to help in grasping slippery prey.

[edit] Hadrocodium
 Hadrocodium skull. The jaw joint is fully mammalian (squamosal-dentary only) and farther forward
than in earlier transitional forms.
The consensus family tree above shows Hadrocodium as an "aunt" of true mammals, while
symmetrodonts and kuehneotheriids are more closely related to true mammals. But fossils of symmetrodonts
and kuehneotheriids are so few and fragmentary that they are poorly understood and may be paraphyletic.
[26] On the other hand there are good fossils of Hadrocodium (about 195M years ago in the very early
Jurassic) and they have some important features: [27]
• The jaw joint consists only of the squamosal and dentary bones, and the jaw contains no
smaller bones to the rear of the dentary, unlike the therapsid design.
• In therapsids and most mammaliformes the eardrum stretched over a trough at the rear of
the lower jaw. But Hadrocodium had no such trough, which suggests its ear was part of the cranium,
as it is in mammals — and hence that the former articular and quadrate had migrated to the middle
ear and become the malleus and incus. On the other hand the dentary has a "bay" at the rear that
mammals lack. This suggests that Hadrocodium's dentary bone retained the same shape that it
would have had if the articular and quadrate had remained part of the jaw joint, and therefore that
Hadroconium or a very close ancestor may have been the first to have a fully mammalian middle ear.
• Therapsids and earlier mammaliforms had their jaw joints very far back in the skull, partly
because the ear was at the rear end of the jaw but also had to be close to the brain. This
arrangement limited the size of the braincase, because it forced the jaw muscles to run round and
over it. Hadrocodium's braincase and jaws were no longer bound to each other by the need to
support the ear, and its jaw joint was further forward. In its descendants or those of animals with a
similar arrangement, the brain case was free to expand without being constrained by the jaw and the
jaw was free to change without being constrained by the need to keep the ear near the brain — in
 other words it now became possible for mammal-like animals both to develop large brains and to
adapt their jaws and teeth in ways that were purely specialized for eating.

[edit] The earliest true mammals

This part of the story introduces new complications, since true mammals are the only group that still
has living members:
• One has to distinguish between extinct groups and those that have living representatives.
• One often feels compelled to try to explain the evolution of features that do not appear in
fossils. This endeavor often involves Molecular phylogenetics, a technique that has become popular
since the mid-1980s but is still often controversial because of its assumptions, especially about the
reliability of the molecular clock.

[edit] Family tree of early true mammals

(based on Mammalia: Overview - Palaeos; X marks extinct groups)
Mammals
 Australosphenida

Ausktribosphenidae X

Monotremes

Triconodonta
X

Spalacotheroidea
X

Cladotheria
Dryolestoidea
X

Theria

Metatheria
 Eutheria

[edit] Australosphenida and Ausktribosphenidae

Australosphenida and Ausktribosphenidae
Ausktribosphenidae is a group name that has been given to some rather puzzling finds that:[28]
• appear to have tribosphenic molars, a type of tooth that is otherwise known only in placentals
and marsupials.[29]
• come from mid Cretaceous deposits in Australia — but Australia was connected only to
Antarctica, and placentals originated in the northern hemisphere and were confined to it until
continental drift formed land connections from North America to South America, from Asia to Africa
and from Asia to India (the late Cretaceous map at [1] shows how the southern continents are
separated).
• are represented only by skull and jaw fragments, which is not very helpful.
Australosphenida is a group that has been defined in order to include the Ausktribosphenidae and
monotremes. Asfaltomylos (mid- to late Jurassic, from Patagonia) has been interpreted as a basal
australosphenid (animal that has features shared with both Ausktribosphenidae and monotremes; lacks
features that are peculiar to Ausktribosphenidae or monotremes; also lacks features that are absent in
Ausktribosphenidae and monotremes) and as showing that australosphenids were widespread throughout
Gondwanaland (the old Southern hemisphere super-continent).[30]
 But recent analysis of Teinolophos suggests Teinolophos (about 115M years ago) was a "crown
group" (advanced and relatively specialised) monotreme, so the basal (most primitive) monotremes must
have appeared considerably earlier; that some alleged Australosphenids were also "crown group"
monotremes (e.g. Steropodon); and that other alleged Australosphenids (e.g. Ausktribosphenos, Bishops,
Ambondro, Asfaltomylos) are therefore more closely related to and possibly members of the Therian
mammals (group that includes marsupials and placentals, see below).[31]

[edit] Monotremes
The earliest known monotreme is Teinolophos, which lived about 123M years ago in Australia.
Recent analysis suggest that it was not a basal (primitive, ancestral) monotreme but a full-fledged platypus,
and therefore that the platypus and echidna lineages diverged considerably earlier and that basal
monotremes were even earlier.[31]
A more recent study, however, has suggested that the Teinolophis was really a basal monotreme
and that the radiation of modern monotremes occurred more recently. Platypus fossils are known to date
back to the Paleocene; their semi-aquatic lifestyle prevented this group from being outcompeted by the
marsupials that invaded Australia millions of years ago since the reproductive mode of marsupials prevents
them from taking to the water. Echidnas were found to have diverged from the platypus lineage as recently
as 19-48M when they made their transition from semi-aquatic to terrestrial lifestyle.[32]
Monotremes have some features that may be inherited from the original amniotes:
• they use the same orifice to urinate, defecate and reproduce ("monotreme" means "one
hole") — as lizards and birds also do.
• they lay eggs that are leathery and uncalcified, like those of lizards, turtles and crocodilians.
 Unlike in other mammals, female monotremes do not have nipples and feed their young by
"sweating" milk from patches on their bellies.
Of course these features are not visible in fossils, and the main characteristics from paleontologists'
point of view are:[28]
• a slender dentary bone in which the coronoid process is small or non-existent.
• the external opening of the ear lies at the posterior base of the jaw.
• the jugal bone is small or non-existent.
• a primitive pectoral girdle with strong ventral elements: coracoids, clavicles and interclavicle.
Note: therian mammals have no interclavicle.[33]
• sprawling or semi-sprawling forelimbs.

[edit] Theria
Theria ("beasts") is a name applied to the hypothetical group from which both metatheria (which
include marsupials) and eutheria (which include placentals) descended. Although no convincing fossils of
basal therians have been found (just a few teeth and jaw fragments), metatheria and eutheria share some
features that one would expect to have been inherited from a common ancestral group:[34]
Therian form of crurotarsal ankle. Adapted with permission from Palaeos
• no interclavicle.[33]
• coracoid bones non-existent or fused with the shoulder blades to form coracoid processes.
• a type of crurotarsal ankle joint in which: the main joint is between the tibia and astragalus;
the calcaneum has no contact with the tibia but forms a heel to which muscles can attach. (The other
 well-known type of crurotarsal ankle is seen in crocodilians and works differently — most of the
bending at the ankle is between the calcaneum and astragalus).
• tribosphenic molars.[29]
Tribosphenic molars have been found in fossils from Madagascar, which indicates that therian
mammals are at least 167 million years old.[35]

[edit] Metatheria
The living Metatheria are all marsupials ("animals with pouches"). A few fossil genera such as the
Mongolian late Cretaceous Asiatherium may be marsupials or members of some other metatherian group(s).
[36][37]
The oldest known marsupial is Sinodelphys, found in 125M-year old early Cretaceous shale in
China's northeastern Liaoning Province. The fossil is nearly complete and includes tufts of fur and imprints of
soft tissues.[38]
Didelphimorphia (common opossums of the Western Hemisphere) first appeared in the late
Cretaceous and still have living representatives, probably because they are mostly semi-arboreal
unspecialized omnivores.[39]
The best-known feature of marsupials is their method of reproduction:
• The mother develops a kind of yolk sack in her womb that delivers nutrients to the embryo.
Embryos of bandicoots, koalas and wombats additionally form placenta-like organs that connect
them to the uterine wall, although the placenta-like organs are smaller than in placental mammals
and it is not certain that they transfer nutrients from the mother to the embryo.[40]
 • Pregnancy is very short, typically 4 to 5 weeks. The embryo is born at a very young age of
development, and is usually less than 2 in (5.1 cm) long at birth. It has been suggested that the short
pregnancy is necessary to reduce the risk that the mother's immune system will attack the embryo.
• The newborn marsupial uses its forelimbs (with relatively strong hands) to climb to a nipple,
which is usually in a pouch on the mother's belly. The mother feeds the baby by contracting muscles
over her mammary glands, as the baby is too weak to suck. The newborn marsupial's need to use its
forelimbs in climbing to the nipple has prevented the forelimbs from evolving into paddles or wings
and has therefore prevented the appearance of aquatic or truly flying marsupials (although there are
several marsupial gliders).

Palate of thylacine, showing one of the paired palatal fenestrae (top left), which are a signature
feature of marsupials.
Although some marsupials look very like some placentals (the thylacine or "marsupial wolf" is a good
example), marsupial skeletons have some features that distinguish them from placentals:[41]
• Some, including the thylacine, have 4 molars. No placentals have more than 3.
• All have a pair of palatal fenestrae, window-like openings on the bottom of the skull (in
addition to the smaller nostril openings).
 Marsupials also have a pair of marsupial bones (sometimes called "epipubic bones"), which support
the pouch in females. But these are not unique to marsupials, since they have been found in fossils of
multituberculates, monotremes, and even eutherians — so they are probably a common ancestral feature that
disappeared at some point after the ancestry of living placental mammals diverged from that of marsupials.
[42][43] Some researchers think the epipubic bones' original function was to assist locomotion by supporting
some of the muscles that pull the thigh forwards.[44]

[edit] Eutheria
Main article: Eutheria
The living Eutheria ("true beasts") are all placentals. But the earliest known eutherian, Eomaia, found
in China and dated to 125M years ago, has some features that are more like those of marsupials (the
surviving metatherians):[45]
 Fossil of Eomaia in the Hong Kong Science Museum.
• Epipubic bones extending forwards from the pelvis, which are not found in any modern
placental, but are found in all other mammals — non-placental eutherians, marsupials, monotremes
and mammaliformes — and even in the cynodont therapsids that are closest to mammals. Their
function is to stiffen the body during locomotion.[46] This stiffening would be harmful in pregnant
placentals, whose abdomens need to expand.[47]
• A narrow pelvic outlet, which indicates that the young were very small at birth and therefore
pregnancy was short, as in modern marsupials. This suggests that the placenta was a later
development.
• 5 incisors in each side of the upper jaw. This number is typical of metatherians, and the
maximum number in modern placentals is 3, except for homodonts such as the armadillo. But
Eomaia's molar to premolar ratio (it has more pre-molars than molars) is typical of eutherians,
including placentals, and not normal in marsupials.
Eomaia also has a Meckelian groove, a primitive feature of the lower jaw that is not found in modern
placental mammals.
These intermediate features are consistent with molecular phylogenetics estimates that the
placentals diversified about 110M years ago, 15M years after the date of the Eomaia fossil.
Eomaia also has many features that strongly suggest it was a climber, including several features of
the feet and toes; well-developed attachment points for muscles that are used a lot in climbing; and a tail that
is twice as long as the rest of the spine.
 Placentals' best-known feature is their method of reproduction:
• The embryo attaches itself to the uterus via a large placenta via which the mother supplies
food and oxygen and removes waste products.
• Pregnancy is relatively long and the young are fairly well-developed at birth. In some species
(especially herbivores living on plains) the young can walk and even run within an hour of birth.
It has been suggested that the evolution of placental reproduction was made possible by retroviruses
that:[48]
• make the interface between the placenta and uterus into a syncytium, i.e. a thin layer of cells
with a shared external membrane. This allows the passage of oxygen, nutrients and waste products
but prevents the passage of blood and other cells, which would cause the mother's immune system
to attack the fetus.
• reduce the aggressiveness of the mother's immune system (which is good for the foetus but
makes the mother more vulnerable to infections).
From a paleontologist's point of view, eutherians are mainly distinguished by various features of their
teeth,[49] ankles and feet.[50]

[edit] Expansion of ecological niches in the Mesozoic

There is still some truth in the "small, nocturnal insectivores" stereotype but recent finds, mainly in
China, show that some mammaliforms and true mammals were larger and had a variety of lifestyles. For
example:
 • Castorocauda, a member of Docodonta which lived in the middle Jurassic about 164 million
years, was about 42.5 cm (16.7 in) long, weighed 500–800 g (18–28 oz), had limbs that were adapted
for swimming and digging and teeth adapted for eating fish.[25]
• Multituberculates, are allotherians that survived for over 125 million years (from mid Jurassic,
about 160M years ago, to early Oligocene, about 35M years ago) are often called the "rodents of the
Mesozoic", because they had continuously-growing incisors like those of modern rodents.[23]

Repenomamus sometimes preyed on young dinosaurs

• Fruitafossor, from the late Jurassic period about 150 million years ago, was about the size of
a chipmunk and its teeth, forelimbs and back suggest that it broke open the nest of social insects to
prey on them (probably termites, as ants had not yet appeared).[51]
 • Volaticotherium, allotherians from the boundary the early Cretaceous about 125M years ago,
is the earliest-known gliding mammal and had a gliding membrane that stretched out between its
limbs, rather like that of a modern flying squirrel. This also suggests it was active mainly during the
day.[52]
• Repenomamus, tricodonts from the early Cretaceous 130 million years ago, was a stocky,
badger-like predator that sometimes preyed on young dinosaurs. Two species have been
recognized, one more than 1 m (39 in) long and weighing about 12–14 kg (26–31 lb), the other less
than 0.5 m (20 in) long and weighing 4–6 kg (8.8–13 lb).[53][54]

[edit] Evolution of major groups of living mammals

There are currently vigorous debates between traditional paleontologists ("fossil-hunters") and
molecular phylogeneticists about how and when the true mammals diversified, especially the placentals.
Generally the traditional paelontologists date the appearance of a particular group by the earliest known
fossil whose features make it likely to be a member of that group, while the molecular phylogeneticists
suggest that each lineage diverged earlier (usually in the Cretaceous) and that the earliest members of each
group were anatomically very similar to early members of other groups and differed only in their genes.
These debates extend to the definition of and relationships between the major groups of placentals — the
controversy about Afrotheria is a good example.
[edit] Fossil-based family tree of placental mammals
Here is a very simplified version of a typical family tree based on fossils, based on Cladogram of
Mammalia - Palaeos. It tries to show the nearest thing there is at present to a consensus view, but some
paleontologists have very different views, for example:[55]
• The most common view is that placentals originated in the southern hemisphere, but some
paleontologists argue that they first appeared in Laurasia (old supercontinent containing modern
Asia, N. America and Europe).
• Paleontologists differ about when the first placentals appeared, with estimates ranging from
20M years before the end of the Cretaceous to just after the end of the Cretaceous. And molecular
biologists argue for a much earlier origin.
• Most paleontologists suggest that placentals should be divided into Xenarthra and the rest,
but a few think these animals diverged later.
For the sake of brevity and simplicity the diagram omits some extinct groups in order to focus on the
ancestry of well-known modern groups of placentals — X marks extinct groups. The diagram also shows the
following:
• the age of the oldest known fossils in many groups, since one of the major debates between
traditional paleontologists and molecular phylogeneticists is about when various groups first became
distinct.
• well-known modern members of most groups.
 Eutheria

Epitheria (latest Cretaceous)

Anagalida
 Zalambdalestidae X
(late Cretaceous)

Macroscelidea
(late Eocene)
(elephant shrews)
Anagaloidea
X

Glires (early Paleocene)

Lagomorpha
(Eocene)
(rabbits, hares, pikas)
Rodentia (late
Paleocene)
(mice & rats, squirrels,
porcupines)
 Archonta
Scandentia (mid
Eocene)
(tree shrews)
Primatomorpha

Plesiadapiformes X

Primates (early Paleocene)

(tarsiers, lemurs, monkeys, apes including humans)

Dermoptera (late Eocene)

(colugos)

Chiroptera (late Paleocene)

(bats)

Carnivora
(early Paleocene)
(cats, dogs, bears,
seals)
Ungulatomorpha (late Cretaceous)
Eparctocyona (late Cretaceous)
(some
extinct groups)
X
 Arctostylopida
X (late Paleocene)

Mesonychia
X (mid Paleocene)
(predators /
scavengers, but not
closely related to
modern carnivores)
Cetartiodactyla
Cetacea
(early Eocene)
(whales, dolphins,
porpoises)
Artiodactyla
(early Eocene)
(even-toed
ungulates: pigs,
hippos, camels,
giraffes, cattle,
deer)
 Altungulata
Hilalia
X

Perissodactyla (late Paleocene)

(odd-toed ungulates: horses, rhinos, tapirs)

Tubulidentata (early Miocene)

(aardvarks)

Paenungulata ("not quite ungulates")

Hyracoidea (early
Eocene)
(hyraxes)
Sirenia (early
Eocene)
(manatees, dugongs)

Proboscidea (early
Eocene)
(elephants)
This family tree contains some surprises and puzzles. For example:
• The closest living relatives of cetaceans (whales, dolphins, porpoises) are artiodactyls,
hoofed animals, which are almost all pure vegetarians.
• Bats are fairly close relatives of primates.
• The closest living relatives of elephants are the aquatic sirenians, while their next relatives
are hyraxes, which look more like well-fed guinea pigs.
• There is little correspondence between the structure of the family (what was descended from
what) and the dates of the earliest fossils of each group. For example the earliest fossils of
perissodactyls (the living members of which are horses, rhinos and tapirs) date from the late
Paleocene but the earliest fossils of their "sister group" the Tubulidentata date from the early
Miocene, nearly 50M years later. Paleontologists are fairly confident about the family relationships,
which are based on cladistic analyses, and believe that fossils of the ancestors of modern aardvarks
have simply not been found yet.
 [edit] Family tree of placental mammals according to molecular
phylogenetics
Molecular phylogenetics uses features of organisms' genes to work out family trees in much the
same way as paleontologists do with features of fossils — if two organisms' genes are more similar to each
other than to those of a third organism, the two organisms are more closely related to each other than to the
third.
Molecular phylogeneticists have proposed a family tree that is very different from the one with which
paleontologists are familiar. Like paleontologists, molecular phylogeneticists have different ideas about
various details, but here is a typical family tree according to molecular phylogenetics:[56][57] Note that the
diagram shown here omits extinct groups, as one cannot extract DNA from fossils.
Eutheria
Atlantogenata "born round the Atlantic ocean")
Xenarthra
(armadillos, anteaters,
sloths)
Afrotheria
Afroinsectiphilia (golden moles, tenrecs, otter
shrews)

unnamed
Macroscelidea (elephant
shrews)

Tubulidentata (aardvarks)

Paenungulata ("not quite ungulates")

Hyracoidea (hyraxes)

Proboscidea (elephants)

Sirenia (manatees, dugongs)

 Boreoeutheria ("northern true / placental mammals")
Laurasiatheria

Erinaceomorpha (hedgehogs, gymnures)

Soricomorpha (moles, shrews,

solenodons)

Cetartiodactyla

Pegasoferae

Pholidota (pangolins)

Chiroptera (bats)
 Carnivora (cats, dogs, bears, seals)

Perissodactyla (horses, rhinos,

tapirs).

Euarchontoglires
Glires

Lagomorpha(rabbits, hares, pikas)

Rodentia (late Paleocene)(mice & rats, squirrels,

porcupines)

Euarchonta
 Scandentia (tree shrews)

Dermoptera (colugos)

Primates (tarsiers, lemurs, monkeys, apes including

humans)

Here are the most significant of the many differences between this family tree and the one familiar to
paleontologists:
 • The top-level division is between Atlantogenata and Boreoeutheria, instead of between
Xenarthra and the rest. But some molecular phylogeneticists have proposed a 3-way top-level split
between Xenarthra, Afrotheria and Boreoeutheria.
• Afrotheria contains several groups that are only distantly related according to the
paleontologists' version: Afroinsectiphilia ("African insectivores"), Tubulidentata (aardvarks, which
paleontologists regard as much closer to odd-toed ungulates than to other members of Afrotheria),
Macroscelidea (elephant shrews, usually regarded as close to rabbits and rodents). The only
members of Afrotheria that paleontologists would regard as closely related are Hyracoidea (hyraxes),
Proboscidea (elephants) and Sirenia (manatees, dugongs).
• Insectivores are split into 3 groups: one is part of Afrotheria and the other two are distinct
sub-groups within Boreoeutheria.
• Bats are closer to Carnivora and odd-toed ungulates than to primates and Dermoptera
(colugos).
• Perissodactyla (odd-toed ungulates) are closer to Carnivora and bats than to Artiodactyla
(even-toed ungulates).
The grouping together of the Afrotheria has some geological justification. All surviving members of
the Afrotheria originate from South American or (mainly) African lineages — even the Indian elephant, which
diverged from an African lineage about 7.6 million years ago.[58] As Pangaea broke up Africa and South
America separated from the other continents less than 150M years ago, and from each other between 100M
and 80M years ago.[59][60] The earliest known eutherian mammal is Eomaia, from about 125M years ago.
So it would not be surprising if the earliest eutherian immigrants into Africa and South America were isolated
there and radiated into all the available ecological niches.
 Nevertheless these proposals have been controversial. Paleontologists naturally insist that fossil
evidence must take priority over deductions from samples of the DNA of modern animals. More surprisingly,
these new family trees have been criticised by other molecular phylogeneticists, sometimes quite harshly:[61]
• Mitochondrial DNA's mutation rate in mammals varies from region to region — some parts
hardly ever change and some change extremely quickly and even show large variations between
individuals within the same species.[62][63]
• Mammalian mitochondrial DNA mutates so fast that it causes a problem called "saturation",
where random noise drowns out any information that may be present. If a particular piece of
mitochondrial DNA mutates randomly every few million years, it will have changed several times in
the 60 to 75M years since the major groups of placental mammals diverged.[64]

[edit] Timing of placental evolution

Recent molecular phylogenetic studies suggest that most placental orders diverged about 100M to
85M years ago, but that modern families first appeared in the late Eocene and early Miocene.[65]
Some paleontologists object that no placental fossils have been found from before the end of the
Cretaceous — for example Maelestes gobiensis, from about 75M years ago, is a eutherian but not a true
placental.[66] Many Cretaceous fossil sites contain well-preserved lizards, salamanders, birds, and
mammals, but not the modern forms of mammals. It is likely that they simply did not exist, and that the
molecular clock runs fast during major evolutionary radiations.[67] On the other hand there is fossil evidence
from 85 million years ago of hoofed animals that may be ancestors of modern ungulates.[68]
Fossils of the earliest members of most modern groups date from the Paleocene, a few date from
later and very few from the Cretaceous, before the extinction of the dinosaurs. But some paleontologists,
influenced by molecular phylogenetic studies, have used statistical methods to extrapolate backwards from
fossils of members of modern groups and concluded that primates arose in the late Cretaceous.[69] However
statistical studies of the fossil record confirm that mammals were restricted in size and diversity right to the
end of the Cretaceous, and rapidly grew in size and diversity during the Early Paleocene.[70][71]

[edit] Evolution of mammalian features

[edit] Jaws and middle ears
See also Evolution of mammalian auditory ossicles
Hadrocodium, whose fossils date from the early Jurassic, provides the first clear evidence of fully
mammalian jaw joints and middle ears, in which the jaw joint is formed by the dentary and squamosal bones
while the articular and quadrate move to the middle ear, where they are known as the incus and malleus.
Curiously it is usually classified as a member of the mammaliformes rather than a as a true mammal.
One analysis of the monotreme Teinolophos suggested that this animal had a pre-mammalian jaw
joint formed by the angular and quadrate bones and that the typical mammalian middle ear evolved twice
independently, in monotremes and in therian mammals, but this idea has been disputed.[72] In fact 2 of the
suggestion's authors co-authored a later paper that reinterpreted the same features as evidence that
Teinolophos was a full-fledged platypus, which means it would have had a mammalian jaw joint and middle
ear.[31]
[edit] Milk production (lactation)
It has been suggested that lactation's original function was to keep eggs moist. Much of the argument
is based on monotremes (egg-laying mammals):[73][74][75]
• Monotremes do not have nipples but secrete milk from a hairy patch on their bellies.
• During incubation, monotremes' eggs are covered in a sticky substance whose origin is not
known. Before the eggs are laid, their shells have only three layers. Afterwards a fourth layer
appears, and its composition is different from that of the original three. The sticky substance and the
fourth layer may be produced by the mammary glands.
• If so, that may explain why the patches from which monotremes secrete milk are hairy — it is
easier to spread moisture and other substances over the egg from a broad, hairy area than from a
small, bare nipple.

[edit] Hair and fur

See also: Evolution of hair
The first clear evidence of hair or fur is in fossils of Castorocauda, from 164M years ago in the mid
Jurassic.
From 1955 onwards some scientists have interpreted the foramina (passages) in the maxillae (upper
jaws) and premaxillae (small bones in front of the maxillae) of cynodonts as channels that supplied blood
vessels and nerves to vibrissae (whiskers), and suggested that this was evidence of hair or fur.[76][77] But
foramina do not necessarily show that an animal had vibrissae — for example the modern lizard Tupinambis
has foramina that are almost identical to those found in the non-mammalian cynodont Thrinaxodon.[9][78]
[edit] Erect limbs
The evolution of erect limbs in mammals is incomplete — living and fossil monotremes have sprawling
limbs. In fact some scientists think that the parasagittal (non-sprawling) limb posture is a synapomorphy
(distinguishing characteristic) of the Boreosphenida, a group that contains the Theria and therefore includes
the last common ancestor of modern marsupials and placentals — and therefore that all earlier mammals had
sprawling limbs.[79]
Sinodelphys (the earliest known marsupial) and Eomaia (the earliest known eutherian) lived about
125M years ago, so erect limbs must have evolved before then.

[edit] Warm-bloodedness
"Warm-bloodedness" is a complex and rather ambiguous term, because it includes some or all of the
following:
• Endothermy, i.e. the ability to generate heat internally rather than via behaviors such as
basking or muscular activity.
• Homeothermy, i.e. maintaining a fairly constant body temperature.
• Tachymetabolism, i.e. maintaining a high metabolic rate, particularly when at rest. This
requires a fairly high and stable body temperature, since biochemical processes run about half as
fast if an animal's temperature drops by 10°C; most enzymes have an optimum operating
temperature and their efficiency drops rapidly outside the preferred range.
Since we can't know much about the internal mechanisms of extinct creatures, most discussion
focuses on homeothermy and tachymetabolism.
 Modern monotremes have a lower body temperature and more variable metabolic rate than
marsupials and placentals.[80] So the main question is when a monotreme-like metabolism evolved in
mammals. The evidence found so far suggests Triassic cynodonts may have had fairly high metabolic rates,
but is not conclusive.

[edit] Respiratory turbinates

Modern mammals have respiratory turbinates, convoluted structures of thin bone in the nasal cavity.
These are lined with mucous membranes that warm and moisten inhaled air and extract heat and moisture
from exhaled air. An animal with respiratory turbinates can maintain a high rate of breathing without the
danger of drying its lungs out, and therefore may have a fast metabolism. Unfortunately these bones are very
delicate and therefore have not yet been found in fossils. But rudimentary ridges like those that support
respiratory turbinates have been found in Triassic therapsids such as Thrinaxodon and Diademodon, which
suggests that they may have had fairly high metabolic rates. [76] [81][82]

[edit] Bony secondary palate

Mammals have a secondary bony palate, which separates the respiratory passage from the mouth,
allowing them to eat and breathe at the same time. Secondary bony palates have been found in the more
advanced cynodonts and have been used as evidence of high metabolic rates.[76][77][83] But some cold-
blooded vertebrates have secondary bony palates (crocodilians and some lizards), while birds, which are
warm-blooded, do not have them.[9]
 [edit] Diaphragm
A muscular diaphragm helps mammals to breathe, especially during strenuous activity. For a
diaphragm to work, the ribs must not restrict the abdomen, so that expansion of the chest can be
compensated for by reduction in the volume of the abdomen and vice versa. The advanced cynodonts have
very mammal-like rib cages, with greatly reduced lumbar ribs. This suggests that these animals had
diaphragms, were capable of strenuous activity for fairly long periods and therefore had high metabolic rates.
[76][77] On the other hand these mammal-like rib cages may have evolved to increase agility.[9] But the
movement of even advanced therapsids was "like a wheelbarrow", with the hindlimbs providing all the thrust
while the forelimbs only steered the animal, in other words advanced therapsids were not as agile as either
modern mammals or the early dinosaurs.[84] So the idea that the main function of these mammal-like rib
cages was to increase agility is doubtful.

[edit] Limb posture

The therapsids had sprawling forelimbs and semi-erect hindlimbs.[77][85] This suggests that
Carrier's constraint would have made it rather difficult for them to move and breathe at the same time, but not
as difficult as it is for animals such as lizards, which have completely sprawling limbs.[86] But cynodonts
(advanced therapsids) had costal plates that stiffened the rib cage and therefore may have reduced sideways
flexing of the trunk while moving, which would have made it a little easier for them to breathe while moving .
[87] These facts suggest that advanced therapsids were significantly less active than modern mammals of
similar size and therefore may have had slower metabolisms.
 [edit] Insulation (hair and fur)
Insulation is the "cheapest" way to maintain a fairly constant body temperature. So possession of hair
or fur would be good evidence of homeothermy but would not be such strong evidence of a high metabolic
rate.[88] [89]
The first clear evidence of hair or fur is in fossils of Castorocauda, from 164M years ago in the mid
Jurassic;[25] arguments that advanced therapsids had hair are unconvincing.

[edit] References
1. ^ a b "Amniota - Palaeos". http://www.palaeos.org/Amniota.
2. ^ a b "Synapsida overview - Palaeos".
http://www.palaeos.com/Vertebrates/Units/Unit390/000.html.
3. ^ a b Mammalia: Overview - Palaeos
4. ^ Cowen, R. (2000). History of Life. Oxford: Blackwell Science. pp. 432.
5. ^ K. A. Kermack, Frances Mussett and H. W. RIgney (January 1981). "The skull of
Morganucodon". Zoological Journal of the Linnean Society 71 (1): page 148. doi:10.1111/j.1096-
3642.1981.tb01127.x.
6. ^ "Synapsida: Varanopseidae - Palaeos".
http://www.palaeos.com/Vertebrates/Units/Unit390/200.html.
7. ^ a b "Therapsida - Palaeos".
http://www.palaeos.com/Vertebrates/Units/400Therapsida/100.html.
 8. ^ a b Kermack, D.M.; Kermack, K.A. (1984). The evolution of mammalian characters. Croom
Helm. ISBN 079915349.
9. ^ a b c d Bennett, A.F.; Ruben, J.A. (1986). "The metabolic and thermoregulatory status of
therapsids". in Hotton III, N; MacLean, P.D.; Roth, J.J. et al.. The ecology and biology of mammal-
like reptiles. Washington: Smithsonian Institution Press, Washington. pp. 207–218.
10.^ "Dinocephalia - Palaeos".
http://www.palaeos.com/Vertebrates/Units/400Therapsida/300.html.
11.^ "Neotherapsida - Palaeos".
http://www.palaeos.com/Vertebrates/Units/400Therapsida/700.html.
12.^ "Theriodontia - Paleos".
http://www.palaeos.com/Vertebrates/Units/400Therapsida/400.800.html.
13.^ a b "Cynodontia Overview - Palaeos".
http://www.palaeos.com/Vertebrates/Units/410Cynodontia/410.000.html.
14.^ Groenewald, G.H., Welman, J. and MacEachern, J.A. (April 2001). "Vertebrate Burrow
Complexes from the Early Triassic Cynognathus Zone (Driekoppen Formation, Beaufort Group) of
the Karoo Basin, South Africa". PALAIOS 16 (2): 148–160. doi:10.1669/0883-
1351(2001)016<0148:VBCFTE>2.0.CO;2.
http://palaios.sepmonline.org/cgi/content/abstract/16/2/148. Retrieved 2008-07-07.
15.^ "Olenekian Age of the Triassic - Palaeos".
http://www.palaeos.com/Mesozoic/Triassic/Olenekian.html.
16.^ "The Triassic Period - Palaeos". http://www.palaeos.com/Mesozoic/Triassic/Triassic.htm.
17.^ Ruben, J.A., and Jones, T.D. (2000). "Selective Factors Associated with the Origin of Fur
and Feathers". American Zoologist 40 (4): 585–596. doi:10.1093/icb/40.4.585.
http://icb.oxfordjournals.org/cgi/content/full/40/4/585.
 18.^ Raichle, M.E.; Gusnard, D.A. (August 6, 2002). "Appraising the brain's energy budget".
PNAS 99 (16): 10237–10239. doi:10.1073/pnas.172399499. PMID 12149485. PMC 124895.
http://www.pnas.org/cgi/content/full/99/16/10237.
19.^ "Brain power". New Scientist. 2006.
http://www.newscientist.com/blog/shortsharpscience/2006/09/brain-power.html.
20.^ Travis, J (October 2003). "Visionary research: scientists delve into the evolution of color
vision in primates". Science News 164 (15).
http://findarticles.com/p/articles/mi_m1200/is_15_164/ai_110266608.
21.^ a b Cifelli, R.L. (November 2001). "Early mammalian radiations". Journal of Paleontology
75: 1214. doi:10.1666/0022-3360(2001)075<1214:EMR>2.0.CO;2.
http://findarticles.com/p/articles/mi_qa3790/is_200111/ai_n8958762/pg_6.
22.^ Luo, Z.-X. (2007). "Transformation and diversification in early mammal evolution". Nature
450 (7172): 1011–1019. doi:10.1038/nature06277. PMID 18075580.
23.^ a b c "Mammaliformes - Palaeos".
http://www.palaeos.com/Vertebrates/Units/Unit420/420.100.html.
24.^ "Morganucodontids & Docodonts - Palaeos".
http://www.palaeos.com/Vertebrates/Units/Unit420/420.200.html.
25.^ a b c Ji, Q.; Luo, Z-X, Yuan, C-X, and Tabrum, A.R. (February 2006). "A Swimming
Mammaliaform from the Middle Jurassic and Ecomorphological Diversification of Early Mammals".
Science 311 (5764): 1123. doi:10.1126/science.1123026. PMID 16497926.
http://www.sciencemag.org/cgi/content/abstract/311/5764/1123. See also the news item at "Jurassic
"Beaver" Found; Rewrites History of Mammals".
http://news.nationalgeographic.com/news/2006/02/0223_060223_beaver.html.
 26.^ "Symmetrodonta - Palaeos".
http://www.palaeos.com/Vertebrates/Units/Unit420/420.300.html.
27.^ Luo, Z-X., Crompton, A.W., and Sun, A-L. (May 2001). "A New Mammaliaform from the
Early Jurassic and Evolution of Mammalian Characteristics". Science 292 (5521): 1535–1540.
doi:10.1126/science.1058476. PMID 11375489.
http://www.sciencemag.org/cgi/content/full/292/5521/1535. Retrieved 2008-09-08.
28.^ a b "Mammalia - Palaeos".
http://www.palaeos.com/Vertebrates/Units/430Mammalia/430.100.html.
29.^ a b Jacobs, L.L., Winkler, D.A., and Murry P.A. (July 1, 1989). "Modern Mammal Origins:
Evolutionary Grades in the Early Cretaceous of North America". Proceedings of the National
Academy of Sciences of the USA 86 (13): 4992–4995. doi:10.1073/pnas.86.13.4992. PMID 2740336.
PMC 297542. http://links.jstor.org/sici?sici=0027-
8424%2819890701%2986%3A13%3C4992%3AMMOEGI%3E2.0.CO%3B2-
9&size=LARGE&origin=JSTOR-enlargePage.
30.^ Rauhut, O.W.M., Martin, T., Ortiz-Jaureguizar, E. and Puerta, P. (14 March 2002). "A
Jurassic mammal from South America". Nature 416 (416): 165–168. doi:10.1038/416165a.
PMID 11894091. http://www.nature.com/nature/journal/v416/n6877/full/416165a.html.
31.^ a b c Rowe, T., Rich, T.H., Vickers-Rich, P., Springer, M., and Woodburne, M.O. (January
2008). "The oldest platypus and its bearing on divergence timing of the platypus and echidna
clades". Proceedings of the National Academy of Sciences 105 (4): 1238–1242.
doi:10.1073/pnas.0706385105. PMID 18216270. PMC 2234122.
http://www.pnas.org/cgi/content/full/105/4/1238.
32.^ "Molecules, morphology, and ecology indicate a recent, amphibious ancestry for echidnas".
http://www.pnas.org/content/106/40/17089.
 33.^ a b "Appendicular Skeleton". http://courses.washington.edu/chordate/453lectures/set2/453-
appendskel-06.htm.
34.^ "Mammalia: Spalacotheroidea & Cladotheria - Palaeos".
http://www.palaeos.com/Vertebrates/Units/430Mammalia/430.500.html.
35.^ Flynn, J.J., Parrish, J.M. Rakotosamimanana, B., Simpson, W.F., and Wyss, A.R.
(September 1999). "A Middle Jurassic mammal from Madagascar". Nature 401: 57–60.
doi:10.1038/43420. http://www.nature.com/nature/journal/v401/n6748/abs/401057a0.html. Retrieved
2008-09-08.
36.^ "Metatheria - Palaeos". http://www.palaeos.com/Vertebrates/Units/Unit440/440.100.html.
37.^ Szalay, F.S.; Trofimov, B.A. (1996). "The Mongolian Late Cretaceous Asiatherium, and the
early phylogeny and paleobiogeography of Metatheria" ([dead link] — Scholar search). Journal of
Vertebrate Paleontology 16 (3): 474–509. http://www.vertpaleo.org/jvp/16-474-509.html.
38.^ "Oldest Marsupial Fossil Found in China". National Geographic News. 2003-12-15.
http://news.nationalgeographic.com/news/2003/12/1215_031215_oldestmarsupial.html.
39.^ "Didelphimorphia - Palaeos".
http://www.palaeos.com/Vertebrates/Units/Unit440/440.100.html#Didelphimorphia.
40.^ "Family Peramelidae (bandicoots and echymiperas)".
http://animaldiversity.ummz.umich.edu/site/accounts/information/Peramelidae.html.
41.^ "Species is as species does... Part II". http://lancelet.blogspot.com/2005/12/species-is-as-
species-does-part-ii.html.
42.^ "Marsupials". http://paleo.amnh.org/bjburger/fossilmammal/ma3.html.
43.^ Novacek, M.J.; Rougier, G.W.; Wible, J.R.; McKenna, M.C.; Dashzeveg, D; Horovitz, I
(1997). "Epipubic bones in eutherian mammals from the late Cretaceous of Mongolia". Nature 389
(6650): 440–441. doi:10.1038/39020. PMID 9333234.
 44.^ White, T.D. (August 9, 1989). "An analysis of epipubic bone function in mammals using
scaling theory". Jornal of Theoretical Biology 139 (3): 343–57. doi:10.1016/S0022-5193(89)80213-9.
PMID 2615378.
45.^ "Eomaia scansoria: discovery of oldest known placental mammal".
http://www.evolutionpages.com/Eomaia%20scansoria.htm.
46.^ Reilly, S.M., and White, T.D. (January 2003). "Hypaxial Motor Patterns and the Function of
Epipubic Bones in Primitive Mammals". Science 299 (5605): 400–402. doi:10.1126/science.1074905.
PMID 12532019. http://www.sciencemag.org/cgi/content/full/299/5605/400. Retrieved 2008-09-24.
47.^ Novacek, M.J., Rougier, G.W, Wible, J.R., McKenna, M.C, Dashzeveg, D.,and Horovitz, I.
(October 1997). "Epipubic bones in eutherian mammals from the Late Cretaceous of Mongolia".
Nature 389 (6650): 483–486. doi:10.1038/39020. PMID 9333234.
http://www.nature.com/nature/journal/v389/n6650/full/389483a0.html. Retrieved 2008-09-24.
48.^ Fox, D (1999). "Why we don't lay eggs". New Scientist.
http://www.newscientist.com/article/mg16221904.800-why-we-dont-lay-eggs.html.
49.^ "Eutheria - Palaeos".
http://www.palaeos.com/Vertebrates/Units/450Eutheria/450.100.html#Eutheria.
50.^ Ji, Q., Luo, Z-X., Yuan, C-X.,Wible, J.R., Zhang, J-P.,and Georgi, J.A. (April 2002). "The
earliest known eutherian mammal". Nature 416 (6883): 816–822. doi:10.1038/416816a.
PMID 11976675. http://www.nature.com/nature/journal/v416/n6883/full/416816a.html. Retrieved
2008-09-24.
51.^ Luo, Z.-X., Wible, J.R. (2005). "A Late Jurassic Digging Mammal and Early Mammal
Diversification". Science 308 (5718): 103–107.. doi:10.1126/science.1108875. PMID 15802602.
 52.^ Meng, J., Hu, Y., Wang, Y., Wang, X., Li, C. (December 2006). "A Mesozoic gliding
mammal from northeastern China". Nature 444 (7121): 889–893. doi:10.1038/nature05234.
PMID 17167478. http://www.nature.com/nature/journal/v444/n7121/abs/nature05234.html.
53.^ Li, J., Wang, Y., Wang, Y., Li, C. (2000). "A new family of primitive mammal from the
Mesozoic of western Liaoning, China". Chinese Science Bulletin 46 (9): 782–785.
doi:10.1007/BF03187223. abstract, in English
54.^ Hu, Y., Meng, J., Wang, Y., Li, C. (2005). "Large Mesozoic mammals fed on young
dinosaurs". Nature 433 (7022): 149–152. doi:10.1038/nature03102. PMID 15650737.
http://www.nature.com/cgi-taf/DynaPage.taf?
file=/nature/journal/v433/n7022/abs/nature03102_fs.html&dynoptions=doi1105989664.
55.^ Wible, J.R., Rougier, G.W., Novacek, M.J., and Asher, R.J. (2007). "Cretaceous eutherians
and Laurasian origin for placental mammals near the K/T boundary". Nature 447 (447): 1003–1006.
doi:10.1038/nature05854. PMID 17581585.
http://www.nature.com/nature/journal/v447/n7147/full/nature05854.html.
56.^ Murphy, W.J., Eizirik, E., Springer, M.S et al. (14 December 2001). "Resolution of the Early
Placental Mammal Radiation Using Bayesian Phylogenetics". Science 294 (5550): 2348–2351.
doi:10.1126/science.1067179. PMID 11743200.
57.^ Kriegs, J.O., Churakov, G., Kiefmann, M., et al. (2006). "Retroposed Elements as Archives
for the Evolutionary History of Placental Mammals". PLoS Biol 4 (4): e91.
doi:10.1371/journal.pbio.0040091. PMID 16515367. (pdf version)
58.^ "Scientists map elephant evolution". BBC News. 2007-07-24.
http://news.bbc.co.uk/2/hi/science/nature/6913934.stm. Retrieved 2008-08-11.
59.^ Historical perspective (the Dynamic Earth, USGS)
60.^ Cretaceous map
 61.^ Insectivora Overview - Palaeos
62.^ Springer, M.S.; Douzery, E. (1996). "Secondary Structure and patterns of evolution among
mammalian mitochondrial 12S rRNA molecules". J. Mol. Evol. 43 (4): 357–373.
doi:10.1007/BF02339010. PMID 8798341.
63.^ Springer, M.S.; Hollar, L.J.; Burk, A. (1995). "Compensatory substitutions and the evolution
of the mitochondrial 12S rRNA gene in mammals". Mol. Biol. Evol. 12 (6): 1138–1150.
PMID 8524047.
64.^ Li, W-H (1997). Molecular Evolution. Sinauer Associates.
65.^ Bininda-Emonds, O.R.P.; Cardillo, M.; Jones, K.E.; 'et al.', Ross D. E.; Beck, Robin M. D.;
Grenyer, Richard; Price, Samantha A.; Vos, Rutger A. et al. (2007). "The delayed rise of present-day
mammals". Nature 446 (446): 507–511. doi:10.1038/nature05634. PMID 17392779.
http://scienceblogs.com/pharyngula/2007/03/dont_blame_the_dinosaurs.php.
66.^ Dinosaur Extinction Spurred Rise of Modern Mammals
67.^ Benton, M.J. (December 1999). "Early origins of modern birds and mammals: molecules vs.
morphology". BioEssays 21 (12): 1043–1051. doi:10.1002/(SICI)1521-1878(199912)22:1<1043::AID-
BIES8>3.0.CO;2-B. PMID 10580989.
68.^ Archibald, J.D. (May 1996). "Fossil Evidence for a Late Cretaceous Origin of "Hoofed"
Mammals". Science 272 (5265): 1150–1153. doi:10.1126/science.272.5265.1150. PMID 8662448.
http://www.sciencemag.org/cgi/content/abstract/272/5265/1150. Retrieved 2008-09-08.
69.^ Martin, R.D.; Soligo, C.; Tavaré, S. (2007). "Primate Origins: Implications of a Cretaceous
Ancestry" (PDF). Folia Primatologica 78 (78): 277–296. doi:10.1159/000105145. PMID 17855783.
http://content.karger.com/ProdukteDB/produkte.asp?
Aktion=ShowPDF&ArtikelNr=105145&Ausgabe=233328&ProduktNr=223842&filename=105145.pdf.
— a similar paper by these authors is free online at New light on the dates of primate origins and
divergence
70.^ Alroy J. (March 1999). "The fossil record of North American mammals: evidence for a
Paleocene evolutionary radiation". Systematic biology 48 (1): 107–18.
doi:10.1080/106351599260472. PMID 12078635.
71.^ Archibald J.D., and Deutschman D.H. (June 2001). "Quantitative Analysis of the Timing of
the Origin and Diversification of Extant Placental Orders". Journal of Mammalian Evolution 8 (2): 107–
124. doi:10.1023/A:1011317930838.
http://www.ingentaconnect.com/content/klu/jomm/2001/00000008/00000002/00342277. Retrieved
2008-09-24.
72.^ Rich, T.H., Hopson, J.A., Musser, A.M., Flannery., T.GF., and Vickers-Rich, P. (11
February 2005). "Independent Origins of Middle Ear Bones in Monotremes and Therians". Science
307 (5711): 910–914. doi:10.1126/science.1105717. PMID 15705848.
http://www.sciencemag.org/cgi/content/abstract/307/5711/910. For other opinions see "Technical
comments" linked from same Web page
73.^ Oftedal, O.T. (2002). "The mammary gland and its origin during synapsid evolution".
Journal of Mammary Gland Biology and Neoplasia 7 (3): 225–252. doi:10.1023/A:1022896515287.
PMID 12751889.
74.^ Oftedal, O.T. (2002). The origin of lactation as a water source for parchment-shelled
eggs=Journal of Mammary Gland Biology and Neoplasia. 7. pp. 253–266.
75.^ Lactating on Eggs
76.^ a b c d Brink, A.S. (1955). "A study on the skeleton of Diademodon". Palaeontologia
Africana 3: 3–39.
 77.^ a b c d Kemp, T.S. (1982). Mammal-like reptiles and the origin of mammals. London:
Academic Press. pp. 363.
78.^ Estes, R. (1961). "Cranial anatomy of the cynodont reptile Thrinaxodon liorhinus". Bulletin
of the Museum of Comparative Zoology (1253): 165–180.
79.^ Kielan−Jaworowska, Z.; Hurum, J.H.. (2006). "Limb posture in early mammals: Sprawling
or parasagittal" (PDF). Acta Palaeontologica Polonica 51 (3): 10237–10239.
http://www.app.pan.pl/archive/published/app51/app51-393.pdf. Retrieved 2008-09-24.
80.^ Paul, G.S. (1988). Predatory Dinosaurs of the World. New York: Simon and Schuster.
pp. 464.
81.^ Hillenius, W.H. (1992). "The evolution of nasal turbinates and mammalian endothermy".
Paleobiology 18 (1): 17–29.
82.^ Ruben, J. (1995). "The evolution of endothermy in mammals and birds: from physiology to
fossils". Annual Review of Physiology 57: 69–95. doi:10.1146/annurev.ph.57.030195.000441.
PMID 7778882.
83.^ McNab, B.K. (1978). "The evolution of endothermy in the phylogeny of mammals".
American Naturalist 112: 1–21. doi:10.1086/283249.
84.^ Ccowen, R. (2000). History of Life. Oxford: Blackwell Science. pp. 432.
85.^ Jenkins, F.A., Jr (1971). "The postcranial skeleton of African cynodonts". Bulletin of the
Peabody Museum of Natural History (36): 1–216.
86.^ Pough, F.H; Heiser, J.B.; McFarland, W.N. (1996). Vertebrate Life. New Jersey: Prentice-
Hall. pp. 798.
87.^ Sidor, C.A.; Hopson, J.A. (1998). "Ghost lineages and "mammalness": assessing the
temporal pattern of character acquisition in the Synapsida". Paleobiology (24): 254–273.
 88.^ Schmidt-Nielsen, K. (1975). Animal physiology: Adaptation and environment. Cambridge:
Cambridge University Press. pp. 699.
89.^ Withers, P.C. (1992). Comparative Animal Physiology. Fort Worth: Saunders College.
pp. 949.

[edit] Bibliography
• Robert L. Carroll, Vertebrate Paleontology and Evolution, W. H. Freeman and Company,
New York, 1988 ISBN 0-716-71822-7. Chapters XVII through XXI
• Nicholas Hotton III, Paul D. MacLean, Jan J. Roth, and E. Carol Roth, editors, The Ecology
and Biology of Mammal-like Reptiles, Smithsonian Institution Press, Washington and London, 1986
ISBN 0-87474-524-1
• T. S. Kemp, The Origin and Evolution of Mammals, Oxford University Press, New York, 2005
ISBN 0-19-850760-7
• Zofia Kielan-Jaworowska, Richard L. Cifelli, and Zhe-Xi Luo, Mammals from the Age of
Dinosaurs: Origins, Evolution, and Structure, Columbia University Press, New York, 2004 ISBN 0-
231-11918-6. Comprehensive coverage from the first mammals up to the time of the K-T mass
extinction.
• Zhe-Xi Luo, "Transformation and diversification in early mammal evolution", Nature volume
450 number 7172 (13 December 2007) pages 1011–1019. doi:10.1038/nature06277. A survey article
with 98 references to the scientific literature.
 [edit] External links
• The Cynodontia covers several aspects of the evolution of cynodonts into mammals, with
plenty of references.
Retrieved from "http://en.wikipedia.org/wiki/Evolution_of_mammals"

Categories: Evolution by taxon | Mammals | Synapsids | Mammalogy | Paleontology | Mammal

anatomy | Prehistoric mammals
 W000

Histoire évolutive des mammifères

Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Fichier:Thrinaxo BW.jpg
Reconstitution d'un Thrinaxodon, un membre du groupe des cynodontes qui inclut les ancêtres des
mammifères.
Selon la théorie de l'évolution, les mammifères sont issus des synapsides (des reptiles mammaliens)
selon un processus graduel qui a pris approximativement 70 Ma, du milieu du Permien jusqu'au milieu du
Jurassique, et au cours du Trias.
 D’un point de vue purement cladistique, les mammifères sont alors vus comme les survivants des
synapsides.
Sommaire
[masquer]
• 1 Les ancêtres des mammifères
• 1.1 Les amniotes
• 1.2 Les synapsides
• 1.3 Les thérapsides
• 1.3.1 Arbre d'évolution des thérapsides
• 2 Le tournant du Trias
• 3 Des cynodontes aux vrais mammifères
• 3.1 Des incertitudes
• 3.2 Mammifères ou mammaliens ?
• 3.3 Arbre phylogénique des cynodontes vers les
mammifères[15].
• 3.3.1 Multituberculés
• 3.3.2 Les Morganucodontes
• 3.3.3 Les Docodontes
• 3.3.4 Hadrocodium
• 4 Les premiers vrais mammifères
• 4.1 Arbre d’évolution des premiers vrais
mammifères
• 4.2 Australosphenida et Ausktribosphenidae
Les ancêtres des mammifères [modifier]
Voici ci-dessous l’arbre phylogénique (d'évolution) très simplifié des mammifères. Nous en verrons
ensuite les détails et les incertitudes qui font encore débat.
─o Tétrapodes
├─o Amphibiens
└─o Amniotes
├─o Sauropsides (Oiseaux, Crocodiliens, ...)
└─o Synapsides
└─o Thérapsides
└─o Cynodontes
└─o Mammifères

Les amniotes [modifier]

Contrairement à la plupart des amphibiens[1], les Amniotes ont pu s'affranchir de pondre dans l'eau
grâce à l'amnios, un sac qui protège l'embryon de la déshydratation. Les premiers amniotes sont connus dès
-312 Ma[2], c'est-à-dire au milieu du Carbonifère.
En quelques millions d'années, deux branches se sont distinguées :
• les Synapsides, qui incluent actuellement les mammifères ;
• les Sauropsides représentés aujourd'hui par les tortues, les lézards, les serpents, les
crocodiles et les oiseaux (les seuls survivants des dinosaures).
 Les synapsides [modifier]

Le crâne d’un synapside a une fosse temporale en position inférieure (en bas à droite sur cette
image).
La présence de fosses temporales est avantageuse car elle permet :
• d’avoir un crâne plus léger et résistant ;
• de probablement améliorer l’insertion des muscles de la mâchoire. Étant plus loin de la
mâchoire, les muscles peuvent être plus longs et donc exercer une plus grande force de traction
avec une plus grande amplitude de mouvement.
Les fossiles terrestres du début du Permien indiquent que les premiers synapsides, les
"Pélycosaures", étaient les vertébrés terrestres les plus communs et les plus répandus en leur temps[3].
Les thérapsides [modifier]
Le groupe des Thérapsides s'est différencié de celui des "pélycosaures" au milieu du Permien et
constituèrent la grande majorité des tétrapodes jusqu'à la fin de cette période. Ils diffèrent des
"pélycosaures" par leur crâne et leur mâchoire, dont la taille plus importante de la fosse temporale et des
incisives de même taille[4].
Les Thérapsides évoluèrent en plusieurs étapes, commençant par des animaux ressemblant à leurs
lointains parents "Pélycosaures" et aboutissant à des animaux ressemblant beaucoup aux mammifères :
• Développement progressif de l’os secondaire du palais osseux[5] qui permettrait une
mastication plus efficace.
La plupart des ouvrages et articles de référence interprètent cette évolution comme un prérequis
vers le métabolisme élevé des mammifères, parce qu’il est ainsi possible pour ces animaux de manger et de
respirer en même temps. Mais quelques scientifiques argumentent que certains ectothermes utilisent un
palais secondaire charnu pour séparer la bouche des voies aériennes et que l'avantage d'un palais "osseux"
est qu'il fournit une surface dure sur laquelle la langue peut manipuler les aliments, facilitant plutôt la
mastication que la respiration[6].
Chez les mammifères, le palais est formé par 2 os spécifiques, mais plusieurs thérapsides du
Permien ont d’autres combinaisons d’os qui font office de palais.
Mais aussi :
• le dentaire (anciennement "os maxillaire inférieur") devient progressivement l’os principal de
la mâchoire inférieure ;
 • une tendance vers des membres para-sagittaux (en dessous du corps) qui augmentent
l’endurance des animaux en limitant les contraintes mécaniques.
Mais cette évolution est irrégulière, par exemple : tous les thérapsides herbivores conservent des
membres latéraux (certaines formes tardives pourraient avoir des membres arrières semi-droits) ; les
thérapsides carnivores du Permien ont des membres antérieurs latéraux, alors certains de la fin du Permien
ont des membres antérieurs semi-érigés.
En fait, les monotrèmes modernes (ornithorynque par exemple) ont toujours des membres semi-érigés.
On note aussi :
• au Trias, des progrès vers la mâchoire et l’oreille interne de type "mammifère" ;
• des éléments plausibles d’existence de poils pour les thérapsides du Trias [réf. souhaitée] mais
pas pour ceux du Permien ;
• certains scientifiques affirment que des signes de lactation sont présents sur les thérapsides
du Trias.[réf. souhaitée]

Arbre d'évolution des thérapsides [modifier]

On ne donne ici qu'un arbre simplifié[4]: seuls les thérapsides les plus importants dans l’évolution
vers les mammifères sont décrits ci-dessous.
─o Thérapsides
├─o Biarmosuchia
└─o
├─o Dinocephalia
 └─o Neotherapsida
├─o Anomodontia
│ └─o Dicynodontes
└─o Theriodontia
├─o Gorgonopsia
└─o Eutheriodontia
└─o Cynodontes
:
└─o Mammifères

Seuls les dicynodontes, thérocéphales et les cynodontes survivront jusqu’à la fin du Trias où ils
disparaissent durant la période d'extinction massive du Trias-Jurassique, hormis les quelques Cynodontes
qui donneront naissance aux Mammifères.
Les Theriodontia disposent d'articulations de mâchoire où l’articulaire est solidement attaché à l’os
carré du crâne. La réduction de taille de l’os carré est une étape importante vers le développement de la
mâchoire qui permet une plus grande ouverture et de la formation de l'oreille interne vers le type partagé par
tous les mammifères. Un taxon de carnivores, les gorgonopsiens, taxon frère des Eutheriodontia dont sont
issus les mammifères, ont pu développer « des impressionnantes dents de sabre ».
Le groupe des cynodontes issu des Eutheriodontia est apparu à la fin du Permien. L’ancêtre de tous
les mammifères appartient à ce groupe. Le sous-groupe le plus souvent considéré est celui des
trithélodontes.
 Les caractéristiques qui rapprochent les cynodontes des mammifères sont la réduction du nombre
d’os de la mâchoire inférieure, la présence d'un palais secondaire osseux, des dents jugales avec un modèle
complexe en couronnes et une grande cavité endocrânienne[7].

Le tournant du Trias [modifier]

La période d'extinction massive du Trias-Jurassique a éliminé environ 70% des espèces de
vertébrés terrestres et la majorité des plantes de surface, avec comme conséquences[8]:
• l’effondrement de l’écosystème et de la chaîne alimentaire dont le rétablissement se fit en 6
millions d’années environ ;
• les espèces dominantes furent contestées dans leur niche écologique, y compris les
cynodontes qui semblaient être en position de force à la fin du Permien.
Mais les cynodontes furent éclipsés par un antique et obscur groupe issu des sauropsidiens, les
archosauriens dont sont membres les crocodiles, les ptérosaures et les dinosaures - donc par conséquent
les oiseaux.
Plusieurs explications sont avancées pour cette extinction massive, mais la plus populaire est que le
début du Trias était plutôt aride et que les archosauriens avaient une supériorité dans la gestion de leur eau.
En effet, tous les sauropsides connus ont une peau avec moins de glandes et excrètent de l’acide urique au
lieu de l'urée (comme les mammifères et donc par déduction les thérapsides) qui nécessite moins d’eau pour
rester suffisamment liquide[9].
 Le tournant du Trias a été progressif. Dans les premiers temps du Trias, les cynodontes étaient les
principaux prédateurs et les Lystrosaurus les principaux herbivores, mais à la mi-Trias, les archosauriens
dominent toutes les grandes niches écologiques aussi bien herbivores que carnassiers.
Mais le tournant du Trias a été un élément moteur[10] de l’évolution des cynodontes en mammifères.
Les seuls descendants des cynodontes ayant réussi à survivre furent des petits, principalement nocturnes,
insectivores[7].
Le fait d’être des chasseurs nocturnes insectivores a eu comme conséquences :
• l’accélération de la tendance des thérapsides vers des dents différenciées spécialisées et
précisément localisées, à cause de la nécessité de capturer des arthropodes et d’écraser leur
carapace (exosquelette) ;
• la vie nocturne nécessite des avancées en termes d’isolation et régulation thermique pour
pouvoir être actif dans la fraicheur de la nuit ;
• l’acuité des sens devient vitale, notamment ceux de l’ouïe et de l’odorat :
• l’évolution vers l’oreille interne de mammifère a été accélérée, et simultanément celle
de la mâchoire, puisque des os de la mâchoire sont devenus des os de l’oreille interne ;
• l’augmentation de la taille des lobes cérébraux spécialisés dans le traitement des
sens. Les tissus cérébraux ayant une énorme consommation d’énergie[11], [12],
l’augmentation de la contrainte alimentaire favorisa les évolutions sur l’isolation (poils), sur la
régulation thermique et sur la nutrition.
• comme effet négatif, la vue devient moins importante (puisque vivant la nuit) et ceci se
retrouve dans le fait que la plupart des mammifères ont une vision des couleurs médiocre, y compris
les petits primates comme les lémuriens[13].
 Des cynodontes aux vrais mammifères [modifier]

Des incertitudes [modifier]

Bien que le « Tournant du Jurassique » ait probablement accéléré l’évolution des mammifères, il est
difficile pour les paléontologues de le constater car les bons fossiles de quasi-mammifères sont extrêmement
rares, principalement à cause de leur très petite taille (plus petits qu’un rat actuel en majorité) et de leurs
niches écologiques de prédilection :
• Les mammifères ont été largement limités aux zones environnementales qui sont peu
propices aux bons fossiles. Les meilleures conditions de fossilisations sur la terre sèche sont les
plaines régulièrement inondées. Les inondations saisonnières recouvrent rapidement les cadavres
d’une couche de boue protectrice qui est plus tard compressée en roche sédimentaire. Mais ces
plaines humides sont dominées par des animaux de taille moyenne à grande, et au Trias les derniers
thérapsides et les quasi-mammifères ne pouvaient pas rivaliser avec les archosauriens dans cette
catégorie de taille ;
• Les os des petits animaux sont plus fragiles et sont souvent détruits par les charognards (y
compris bactéries et moisissures) ou par écrasement et dispersion avant qu'ils soient fossilisés ;
• Les petits fossiles sont plus difficiles à découvrir et plus vulnérables aux agressions de la
nature avant qu’on les découvre.
En fait, on dit, qu’il n’y a pas si longtemps, dans les années 1980, la totalité des fossiles de quasi-
mammifères pouvait tenir dans une boite à chaussures et que la plupart étaient des dents qui sont
plus résistantes que les os[14].
En conséquence :
 • Dans de nombreux cas, il est difficile d’assigner un mammifère du Mésozoïque ou un fossile
de presque mammifère à un genre ;
• Tous les fossiles disponibles d'un genre font rarement un squelette complet et il est donc
difficile de décider quels sont les genres se ressemblant l'un l'autre et donc de là, très probablement
proches parents. Autrement dit, il devient très difficile de les classifier au moyen de la cladistique, qui
est la méthode, actuellement disponible, la moins subjective et la plus fiable.
Donc l'évolution de mammifères dans le Mésozoïque est pleine d'incertitudes, bien qu'il soit certain
que les vrais mammifères (de type actuel) soient apparus au Mésozoïque.

Mammifères ou mammaliens ? [modifier]

Une conséquence de ces incertitudes a été la modification de la définition de « mammifères » par les
paléontologues. Pendant longtemps, un fossile était considéré comme un mammifère s’il possédait le critère
de mâchoire et d’oreille interne vu précédemment (l’articulation de la mâchoire des mammifères est
seulement composée de os temporal et de l’os mandibule, alors que l’os articulaire et l’os carré sont devenus
l'enclume et le marteau de l’oreille interne moyenne).
Mais plus récemment, les paléontologues ont défini les mammifères comme le plus récent ancêtre
commun aux monotrèmes, marsupiaux et placentaires, ainsi que tous leurs descendants. Les
paléontologues ont dû définir un autre clade : les mammaliens (ou mammaliformes) pour classifier tous les
animaux qui sont plus ressemblants aux mammifères que les cynodontes mais plus éloignés que les
monotrèmes, marsupiaux ou placentaires[15].
Bien que cela semble maintenant être l'approche majoritaire, certains paléontologues ne l’admettent
pas parce qu’il ne fait que déplacer la plupart des problèmes dans un nouveau clade sans les résoudre. Ainsi
ce clade mammaliformes inclut quelques animaux avec des joints de mâchoire "mammifères" et certains
avec des joints de mâchoire (os articulaire-os carré) "reptiliens" ; et la nouvelle définition "de mammifère" et
"mammaliformes" dépend des derniers ancêtres communs des deux groupes qui n'ont pas encore été
trouvés[16]. Malgré ces réserves, on adoptera ici l’approche majoritaire et on appellera mammaliformes la
plupart des descendants des cynodontes du Mésozoïque.

Arbre phylogénique des cynodontes vers les mammifères[15]. [modifier]

--Cynodontes
|
`--Mammaliformes
|
+--Allotheria
| |
| `--Multituberculata
|
`--+--Morganucodontidae
|
`--+--Docodontes
|
`--+--Hadrocodium
|
`--Symmetrodonta
|
|--Kuehneotheriidae
 |
`--Mammifères

Multituberculés [modifier]
Les Multituberculés, appelés ainsi pour les nombreuses bosses sur leurs molaires, sont aussi
nommés "les rongeurs du Mésozoïque". C'est un excellent exemple de "l’expérimentation" de l’évolution
pendant cette ère.
À première vue, ils ressemblent aux mammifères modernes : leurs joints de mâchoire consistent en
seulement l'os mandibule et l'os squamosal, et les os carré et articulaires font partie de l’oreille interne
moyenne ; leurs dents sont différenciées. Ils ont des os zygomatique (les pommettes) ; la structure de leur
bassin osseux suggère qu’ils donnaient naissance à des petits minuscules et vulnérables, comme les
marsupiaux modernes. Ils ont vécu pendant un temps incroyablement long, plus de 120 millions d'années
(du milieu du Jurassique, il y a 160 millions d’années, au début de l’Oligocène, il y a 35 millions d’années. Ce
qui en aurait fait les mammifères les plus réussis jusqu'alors.
Mais une étude plus approfondie montre qu'ils diffèrent beaucoup des mammifères modernes[15]:
• leurs "molaires" ont deux rangées parallèles de bosses, à la différence des molaires
tribosphéniques (à trois pointes) des premiers mammifères ;
• la mastication est complètement différente. Les mammifères mâchent avec un mouvement
latéral qui fait que d'habitude les molaires s’emboîtent sur seulement un côté à la fois.
Les mâchoires des Multituberculés étaient incapables de ce mouvement latéral. Ils mâchaient en
frottant leurs dents inférieures d’avant en arrière contre les supérieures comme si la mâchoire était fermée.
 • la partie antérieure (devant) de l'arc zygomatique est formé essentiellement de l'os maxillaire
(anciennement os maxillaire supérieur) plutôt que l'os jugal et le jugal est un petit os dans un petit
créneau dans le processus maxillaire (l'extension) ;
• l’os temporal ne fait pas partie du crâne ;
• le rostre (le museau) est différent de celui de mammifères, en fait il ressemble plutôt à celui
d’un pélycosaurien comme le Dimétrodon. Le museau des Multituberculés est semblable à une boite,
avec les grands maxillaires plats formant les côtés, l’os nasal le sommet et l'os prémaxillaire formant
le devant.

Les Morganucodontes [modifier]

Les premiers Morganucodontes apparaissent à la fin du Trias, il y a environ 250 millions d’années.
Ils sont un excellent exemple de forme transitionnelle puisqu’ils ont aussi bien la jonction de mâchoire os
mandibule-os temporal que articular-os carré[17]. Ils sont aussi l’un des premiers mammaliformes trouvés et
aussi l’un des plus étudiés depuis qu’un nombre inhabituel de fossiles ont été découverts.

Les Docodontes [modifier]

Le membre le plus caractéristique des docodontes est le Castorocauda '("queue de castor")'. Ces
animaux ont vécu au milieu du Jurassique il y a 164 millions d’années. Ils ont été découverts en 2004 et
décrits en 2006[18].
Le Castorocauda n’était pas un docodonte typique (la plupart étaient omnivores) et pas non plus un
vrai mammifère, mais il est extrêmement important dans l’étude de l’évolution des mammifères parce le
premier fossile trouvé était presque complet (un vrai luxe en paléontologie) et cela remis en cause le
stéréotype « petit insectivore nocturne » du mammifère :
• il est visiblement plus grand que la plupart des fossiles mammaliformes du Mésozoïque-
environ 43 cm du nez à la pointe de la queue (queue de 13 cm), et aurait pesé plus de 800
grammes ;
• il donne la preuve irréfutable de la plus ancienne présence de poils actuellement connue.
Auparavant, le fossile « à poils » le plus ancien était le Eomaia, un vrai mammifère d’il y a 125
millions d’années ;
• il a des caractéristiques d’adaptation aquatique comme les os de la queue plats, et des
tissus mous entre les orteils suggérant qu’ils étaient palmés. Avant sa découverte, le plus ancien
mammiforme semi-aquatique connu datait du miocène, 110 millions d’années plus tard ;
• les puissantes pattes avant du Castorocauda semblent faites pour creuser. Cette
caractéristique ainsi que les éperons sur ses chevilles le font ressembler à l'ornithorynque, qui lui
aussi nage et creuse ;
• ses dents semblent adaptées pour manger du poisson : les 2 premières molaires sont
cuspides (bosselées) et rangées en ligne droite. Elles semblent plus appropriées pour saisir et
couper que pour broyer. Ces molaires sont orientées vers l’arrière pour aider dans la prise d’une
proie glissante.

Hadrocodium [modifier]
L’arbre phylogénétique de consensus de la famille vu au début de chapitre, montre "Hadrocodium"
comme "un oncle" de vrais mammifères, tandis que les symmetrodontes et kuehneotheriides sont liés plus
directement aux vrais mammifères.
 Mais les fossiles de Symmetrodontes et de Kuehneothériens sont si peu nombreux et fragmentaires
que ces animaux sont mal compris et pourraient être paraphylétiques. Alors qu’il y a les bons fossiles
d’"Hadrocodium" qui montrent quelques importantes caractéristiques[19]:
• la jonction de la mâchoire est seulement composée de l'os temporal et l’os mandibule
(anciennement maxillaire inférieure), et, à l’opposé de la structure thérapside, la mâchoire ne
contient plus de petits os à l’arrière de l’os mandibule ;
• chez les Thérapsides et la plupart des Mammaliformes, le tympan est tendu sur une cuvette
à l'arrière de la mâchoire inférieure. Mais "Hadrocodium" n'avait pas une telle cuvette, ce qui suggère
que son oreille fait partie du crâne, comme chez les mammifères - et de là que les anciens os
articulaire et carré ont migré dans l’oreille interne moyenne et sont devenus le marteau et l’enclume.
D'autre part, la mâchoire inférieure a une " baie " à l'arrière absente chez les mammifères. Cela
suggère que l'os mandibule "de l'Hadrocodium's" ait conservé la même forme qu'il aurait eue si l'os
articulaire et l’os carré étaient restés dans la partie du joint de mâchoire et donc que "Hadroconium"
ou un ancêtre très proche aurait été le premier à avoir une oreille interne moyenne entièrement
mammifère.
• les Thérapsides et premiers mammaliens avaient leur articulation de mâchoire très en arrière
dans le crâne, en partie parce que l'oreille était derrière la mâchoire, mais devait aussi être proche
du cerveau. Cette organisation limitait la taille du crâne, parce qu'il obligeait les muscles de mâchoire
d'être autour et au-dessus. Du fait de l’évolution de l’oreille, le crâne de l’"Hadrocodium" et ses
mâchoires n’avaient plus cette contrainte et l’articulation de la mâchoire put être plus sur le devant.
Chez ses descendants ou ceux d'animaux ayant une organisation semblable, la boite crânienne était
libre de s'étendre sans être gênée par la mâchoire, et réciproquement la mâchoire était libre de
changer sans être contrainte par le besoin de garder l'oreille près du cerveau.
 Autrement dit, il était maintenant possible pour les animaux mammaliens de développer une plus
grande intelligence, d’adapter leurs mâchoires et de spécialiser leurs dents à leur nourriture.

Les premiers vrais mammifères [modifier]

Cette partie de l’histoire apporte de nouvelles complications, car les vrais mammifères sont le seul
groupe qui a encore des représentants vivants :
• on doit distinguer les groupes éteints et ceux qui ont des représentants vivants ;
• on se sent souvent contraint d’essayer d’expliquer l'évolution des caractéristiques qui
n'apparaissent pas dans des fossiles. Cet effort implique souvent la phylogénétique moléculaire, une
technique qui est devenue populaire depuis le milieu des années 1980, mais qui est souvent
controversée à cause de ses hypothèses de départ, et particulièrement celle de l’horloge
moléculaire.

Arbre d’évolution des premiers vrais mammifères [modifier]

(à partir de Mammalia: Overview - Palaeos)
--Mammifères
|
+--Australosphenida
| |
| +--Ausktribosphenidae (éteint)
| |
| `--Monotrèmes
 |
`--+--Triconodonta (éteint)
|
`--+--Spalacotheroidea (éteint)
|
`--Cladotheria
|
|--Dryolestoidea (éteint)
|
`--Thériens
|
+--Métathériens
|
`--Euthériens

Australosphenida et Ausktribosphenidae [modifier]

Ausktribosphenidae est le nom d’un groupe problématique dans la vision actuelle de l’évolution[20]:
• ils ont apparemment des molaires tribosphéniques (3 pointes), un type de dent seulement
connu chez les placentaires ;
• on les a trouvés dans les couches du milieu du Crétacé en Australie. Mais l’Australie n’a été
en contact qu’avec l’Antarctique, alors que les placentaires sont originaires de l’hémisphère nord où
il furent confinés jusqu’à ce que la dérive des continents crée des ponts entre l’Amérique du Nord et
l’Amérique du Sud, entre l’Asie et l’Afrique et entre l’Asie et l’Inde.
 La carte du monde au Crétacé tardif (Late Cretaceous 94Ma) montre que les continents du sud sont
isolés ;
• les seuls fossiles disponibles sont des fragments de mâchoire et de crâne, ce qui n’est pas
très significatif.
Autrement dit, la distinction entre les groupes Ausktribosphenidae et monotrèmes pourrait être juste
une fiction, à moins que quelqu’un trouve un fossile qui pourrait être décrit précisément comme un
« australosphénide basique ». C'est-à-dire qu’il aurait des caractéristiques communes aux
Ausktribosphenidae et aux monotrèmes, ainsi que des manques par rapport aux caractéristiques communes
de ces descendants Ausktribosphenidae et monotrèmes, et pour finir des manques par rapport aux
caractéristiques spécifiques de ces derniers.

Les Monotrèmes [modifier]

Le plus ancien monotrème connu est le Teinolophos, qui vivait il y a environ 123 millions d’années
en Australie. Les monotrèmes ont des caractéristiques directement hérités de leurs ancêtres amniotes :
• ils ont un cloaque, c’est-à-dire un unique orifice pour uriner, déféquer et se reproduire
(monotrème signifie mono-trou) comme les lézards et les oiseaux ;
• ils déposent des œufs mous non calcifiés qui sont comme du cuir, toujours comme les
lézards, tortues ou crocodiles.
• à la différence des autres mammifères, les monotrèmes femelles n'ont pas de mamelon et
alimentent leurs petits en "suant" le lait sur des « champs mammaires » localisés sur leurs ventres.
Bien sûr, ces caractéristiques ne sont pas visibles à partir de fossiles, et les principaux critères
retenus par les paléontologues sont[21]:
 • un os mandibule fin dans lequel le processus coronoïde de la mandibule est petit ou
inexistant ;
• l’ouverture externe des oreilles se trouve sur la partie postérieure de la mâchoire ;
• l'os jugal est petit ou inexistant ;
• une ceinture scapulaire primitive avec de puissants éléments de respiration : with strong ventral
elements ⇔ avec de puissants éléments de respiration --- [pour ventral, voir Axe dorso-ventral ]
coracoïdes, clavicules et interclavicules. interclavicle ⇔ interclavicule --- [oui]
A Noter que les mammifères de type Thérien n’ont pas d’interclavicules[22].
• des membres antérieurs (avant) latéraux ou semi-latéraux.

Les Thériens [modifier]

Les Thériens (du grec ancien θηρίον / thêrion, animal sauvage, est le nom du groupe réunissant les
métathériens (qui portent leurs petits dans une poche externe comme les marsupiaux) et les euthériens ou
placentaires (qui portent leurs petits en interne dans un placenta). Bien qu’il n’y ait pas de fossiles
convaincants d’un Thérien basique (juste quelques dents et fragments de mâchoires), métathériens et
euthériens partagent quelques caractéristiques qui pourraient être héritées d'un groupe héréditaire
commun[23]:
• pas d’interclavicules[24];
• des os coracoïdes inexistants ou fusionnés avec les omoplates pour former le processus
coracoïde ;
• molaires tribosphéniques ;
 • un type de cheville crurotarsale : l’articulation principale est entre le tibia et l’astragale ; le
calcanéum n'a aucun contact avec le tibia, mais forme un talon auquel les muscles peuvent
s’attacher.
L'autre type bien connu de cheville crurotarsale se trouve chez les crocodiliens et qui fonctionne
différemment ; la plupart de la courbure de la cheville se situe entre le calcanéum et l'astragale).

Les Métathériens [modifier]

Les Métathériens encore vivants de nos jours, sont tous des marsupiaux (du grec marsipos « sac »).
Quelques fossiles du genre comme le mongol Asiatherium de la fin du Crétacé pourraient être des
marsupiaux ou membres d’un autre groupe métathérien[25], [26].
Le plus ancien marsupial connu est le Sinodelphys, trouvé dans du schiste argileux du début du
Crétacé (daté de 125 Ma) de la province Liaoning au nord-est de la Chine. Le fossile est presque complet et
inclut les touffes de fourrure et les empreintes de tissus mous[27].
Didelphimorphia (les opossums communs de l'hémisphère sud et de l’Amérique du Nord) est d'abord
apparu à la fin du Crétacé et a toujours des représentants vivants, probablement parce qu'ils sont surtout
semi-arboricoles, plantigrades et omnivores non spécialisés[28].
La caractéristique la mieux connue des marsupiaux est leur méthode de reproduction :
• la mère développe une sorte de vésicule vitelline dans son utérus qui livre des substances
nutritives à l’embryon. Les embryons de bandicoots, les koalas et les wombats forment en plus un
organe semblable au placenta, bien que plus petit que chez les placentaires qui les connectent à la
 paroi utérine. Il n'est néanmoins pas certain qu'ils transfèrent des substances nutritives de la mère à
l'embryon[29];
• la grossesse est très courte, typiquement de 4 à 5 semaines. L'embryon né à un stade où
commence réellement son développement et mesure d'habitude moins de 5 cm à la naissance. Il a
été suggéré que la grossesse courte est nécessaire pour réduire le risque d’attaque de l’embryon par
le système immunitaire de la mère ;
• les nouveau-nées marsupiaux utilisent les pattes avant (des mains relativement fortes) pour
monter jusqu’à un mamelon, qui est d'habitude dans une poche sur le ventre de la mère. La mère
alimente le bébé en contractant des muscles sur les glandes mammaires, car le bébé est trop faible
pour téter.
Le fait que le nouveau-né marsupial doive utiliser ses pattes avant pour atteindre le mamelon a
probablement empêché l’évolution des pattes avant vers des pagaies ou vers des ailes et a donc
empêché l'apparition de marsupiaux aquatiques ou vraiment volants (bien qu'il y ait plusieurs
marsupiaux planants).
Bien que certains marsupiaux aient une apparence très proche des placentaires, (comme le loup de
Tasmanie), les squelettes des marsupiaux ont quelques caractéristiques spécifiques qui les distinguent des
placentaires[30]:
• certains dont le “loup de Tasmanie”, ont 4 molaires. Les non-placentaires en ont plus de 3.
• une paire de fenêtres palatales, ouvertures semblables à la fenêtre sur le bas du crâne (en
plus des plus petites ouvertures des narines).
• les marsupiaux ont aussi une paire d'os spéciaux (parfois appelé "os épipubis"), qui
supportent la poche chez des femelles. Mais ceux-ci ne sont pas absolument spécifiquement
marsupiaux, puisqu'ils ont été trouvés sur des fossiles de multituberculaires, monotrèmes et même
 euthériens - donc ils ont probablement une caractéristique héréditaire commune qui a disparu à un
certain point après que les ancêtres des mammifères placentaires modernes aient divergé des
marsupiaux[31], [32].

Les Euthériens [modifier]

Les Euthériens actuellement vivants sont tous des placentaires. Mais le tout premier euthérien
connu, Eomaia[33], découvert en Chine et daté de 125 millions d’années, a des caractéristiques qui
ressemblent plus à celles des marsupiaux (les derniers descendants des métathériens)[34]:
• les épipubis se prolongent vers l’avant du bassin, ce qui ne se retrouve pas chez les
placentaires modernes, mais présents chez les marsupiaux, les monotrèmes et les mammaliens
comme les multituberculaires. Autrement dit, ils semblent être une caractéristique primitive qui
pourrait être présente chez les tout premiers placentaires ;
• une ouverture pelvienne (du bassin) étroite qui indique que les nouveau-nés devaient être
très petits et que la gestation était courte, comme chez les marsupiaux actuels. Cela suggère que le
placenta est une évolution ultérieure ;
• cinq incisives de chaque côté de la mâchoire supérieure. Ce nombre est typique des
métathériens alors que le nombre maximal (voir denture) chez les placentaires modernes est de 3,
sauf pour les homodontes comme le tatou. Mais pour Eomaia, le ratio du nombre de molaires sur
celui prémolaires (il a plus de prémolaires que de molaires) est typique d’un placentaire et anormal
pour un marsupial.
Eomaia a aussi une rainure Meckelienne Meckelian groove ⇔ rainure Meckelienne (voirJohann Friedrich
Meckel von Helmsbach), qui est une relique de la mâchoire des pré-mammifères (os articulaire et os carré),
alors que la mâchoire et l’oreille interne de Eomaia sont typiquement mammifères.
• ces caractéristiques intermédiaires sont compatibles avec la phylogénie moléculaire qui
estime que les placentaires se sont diversifiés il y a 110 millions d'années, soit 15 millions d'années
après le fossile "Eomaia".
"Eomaia" a aussi beaucoup de caractéristiques qui suggèrent fortement qu’il était un grimpeur
notamment :
• plusieurs caractéristiques des pieds et des orteils ;
• les points d’attaches des muscles, qui sont utilisés pour grimper, sont bien
développés ;
• et une queue qui est deux fois plus grande que le reste de la colonne vertébrale.
La caractéristique la plus connue des placentaires est leur méthode de reproduction :
• l’embryon est attaché à l’utérus via un grand placenta par lequel la mère fournit
l'alimentation, oxygène et évacue les déchets ;
• la gestation est relativement longue et les nouveau-nés sont assez bien développés à la
naissance, notamment chez les herbivores vivant sur des plaines où le jeune peut marcher et même
courir dans l’heure qui suit la naissance.
Une théorie évoque la possibilité de l’intervention de rétrovirus dans l'évolution vers la reproduction
placentaire[35]. Ces rétrovirus auraient fait l'interface entre le placenta et l'utérus dans un syncytium, c'est-à-
dire une couche mince de cellules avec une membrane externe partagée. Ce syncytium aurait permis le
passage de l'oxygène, des substances nutritives et des déchets, mais empêché le passage de sang et
d'autres cellules qui provoqueraient l’attaque du fœtus par le système immunitaire de la mère. Cela aurait
aussi réduit l'agressivité du système immunitaire de la mère (ce qui est bon pour le fœtus, mais rend la mère
plus vulnérable aux infections).
Mais du point de vue purement paléontologique, les euthériens sont principalement reconnus à leurs
dents[36].

Expansion des niches écologiques au Mésozoïque [modifier]

Il y a toujours une certaine vérité dans le stéréotype du "petit insectivore nocturne". Récemment on a
trouvé, principalement en Chine, certains mammaliens et de vrais mammifères qui étaient plus grands et
plus variés dans leurs styles de vie.
Par exemple :
• Castorocauda, qui a vécu au milieu du Jurassique il y a 164 Ma, long d’environ 43 cm, pour
800 grammes, avec des membres adaptés pour nager et pour creuser, des dents adaptées au
poisson[37].
• le Multituberculaire, qui survécut pendant plus de 120 Ma (du milieu du Jurassique, il y a 160
Ma, jusqu’au début de l'Oligocène, il y a de 35 Ma), est souvent appelé "le rongeur du Mésozoïque",
parce qu'il avait des incisives à croissance continue comme ceux des rongeurs modernes[15].
• Fruitafossor, de la dernière période Jurassique il y a environ 150 Ma, était de la taille d'un
tamia et ses dents, ses pattes avant et arrières suggèrent qu'il forait les nids d'insectes sociaux pour
les manger (probablement des termites, puisque les fourmis n'étaient pas encore apparues[38]).
 • Volaticotherium, à la limite du Jurassique et du Crétacé, il y a 140-120 Ma, est le tout premier
mammifère planant connu. Il avait une membrane entre ses membres, comme celle d'un écureuil
volant moderne. Cela évoque aussi qu’il était actif principalement le jour[39].
• Repenomamus, du début du Crétacé, il y a 128-139 Ma, était un prédateur trapu, semblable
au blaireau, qui chassait parfois de jeunes dinosaures. Deux espèces ont été reconnues, l’une de
plus d’un mètre de long et pesant environ 12–14 kg, l'autre de moins de 0,5 m de long et pesant de 4
à 6 kg[40], [41].

L’évolution des principaux groupes de mammifères vivants [modifier]

Il y a actuellement de vigoureux débats entre les paléontologues traditionnels ("les chasseurs de
fossile") et les phylogénéticiens moléculaires sur quand et comment les vrais mammifères se sont diversifiés,
et particulièrement les placentaires.
Généralement les paléontologues traditionnels datent l'apparition d'un groupe particulier par le plus
ancien fossile connu dont les caractéristiques le font probablement être un membre de ce groupe. Tandis
que phylogénéticiens affirme que chaque origine a divergé plus tôt (d'habitude dans le Crétacé) et que les
premiers membres de chaque groupe étaient anatomiquement très semblables aux premiers membres des
autres groupes divergeant et ne différaient que dans leurs gènes.
Ces débats s'étendent à la définition des liens entre les principaux groupes de placentaires - la
controverse sur Afrotheria est un bon exemple.
Arbre d’évolution des placentaires construit à partir des fossiles [modifier]
Voici une version très simplifiée d’un arbre phylogénétique basé sur les fossiles[42]. Les
paléontologues sont d’accords sur l’ensemble, en particulier sur les groupes principaux, mais il y a des
controverses sur les détails.
Par brièveté et simplicité, le diagramme omet quelques groupes éteints pour se concentrer sur la
généalogie des groupes modernes bien connus de placentaires
Le diagramme montre aussi :
• L'âge des plus vieux fossiles connus pour la plupart des groupes puisque l’un des grands
sujets de débat entre paléontologues traditionnels et phylogénéticiens moléculaire est à partir de
quand on considère des divers groupes comme devenus distincts.
• Les membres modernes typiques de la plupart des groupes.
--Euthériens
|
+--Xénarthres (Paléocène)
| (tatous, fourmiliers, paresseux)
|
`--+--Pholidota (début du Éocène)
| (pangolins)
|
`--Épithériens (Crétacé tardif)
|
|--(plusieurs groupes éteints)
 |
`--+--Insectivores (Crétacé tardif)
| (hérissons, musaraignes, taupes, tangues)
|
`--+--+--Anagalida
| | |
| | +--Zalambdalestidae (éteint) (Crétacé tardif)
| | |
| | `--+--Macroscelidea (Éocène tardif)
| | | (musaraignes éléphant)
| | |
| | `--+--Anagaloidea (éteint)
| | |
| | `--Glires (début Paléocène)
| | |
| | +--Logomorphes (Éocène)
| | | (lapins, lièvres, pikas)
| | |
| | `--Rodentia (Paléocène tardif)
| | (souris et rats, écureuils, porcs-
épics)
| |
| `--Archonta
| |
| |--+--Scandentia (mi-Éocène)
| | | (musaraignes arboricoles)
| | |
| | `--Primatomorpha
| | |
| | +--Plésiadapiformes (éteint)
| | |
| | `--Primates (début du Paléocène)
| | (tarsiers, lémuriens, singes, hommes)
| |
| `--+--Dermoptera (Éocène tardif)
| | (dermoptères)
| |
| `--Chiroptères (Paléocène tardif)
| (chauve-souris)
|
`--+--Ferae (début du Paléocène)
| (chats, chiens, ours, phoques)
|
`--Ungulatomorpha (Crétacé tardif)
|
+--Eparctocyona (Crétacé tardif)
| |
| +--(groupes éteints)
 | |
| `--+--Arctostylopida (éteint) (Paléocène
tardif)
| |
| `--+--Mesonychia (éteint) (mi-Paléocène)
| | (prédateurs/charognards, mais pas
directement liés aux carnivores modernes)
| |
| `--Cetartiodactyla
| |
| +--Cétacés (début de l’Éocène)
| | (baleines, dauphins, marsouins)
| |
| `--Artiodactyla (début de l’Éocène)
| (ongulés artiodactyles : porcs,
hippopotames, chameaux,
|
girafes, vaches, cerfs)
|
`--Altungulata
|
+--Hilalia (éteint)
|
 `--+--+--ongulés périssodactyles (Paléocène
tardif)
| | (Perissodactyla : chevaux,
rhinocéros, tapirs)
| |
| `--Tubulidentata (début du Miocène)
| (Oryctérope du Cap)
|
`--Paenungulata ("non-ongulés")
|
+--Hyracoidea (début de l’Éocène)
| (Damans)
|
`--+--Siréniens (début de l’Éocène)
| (lamantins, dugongs)
|
`--Proboscidea (début de l’Éocène)
(éléphants)

Cet arbre généalogique contient quelques surprises et énigmes.

Par exemple :
 • Les plus proches parents vivants des cétacés (baleines, dauphins, marsouins) sont les
artiodactyles, des animaux à sabots (cochons, chameaux, ruminants) qui sont presque tous des
végétariens exclusifs ;
• les chauves-souris sont des parents assez proches des primates ;
• les plus proches parents vivants des éléphants sont les siréniens aquatiques (dugongs et
lamantins), tandis que leurs parents suivants sont les damans qui ressemblent plutôt à de gros
cobayes ;
• il y a peu de correspondance entre la structure de la famille (qui descend de qui) et les dates
de premiers fossiles de chaque groupe. Par exemple les fossiles les plus anciens de périssodactyles
(dont les membres vivants sont les chevaux, les rhinocéros et les tapirs) datent de la fin du
Paléocène, mais les premiers fossiles du "groupe cousin" des Tubulidentés datent du premier
Miocène, presque 50 millions des années plus tard.
Les paléontologues sont assez confiants sur la fiabilité des relations entre les groupes issus de
l’analyse cladistique.
Ils pensent que les « trous » constatés comme les fossiles des ancêtres de l'Oryctérope du Cap
moderne n'ont tout simplement pas encore été trouvés.

Arbre d’évolution des placentaires d’après la phylogénétique moléculaire

[modifier]
La phylogénie moléculaire utilise les propriétés des gènes des organismes vivants pour élaborer des
arbres d’évolution de la même manière que les paléontologues le font avec les fossiles. Si 2 fossiles ou 2
gènes sont plus semblables entre eux que par rapport à un 3e, alors les 2 organismes sont plus proches
parents que le troisième. Les phylogénéticiens moléculaires ont proposé un arbre d’évolution qui est différent
de celui des paléontologues.
Comme partout, les détails sont sujets à discussions et controverses, mais l’arbre présenté ici est
celui qui fait le plus consensus[43], [44].
Remarquons que le schéma ci-dessous omet les groupes disparus car on ne peut extraire l’ADN à
partir des fossiles.
--Euthériens
|
+--Atlantogenata ("Né autour de l’océan Atlantique")
| |
| +--Xénarthres (tatous, fourmiliers, paresseux)
| |
| `--Afrothériens
| |
| +--Afroinsectiphilia (Taupes dorés, tangue, potamogale)
| |
| +--Pseudungulata ("faux ongulés")
| | |
| | +--Macroscelidea (musaraignes éléphants)
| | |
| | `--Tubulidentata (Oryctérope du Cap)
| |
| `--Paenungulata ("Pas tout à fait ongulés")
 | |
| +--Hyracoidea (damans)
| |
| +--Proboscidea (éléphants)
| |
| `--Siréniens (lamantins, dugongs)
|
`--Boreoeutheria ("les Euthériens (placentaires) du Nord")
|
+--Laurasiatheria
| |
| +--Erinaceomorpha (hérissons)
| |
| +--Soricomorpha (taupes, musaraignes, solénodons)
| |
| +--Cetartiodactyla (cétacés et ongulés à nombre de doigts
pair)
| |
| `--Pegasoferae
| |
| +--Pholidota (pangolins)
| |
| +--Chiroptères (chauve-souris)
| |
 | +--Carnivora (chats, chiens, ours, phoques)
| |
| `--Perissodactyla (chevaux, rhinocéros, tapirs)
|
`--Euarchontoglires
|
+--Glires
| |
| +--Lagomorpha (lapins, lièvres, pikas)
| |
| `--Rodentia (souris rats, écureuils, porcs-épics)
|
`--Euarchonta
|
|--Scandentia (toupailles et ptilocerques)
|
|--Dermoptera (écureuils volants)
|
`--Primates (tarsiers, lémuriens, singes, grands singes,
humains)

Les plus significatives des nombreuses différences entre cet arbre d’évolution et celui des
paléontologues sont :
 • La division au niveau supérieur est entre Atlantogenata et Boreoeutheria, au lieu d'entre
Xénarthres et le reste. Mais certains phylogénéticiens moléculaire ont proposé une scission à 3 voies
de niveau supérieur entre Xénarthres, Afrothériens et Boreoeutheria ;
• Afrotheria contient plusieurs groupes qui sont seulement de lointains cousins selon la
version des paléontologues : Afroinsectiphilia ("insectivores africains"), Tubulidentata (Oryctérope du
Cap, que les paléontologues considèrent comme proche des ongulés au nombre de doigts pair, qu’à
d'autres membres d'Afrotheria), Macroscélididé (des rats à trompe, d'habitude considérés comme
proches des lapins et des rongeurs). Les seuls membres d'Afrotheria que les paléontologues
considéreraient comme étroitement associés sont les Hyracoidea (damans), Proboscidiens
(éléphants) et siréniens (lamantins, dugongs) ;
• les insectivores sont divisés en 3 groupes : le premier fait partie d'Afrotheria et les deux
autres sont des sous-groupes distincts dans Boreoeutheria ;
• les chauves-souris sont plus proches des carnivores et des ongulés au nombre de doigts
impairs qu'aux primates et aux dermoptères ;
• les Périssodactyles (des ongulés au nombre de doigts impair) sont plus proches des
carnivores et des chauves-souris que des Artiodactyles (ongulés au nombre de doigts pair) ;
La réunion des groupes sous Afrotheria (« les Thériens de l’Afrique ») a un peu de justification
géologique. Tous les membres survivants d'Afrotheria vivent principalement en Amérique du Sud ou en
Afrique.
Comme la rupture de la Pangée a éloigné l'Afrique et l'Amérique du Sud des autres continents il y a moins de
150 millions d’années et que l’un et l’autre se sont séparés il y a entre 80 et 100 millions d’années[45], [46].
Le tout premier mammifère Euthérien connu est Eomaia, apparu il y a environ 125 millions d’années. Ainsi il
ne serait pas surprenant que les premiers migrants euthériens en Afrique et en Amérique du Sud se soient
trouvés isolés et aient investi (voir Radiation évolutive) toutes les niches écologiques présentes.
Néanmoins, ces propositions sont controversées. Les Paléontologues insistent naturellement que les
preuves par les fossiles sont prioritaires par rapport aux déductions à partir d’ADN. Plus surprenant, ce
nouvel arbre phylogénétique a été critiqué aussi par d’autres phylogénéticiens moléculaires, et parfois
durement[47], car :
• le taux de mutation de l’ADN mitochondrial chez les mammifères varie d’une région à l’autre.
Dans certaine, l’ADN ne change quasiment pas et dans d’autres il change très rapidement. On
constate même de grandes variations entre les individus d’une même espèce[48], [49].
L’ADN mitochondrial des mammifères mute si vite qu’il cause des problèmes de «saturation », où le
bruit de fond parasite les informations qui pourraient être utiles. Par exemple, si un locus (emplacement
précis et invariable sur un chromosome) mute aléatoirement tous les quelques millions d’années, il aura
changé plusieurs fois depuis le moment où les principaux groupes de mammifères ont divergé (de -60 à -70
millions d’années) masquant ainsi l'information recherchée[50].

Chronologie de l’évolution des Placentaires [modifier]

Des récentes études phylogénétiques évoquent l’hypothèse que la majorité des ordres placentaires
ont divergé il y a entre 100 et 85 millions d’années, mais que les familles modernes apparaissent à la fin de
l’Éocène et au début du Miocène[51].
 Les paléontologues objectent qu’aucun fossile de placentaires n’a été trouvé d’avant la fin du
Crétacé, et que même le fossile le plus récent Maelestes gobiensis, datant d’il y a 75 millions d’années, est
un euthérien et pas un vrai mammifère[52].
Les fossiles des premiers membres de la plupart des groupes modernes datent du Paléocène,
quelques-uns un peu plus tard et très peu du Crétacé, juste avant l'extinction des dinosaures.

Évolution des caractéristiques des mammifères [modifier]

La mâchoire et l’oreille interne moyenne [modifier]

Hadrocodium dont les fossiles datent du début du Jurassique, fournissent la première preuve claire
d’une articulation de mâchoire et d’une oreille moyenne entièrement mammifères.
Curieusement il est d'habitude classé comme membre des mammaliens plutôt que comme un vrai
mammifère.

La production de lait (lactation) [modifier]

Certains font l’hypothèse que la première fonction de la lactation était de couver les œufs en les
gardant chauds et humides.
Cette argumentation a pour référence les monotrèmes (mammifères pondeurs)[53], [54], [55]:
Poils et fourrure [modifier]
La plus ancienne preuve tangible de poils a été trouvé sur le fossile de Castorocauda daté de 164
Ma, c'est-à-dire du milieu du Jurassique.
Peu avant 1955, quelques chercheurs ont interprété la présence de foramina dans le maxillaire et les
prémaxillaires des cynodontes comme étant des canaux qui permettaient le passage des vaisseaux
sanguins et des nerfs pour les vibrisses. Ce qui serait alors selon eux la preuve de poils ou de fourrure[56],
[57]. Mais un foramen ne prouve pas nécessairement qu'un animal avait des moustaches. Par exemple les
Tupinambis, un genre de lézard ont des foramina maxilliaire presque identique à ceux trouvés chez le
cynodonte Thrinaxodon[58], [59].

Les membres droits [modifier]

erect ⇔ droit
L’évolution vers des membres droits est incomplète chez les mammifères. Les monotrèmes vivants et
fossiles ont des pattes latérales sprawling ⇔ latéral. En fait, des chercheurs pensent que la position
parasagitale (non latérale) est un synapomorphisme (une caractéristique particulière) des Boreosphenida,
un groupe qui contient les Thériens et donc inclut le plus ancien ancêtre commun aux marsupiaux et
placentaires modernes[60] Sinodelphys (le plus ancien marsupial connu) et Eomaia (le plus ancien
euthériens connu) vivaient il y a environ 125 millions d’années, donc la divergence a dû se faire avant eux.

Le sang chaud [modifier]

Le sang chaud est une notion complexe et plutôt ambigüe, parce qu’elle inclut l’une ou toutes ces
composantes :
 • ’’Endothermie’’, c'est-à-dire la capacité de générer en interne plutôt que par un
comportement comme l’activité musculaire ou se chauffer au Soleil ;
• ’’Homéothermie’’, c'est-à-dire la capacité de maintenir sa température corporelle à peu près
constante ;
• ’’Tachymétabolisme’’ c'est-à-dire la capacité de maintenir un métabolisme élevé, en
particulier au repos. Ce qui nécessite de maintenir une température corporelle stable et élevée car la
plupart des enzymes ont une plage de température de travail optimale, et leur efficacité faiblit
rapidement en dehors sa température de prédilection. Ainsi les réactions biochimiques sont 2 fois
plus lentes lorsque la température corporelle chute de 10°C.
Comme nous ne pouvons pas connaitre les mécanismes internes des espèces disparues, la plupart
des discussions se concentrent sur homéothermie et le tachymétabolisme.
Les monotrèmes actuels ont une température corporelle plus basse et un niveau de métabolisme
plus variable que les marsupiaux et les placentaires[61]. La principale question est de savoir comment le
métabolisme de type monotrème a évolué vers celui des mammifères. Les éléments trouvés jusqu’ici font
penser que le cynodontes du Trias avait un métabolisme assez élevé, mais sans preuve évidente.

Cornet nasal [modifier]

Les mammifères modernes ont des cornets nasaux respiratoires, les structures complexes d'os
minces dans la cavité nasale. Ces os sont alignés avec la muqueuse qui réchauffe et humidifie l'air inhalé en
extrayant la chaleur et l'humidité de l'air exhalé. Un animal avec des cornets respiratoires peut maintenir un
haut taux de respiration sans danger de dessécher ses poumons et peut donc avoir un métabolisme élevé.
 Malheureusement ces os nasaux sont très délicats et n'ont pas donc encore été trouvés dans des
fossiles. Mais des arêtes rudimentaires comme ceux que supportent les cornets respiratoires ont été
trouvées chez des thérapsides du Trias comme "Thrinaxodon" et "Diademodon", qui suggèrent qu'ils
puissent avoir eu des métabolismes assez élevés[62], [63], [64].

Le second palais osseux [modifier]

Les mammifères ont un second palais osseux qui séparent les voies respiratoires de la bouche, leur
permettant de manger et de respirer ne même temps. Le second palais osseux a été trouvé chez les
cynodontes les plus évolués, ce qui est considéré comme une preuve d’un métabolisme élevé.[64], [65], [66].
Mais certains vertébrés à sang froid ont un second palais (les crocodiles et certains lézards), alors que les
oiseaux, qui sont “à sang chaud”, n’en ont pas[67].

Le Diaphragme [modifier]
Le muscle diaphragme aide les mammifères à respirer, particulièrement lors d’une activité physique
intense. Pour que le diaphragme puisse fonctionner, les côtes ne doivent pas limiter l'abdomen, pour que
l'expansion de la poitrine puisse être compensée de par la réduction du volume du ventre et inversement.
Les cynodontes évolués avaient une cage thoracique très semblable à celle des mammifères, avec un
nombre réduit de côtes lombaires. Cela suggère que ces animaux avaient un diaphragme et qu’ils étaient
capables d’une activité soutenue, et donc d’un haut métabolisme[64], [65]. D’un autre côté, ces cages
thoraciques de type mammifère pourraient aussi avoir évolué pour améliorer l’agilité[68].
Mais les mouvements des thérapsides même évolués étaient maladroits, avec les pattes arrières qui
fournissaient la poussée et les pattes avant qui servaient seulement à la direction. Autrement dit, les
thérapsides les plus évolués étaient moins agiles que les mammifères actuels ou les premiers
dinosaures[69]. Donc l’idée que la principale fonction de la cage thoracique de type mammifère soit
d’améliorer l’agilité est douteuse.

La posture des membres [modifier]

Les thérapsides ont des pattes latérales ou semi-latérales[70], [65]. Les contraintes mécaniques en
résultant font qu’ils devaient être limités pour bouger et respirer en même temps, mais pas autant que pour
ceux, comme les lézards, qui ont des membres complètement latéraux[71].
Mais les cynodontes (thérapsides évolués) avaient des plaques thoraciques qui rendaient la cage
thoracique plus rigide et donc réduisait les torsions du tronc quand l’animal bougeait et facilitait la
respiration[72]. Au vu de ces éléments, on peut dire que les thérapsides évolués étaient significativement
moins actif que les mammifères actuels de taille similaire, et qu’ils avaient donc un métabolisme moins
élevé.

Isolation (poils et fourrure) [modifier]

L’isolation est la façon la plus simple de maintenir à peu près constante une température corporelle.
Donc la présence de poils et de fourrure semble un bon élément pour conclure à l'homéothermie, mais n’est
pas un élément déterminant d’un métabolisme élevé[73], [74]. Nous avons déjà vu que la première preuve
claire de présence poils ou de fourrure est le fossile de "Castorocauda" d'il y a 164 millions d’années, au
milieu du Jurassique ; les arguments avançant que les thérapsides étaient velus, sont peu convaincants.
Notes et références [modifier]
1. ↑ Le crapaud du Surinam, est un des rares amphibiens à avoir développé d’autres manières
de contourner "l'impossibilité" (la difficulté) de pondre hors de l'eau.
2. ↑ (en)R. A. Kissel et R. R. Reisz. «Synapsid fauna of the Upper Pennsylvanian Rock Lake
Shale near Garnett, Kansas and the diversity pattern of early amniotes», dans G. arratia, M. V. H.
Wilson & R. Cloutier (dir.), Recent advances in the origin and early radiation of vertebrates , Verlag
Dr. Friedrich Pfeil, Munich, 2004, p. 409-428.
3. ↑ (en)Synapsida overview [archive] sur Palaeos
4. ↑ a et b (en)Therapsida [archive] sur Palaeos
5. ↑ (en) D. M. Kermack & K. A. Kermack, The evolution of mammalian characters, Croom
Helm (ISBN 079915349)
6. ↑ A.F. Bennett & J.A. Ruben, The ecology and biology of mammal-like reptiles, Smithsonian
Institution Press, Washington, 1986, 207-218 p. (ISBN 978-0874745245)
7. ↑ a et b (en)Cynodontia Overview [archive] sur Palaeos
8. ↑ (en)The Rhaetian Age of the Late Triassic Epoch: 204 to 200 million years ago [archive]
sur Palaeos
9. ↑ (en)The Triassic Period [archive] sur Palaeos
10.↑ voir lien : Radiation évolutive
11.↑ Évaluation du budget énergétique du cerveau. (en)M.E. Raichle et D.A. Gusnard,
« Appraising the brain's energy budget », dans PNAS, vol. 99, no 16, August 6, 2002, p. 10237-
10239 [ texte intégral [archive] ]
 12.↑ (en)Brain power [archive], 2006, New Scientist
13.↑ Recherche visionnaire : les scientifiques fouillent dans l'évolution de la perception
chromatique des primates. (en)J Travis, « Visionary research: scientists delve into the evolution of
color vision in primates », dans Science News, vol. 164, no 15, October 2003 [ texte
intégral [archive] ]
14.↑ R.L. Cifelli, « Early mammalian radiations », dans Journal of Paleontology, November
2001 [ texte intégral [archive] ]
15.↑ a, b, c et d (en)Mammaliformes [archive] sur Palaeos
16.↑ R.L. Cifelli, « Early mammalian radiations », dans Journal of Paleontology, November
2001 [ texte intégral [archive] ]
17.↑ (en)Morganucodontids & Docodonts [archive] sur Palaeos
18.↑ (en)Jurassic "Beaver" Found; Rewrites History of Mammals [archive]
19.↑ (en)Symmetrodonta [archive] sur Palaeos
20.↑ (en)Mammalia [archive] sur Palaeos
21.↑ (en)Mammalia [archive] sur Palaeos
22.↑ (en)Appendicular Skeleton [archive]
23.↑ (en)Mammalia: Spalacotheroidea & Cladotheria [archive] sur Palaeos
24.↑ (en)Appendicular Skeleton [archive]
25.↑ (en)Metatheria [archive] sur Palaeos
 26.↑ (en) F.S. Szalay & B.A. Trofimov, « The Mongolian Late Cretaceous Asiatherium, and
the early phylogeny and paleobiogeography of Metatheria », dans Journal of Vertebrate
Paleontology, vol. 16, no 3, 1996, p. 474-509 [ texte intégral [archive] ]
27.↑ (en)Oldest Marsupial Fossil Found in China [archive], 15 décembre 2003, National
Geographic News
28.↑ (en)Didelphimorphia [archive] sur Palaeos
29.↑ (en)Family Peramelidae (bandicoots and echymiperas) [archive]
30.↑ Species is as species does... Part II [archive]
31.↑ Les marsupiaux. (en)Marsupials [archive]
32.↑ L'os épipupis chez les mammifères euthériens au Crétacé tardif en Mongolie. (en)« »
33.↑ Ji et al, les premiers mammifères euthériens connus. The earliest known eutherian
mammal, Nature, 2002, vol. 416, p. 816-822
34.↑ (découverte du plus ancien placenta mammalien.) (en)Eomaia
scansoria: [archive]discovery of oldest known placental mammal.
35.↑ (Pourquoi nous ne pondons pas d’oeufs ?) (en)« »
36.↑ (en)Eutheria [archive] sur Palaeos
37.↑ (en)Jurassic "Beaver" Found; Rewrites History of Mammals [archive], National
Geographic.
38.↑ (en) « A Late Jurassic Digging Mammal and Early Mammal Diversification », dans
Science, vol. 308, 2005, p. 103-107 [ résumé [archive] ]
 39.↑ (en) Meng, J., Hu, Y., Wang, Y., Wang, X., Li, C., « A Mesozoic gliding mammal from
northeastern China », dans Nature, vol. 444, no 7121, Dec 2006, p. 889-893 [ texte
intégral [archive] ]
40.↑ (en) Li, J., Wang, Y., Wang, Y., Li, C., « A new family of primitive mammal from the
Mesozoic of western Liaoning, China », dans Chinese Science Bulletin, vol. 46, no 9, 2000, p. 782-
785 [ résumé [archive] ]
41.↑ Hu, Y., Meng, J., Wang, Y., Li, C., « Large Mesozoic mammals fed on young dinosaurs »,
dans Nature, vol. 433, 2005, p. 149-152 [ texte intégral [archive] ]
42.↑ À partir de Cladogram of Mammalia - Palaeos [archive]
43.↑ Murphy, W.J., Eizirik, E., Springer, M.S et al, « Resolution of the Early Placental Mammal
Radiation Using Bayesian Phylogenetics », dans Science, vol. 294, no 5550, 14 December 2001,
p. 2348-2351
44.↑ Kriegs, J.O., Churakov, G., Kiefmann, M., et al, « Retroposed Elements as Archives for the
Evolutionary History of Placental Mammals », dans PLoS Biol, vol. 4, no 4, 2006, p. e91 (pdf
version [archive])
45.↑ Cartes mondiales successives, depuis la Pangée ( ≅ -225 Ma). (en)Historical
perspective (the Dynamic Earth, USGS) [archive]
46.↑ Carte du Crétacé, à -94 Ma. (en)Cretaceous map [archive]
47.↑ Les insectivores. (en)Insectivora Overview - Palaeos [archive]
48.↑ M.S. Springer et E. Douzery, « Secondary Structure and patterns of evolution among
mammalian mitochondrial 12S rRNA molecules », dans J. Mol. Evol., vol. 43, 1996, p. 357-373
49.↑ M.S. Springer, L.J. Hollar et A. Burk, « Compensatory substitutions and the evolution of the
mitochondrial 12S rRNA gene in mammals », dans Mol. Biol. Evol., vol. 12, 1995, p. 1138-1150
 50.↑ (en) W-H Li, Molecular Evolution, Sinauer Associates, 1997
51.↑ L'apparition tardive des mammifères actuels. (en)O.R.P. Bininda-Emonds, M. Cardillo,
K.E. Jones et 'et al', « The delayed rise of present-day mammals », dans Nature, no 446, 2007,
p. 507-511 [ texte intégral [archive] ]
52.↑ la disparition des dinosaures stimula l'apparition des mammifères modernes.
(en)Dinosaur Extinction Spurred Rise of Modern Mammals [archive]
53.↑ O.T. Oftedal, « The mammary gland and its origin during synapsid evolution », dans
Journal of Mammary Gland Biology and Neoplasia, vol. 7, no 3, 2002, p. 225-252
54.↑ O.T. Oftedal, « The origin of lactation as a water source for parchment-shelled eggs »,
dans Journal of Mammary Gland Biology and Neoplasia, vol. 7, no 3, 2002, p. 253-266
55.↑ L'initiale "lactaion" sur les oeufs. (en)Lactating on Eggs. [archive]
56.↑ A.S. Brink, « A study on the skeleton of Diademodon », dans Palaeontologia Africana,
vol. 3, 1955, p. 3-39
57.↑ T.S. Kemp, Mammal-like reptiles and the origin of mammals, Academic Press, London,
1982, 363 p.
58.↑ Bennett, A. F. and Ruben, J. A. (1986) "The metabolic and thermoregulatory status of
therapsids"; pp. 207-218 in N. Hotton III, P. D. MacLean, J. J. Roth and E. C. Roth (eds), "The
ecology and biology of mammal-like reptiles", Smithsonian Institution Press, Washington
59.↑ « Cranial anatomy of the cynodont reptile Thrinaxodon liorhinus », dans Bulletin of the
Museum of Comparative Zoology, no 1253, 1961, p. 165-180
60.↑ La position des membres chez les premiers mammifères : droits ou parasagittals ?
(en)Z. Kielan−Jaworowska et J.H.. Hurum, « Limb posture in early mammals: Sprawling or
parasagittal ? », dans Acta Palaeontologica Polonica, vol. 51, no 3, 2006, p. 10237-10239 [ texte
intégral [archive] ]
61.↑ (en) G.S. Paul, Predatory Dinosaurs of the World, Simon and Schuster, New York,
1988, p.
62.↑ W.H. Hillenius, « The evolution of nasal turbinates and mammalian endothermy », dans
Paleobiology, vol. 18, no 1, 1992, p. 17-29
63.↑ J. Ruben, « The evolution of endothermy in mammals and birds: from physiology to
fossils », dans Annual Review of Physiology, vol. 57, 1995, p. 69-95
64.↑ a, b et c A.S. Brink, « A study on the skeleton of Diademodon », dans Palaeontologia
Africana, vol. 3, 1955, p. 3-39
65.↑ a, b et c (en) T.S. Kemp, Mammal-like reptiles and the origin of mammals, Academic
Press, London, 1982, p.
66.↑ B.K. McNab, « The evolution of endothermy in the phylogeny of mammals », dans
American Naturalist, vol. 112, 1978, p. 1-21
67.↑ Bennett, A. F. and Ruben, J. A. (1986) "The metabolic and thermoregulatory status of
therapsids"; pp. 207-218 in N. Hotton III, P. D. MacLean, J. J. Roth and E. C. Roth (eds), "The
ecology and biology of mammal-like reptiles", Smithsonian Institution Press, Washington.
68.↑ Bennett, A. F. and Ruben, J. A. (1986) "The metabolic and thermoregulatory status of
therapsids"; pp. 207-218 in N. Hotton III, P. D. MacLean, J. J. Roth and E. C. Roth (eds), "The
ecology and biology of mammal-like reptiles", Smithsonian Institution Press, Washington.
69.↑ (en) R. Ccowen, History of Life, Blackwell Science, Oxford, 2000, p.
70.↑ F.A., Jr Jenkins, « The postcranial skeleton of African cynodonts », dans Bulletin of the
Peabody Museum of Natural History, no 36, 1971, p. 1-216
 71.↑ (en) F.H Pough, J.B. Heiser et W.N. McFarland, Vertebrate Life, Prentice-Hall, New
Jersey, 1996, p.
72.↑ C.A. Sidor et J.A. Hopson, « Ghost lineages and "mammalness": assessing the temporal
pattern of character acquisition in the Synapsida », dans Paleobiology, no 24, 1998, p. 254-273
73.↑ (en) K. Schmidt-Nielsen, Animal physiology: Adaptation and environment, Cambridge
University Press, Cambridge, 1975, p.
74.↑ (en) P.C. Withers, Comparative Animal Physiology, Saunders College, Fort Worth,
1992, p.

Voir aussi [modifier]

Articles connexes [modifier]
• Mammalia (classification phylogénétique)
• Histoire évolutive des cétacés
• Histoire évolutive des hippopotamidés
• Histoire évolutive des siréniens
• Histoire évolutive des équidés
• Histoire évolutive des primates
• Évolution humaine
Articles similaires [modifier]
• Histoire évolutive des oiseaux
• Histoire évolutive des poissons cartilagineux
• Histoire évolutive des gastéropodes
• Domestication

Sources [modifier]
• Cet article est initialement issu d’une traduction de l’article de Wikipédia en anglais intitulé
« Evolution of mammals », dont la version (actuelle) est directement accessible.

Liens externes [modifier]

• (en)The Cynodontia Site couvrant de nombreux aspects de l'évolution des Cynodontes en
mamifères, avec de nombreuses références.
• Emmanuel Gheerbrant, « Le Paléogène et la radiation des mammifères », CNRS.
• Didier Grandperrin, « L’origine et la radiation des mammifères », académie de Orléans-
Tours.
• Frédéric Delsuc, Jean-François Mauffrey, Emmanuel Douzery, « Une nouvelle classification
des mammifères », dans Pour La Science, vol. 303., Janvier 2003 [ texte intégral ]
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Archaeopteryx
From Wikipedia, the free encyclopedia

Jump to: navigation, search

Archaeopteryx
Fossil range: Late Jurassic, 150.8–148.5 Ma
↓
g
r
e
Є
 The Berlin Archaeopteryx

Scientific classification

Kingdom: Animalia

Phylum: Chordata

Class: Aves

Order: Archaeopterygiformes

Family: Archaeopterygidae

Archaeopteryx
Genus:
Meyer, 1861

Species

A. lithographica Meyer, 1861 (type)

Synonyms

• Griphosaurus Wagner, 1861

• Griphornis Woodward, 1862
• Archaeornis Petronievics, 1917
 • Jurapteryx? Howgate, 1985
• Wellnhoferia? Elżanowski, 2001
For species, see below
Archaeopteryx (pronounced /ˌɑrkiːˈɒptərɨks/ AR-kee-OP-ter-iks), sometimes referred to by its
German name Urvogel ("original bird" or "first bird"), is the earliest and most primitive bird known. The name
derives from the Ancient Greek ἀρχαῖος (archaīos) meaning "ancient", and πτέρυξ (ptéryx), meaning "wing".
Archaeopteryx lived during the Late Jurassic Period around 150–148 million years ago, in what is now
southern Germany during a time when Europe was an archipelago of islands in a shallow warm tropical sea,
much closer to the equator than it is now.
Similar in size and shape to a European Magpie, Archaeopteryx could grow to about 0.5 metres
(1.6 ft) in length. Despite its small size, broad wings, and inferred ability to fly or glide, Archaeopteryx has
more in common with small theropod dinosaurs than it does with modern birds. In particular, it shares the
following features with the deinonychosaurs (dromaeosaurs and troodontids): jaws with sharp teeth, three
fingers with claws, a long bony tail, hyperextensible second toes ("killing claw"), feathers (which also suggest
homeothermy), and various skeletal features.[1][2] The features above make Archaeopteryx a clear
candidate for a transitional fossil between dinosaurs and birds.[3][4] Thus, Archaeopteryx plays an important
role not only in the study of the origin of birds but in the study of dinosaurs. The first complete specimen of
Archaeopteryx was announced in 1861, only two years after Charles Darwin published On the Origin of
Species, and it became a key piece of evidence in the debate over evolution. Over the years, nine more
fossils of Archaeopteryx have surfaced. Despite variation among these fossils, most experts regard all the
remains that have been discovered as belonging to a single species, though this is still debated.
 Many of these eleven fossils include impressions of feathers—among the oldest direct evidence of
feathers. Moreover, because these feathers are an advanced form (flight feathers), these fossils are
evidence that the evolution of feathers began before the Late Jurassic.[5]
Contents
[hide]
• 1 Description
• 2 Paleobiology
• 2.1 Plumage
• 2.2 Flight
• 3 Paleoecology
• 4 History of discovery
• 4.1 Taxonomy
• 4.2 Synonyms
• 5 Controversies
• 5.1 Authenticity
• 5.2 Archaeopteryx and Protoavis
• 5.3 Phylogenetic position
• 6 In popular culture
• 7 See also
• 8 References
• 9 Further reading
• 10 External links
[edit] Description
Archaeopteryx was a primitive bird that lived during the early Tithonian stage of the Jurassic Period,
around 150.8–148.5 million years ago.[6] The only specimens of Archaeopteryx that have been discovered
come from the Solnhofen limestone in Bavaria, southern Germany, which is a lagerstätte, a rare and
remarkable geological formation known for its superbly detailed fossils.[7][8]
Eight specimens compared to a human in scale.
 Archaeopteryx was roughly the size of a medium-sized modern-day bird, with broad wings that were
rounded at the ends and a long tail compared to its body length. In all, Archaeopteryx could reach up to
500 millimeters (1.6 ft) in body length. Archaeopteryx feathers, although less documented than its other
features, were very similar in structure and design to modern-day bird feathers.[7] However, despite the
presence of numerous avian features,[9] Archaeopteryx had many theropod dinosaur characteristics. Unlike
modern birds, Archaeopteryx had small teeth[7] as well as a long bony tail, features which Archaeopteryx
shared with other dinosaurs of the time.[10]
 Comparison of the forelimbs of Deinonychus (left) and Archaeopteryx (right), one of many skeletal
similarities between avians and dromaeosaurids.
Because it displays a number of features common to both birds and dinosaurs, Archaeopteryx has
often been considered a link between them—possibly the first bird in its change from a land dweller to a bird.
[7] In the 1970s, John Ostrom, following T. H. Huxley's lead in 1868, argued that birds evolved from theropod
dinosaurs and Archaeopteryx was a critical piece of evidence for this argument; it preserves a number of
avian features, such as a wishbone, flight feathers, wings and a partially reversed first toe, and a number of
dinosaur and theropod features. For instance, it has a long ascending process of the ankle bone, interdental
plates, an obturator process of the ischium, and long chevrons in the tail. In particular, Ostrom found that
Archaeopteryx was remarkably similar to the theropod family Dromaeosauridae.[11][12][13][14][15][16][17]
[18][19]
The first remains of Archaeopteryx were discovered in 1861; just two years after Charles Darwin
published On the Origin of Species. Archaeopteryx seemed to confirm Darwin's theories and has since
become a key piece of evidence for the origin of birds, the transitional fossils debate, and confirmation of
evolution. Indeed, further research on dinosaurs from the Gobi Desert and China has since provided more
evidence of a link between Archaeopteryx and the dinosaurs, such as the Chinese feathered dinosaurs.
Archaeopteryx is close to the ancestry of modern birds, and it shows most of the features one would expect
in an ancestral bird. However, it may not be the direct ancestor of living birds, and it is uncertain how much
evolutionary divergence was already present among other birds at the time.
[edit] Paleobiology
[edit] Plumage
 1880 photo of the Berlin Archaeopteryx specimen, showing leg feathers that were subsequently
removed during preparation
Specimens of Archaeopteryx were most notable for their well-developed flight feathers. They were
markedly asymmetrical and showed the structure of flight feathers in modern birds, with vanes given stability
by a barb-barbule-barbicel arrangement.[20] The tail feathers were less asymmetrical, again in line with the
situation in modern birds and also had firm vanes. The thumb, however, did not yet bear a separately
movable tuft of stiff feathers.
The body plumage of Archaeopteryx is less well documented and has only been properly researched
in the well-preserved Berlin specimen. Thus, as more than one species seems to be involved, the research
into the Berlin specimen's feathers does not necessarily hold true for the rest of the species of
Archaeopteryx. In the Berlin specimen, there are "trousers" of well-developed feathers on the legs; some of
these feathers seem to have a basic contour feather structure but are somewhat decomposed (they lack
barbicels as in ratites),[21] but in part they are firm and thus capable of supporting flight.[22]
 Anatomical illustration comparing the "frond-tail" of Archaeopteryx to the "fan-tail" of a modern bird
There was a patch of pennaceous feathers running along the back which was quite similar to the
contour feathers of the body plumage of modern birds in being symmetrical and firm, though not as stiff as
the flight-related feathers. Apart from that, the feather traces in the Berlin specimen are limited to a sort of
"proto-down" not dissimilar to that found in the dinosaur Sinosauropteryx, being decomposed and fluffy, and
possibly even appeared more like fur than like feathers in life (though not in their microscopic structure).
These occur on the remainder of the body, as far as such structures are both preserved and not obliterated
by preparation, and the lower neck.[21]
However, there is no indication of feathering on the upper neck and head. While these may
conceivably have been nude as in many closely related feathered dinosaurs for which good specimens are
available, this may still be an artifact of preservation. It appears that most Archaeopteryx specimens became
embedded in anoxic sediment after drifting some time on their back in the sea—the head and neck and the tail
are generally bent downwards, which suggests that the specimens had just started to rot when they were
embedded, with tendons and muscle relaxing so that the characteristic shape of the fossil specimens was
achieved. This would mean that the skin was already softened and loose, which is bolstered by the fact that
in some specimens the flight feathers were starting to detach at the point of embedding in the sediment. So it
is hypothesized that the pertinent specimens moved along the sea bed in shallow water for some time before
burial, the head and upper neck feathers sloughing off, while the more firmly attached tail feathers remained.
[23]

[edit] Flight
As in the wings of modern birds, the flight feathers of Archaeopteryx were somewhat asymmetrical
and the tail feathers were rather broad. This implies that the wings and tail were used for lift generation.
However, it is unclear whether Archaeopteryx was simply a glider or capable of flapping flight. The lack of a
bony breastbone suggests that Archaeopteryx was not a very strong flier, but flight muscles might have
attached to the thick, boomerang-shaped wishbone, the platelike coracoids, or perhaps to a cartilaginous
sternum. The sideways orientation of the glenoid (shoulder) joint between scapula, coracoid and humerus—
instead of the dorsally angled arrangement found in modern birds—suggests that Archaeopteryx was unable
to lift its wings above its back, a requirement for the upstroke found in modern flapping flight. Thus, it seems
likely that Archaeopteryx was indeed unable to use flapping flight as modern birds do, but it may well have
utilized a downstroke-only flap-assisted gliding technique.[24]
Archaeopteryx wings were relatively large, which would have resulted in a low stall speed and
reduced turning radius. The short and rounded shape of the wings would have increased drag, but could also
have improved Archaeopteryx' ability to fly through cluttered environments such as trees and brush (similar
wing shapes are seen in birds which fly through trees and brush, such as crows and pheasants). The
presence of "hind wings", asymmetrical flight feathers stemming from the legs similar to those seen in
dromaeosaurids such as Microraptor, would also have added to the aerial mobility of Archaeopteryx. The first
detailed study of the hind wings by Longrich in 2006 suggested that the structures formed up to 12% of the
total airfoil. This would have reduced stall speed by up to 6% and turning radius by up to 12%.[22]
The feathers of Archaeopteryx were asymmetrical. This has been interpreted as evidence for that it
was a flyer, as flightless birds tend to have symmetrical feathers. However, some scientists, including
Thomson and Speakman, have questioned this. They studied more than 70 families of living birds, and found
that some flightless types do have a range of asymmetry in their feathers, and that the feathers of
Archaeopteryx fall into this range.[25]
In 2010, Robert L. Nudds and Gareth J. Dyke in the journal Science published a paper in which they
analysed the rachises of the primary feathers of Confuciusornis and Archaeopteryx. The analysis suggested
that the rachises on these two genera were thinner and weaker than those compared to modern birds.
According to their results, the authors determined that Archaeopteryx, along with Confuciusornis, were
unable to fly.[26] This study, however, has been criticized by Currie and Chiappe. Chiappe suggest it is
difficult to measure the rachis on fossilized feathers, and Currie means that Archaeopteryx and
Confuciusornis must have been able to fly to some degree, as their fossil are preserved in what is believed to
have been in marine or lake sediments, suggesting they must have reached the water by flight to die in it.[27]
In 2004, scientists analyzing a detailed CT scan of the braincase of Archaeopteryx concluded that its
brain was significantly larger than that of most dinosaurs, indicating that it possessed the brain size
necessary for flying. The overall brain anatomy was reconstructed using the scan. The reconstruction
showed that the regions associated with vision took up nearly one-third of the brain. Other well-developed
areas involved hearing and muscle coordination.[28] The skull scan also revealed the structure of the inner
ear. The structure more closely resembles that of modern birds than the inner ear of reptiles. These
characteristics taken together suggest that Archaeopteryx had the keen sense of hearing, balance, spatial
perception and coordination needed to fly.[29]
Archaeopteryx continues to play an important part in scientific debates about the origin and evolution
of birds. Some scientists see it as a semi-arboreal climbing animal, following the idea that birds evolved from
tree-dwelling gliders (the "trees down" hypothesis for the evolution of flight proposed by O.C. Marsh). Other
scientists see Archaeopteryx as running quickly along the ground, supporting the idea that birds evolved
flight by running (the "ground up" hypothesis proposed by Samuel Wendell Williston). Still others suggest that
Archaeopteryx might have been at home both in the trees and on the ground, like modern crows, and this
latter view is what today is considered best-supported by morphological characters. Altogether, it appears
that the species was not particularly specialized for running on the ground or for perching. Considering the
current knowledge of flight-related morphology, a scenario outlined by Elżanowski in 2002, namely that
Archaeopteryx used its wings mainly to escape predators by glides punctuated with shallow downstrokes to
reach successively higher perches, and alternatively to cover longer distances by (mainly) gliding down from
cliffs or treetops, appears quite reasonable.[23]
Critics of the "ground up" hypothesis have claimed that Archaeopteryx could not lift off from the
ground (Vazquez, 1994). This is based on the fact that they don't see clear evidence that Archaeopteryx had
an avian supracoracoideus system (the muscles which push the wings upward as birds flap).

[edit] Paleoecology
The richness and diversity of the Solnhofen limestones in which all specimens of Archaeopteryx have
been found have shed light on an ancient Jurassic Bavaria strikingly different from the present day. The
latitude was similar to Florida, though the climate was likely to have been drier, as evidenced by fossils of
plants with adaptations for arid conditions and lack of terrestrial sediments characteristic of rivers. Evidence
of plants, though scarce, include cycads and conifers while animals found include a large number of insects,
small lizards, pterosaurs and Compsognathus.[30]
The excellent preservation of Archaeopteryx fossils and other terrestrial fossils found at Solnhofen
indicates that they did not travel far before becoming preserved.[31] The Archaeopteryx specimens found are
likely therefore to have lived on the low islands surrounding the Solnhofen lagoon rather than been corpses
that drifted in from farther away. Archaeopteryx skeletons are considerably less numerous in the deposits of
Solnhofen than those of pterosaurs, of which seven genera have been found.[32] The pterosaurs included
such as Rhamphorhynchus, the group which dominated the niche currently occupied by seabirds, and which
became extinct at the end of the Jurassic. The pterosaurs, which also included Pterodactylus, were common
enough that it is unlikely that the specimens found are vagrants from the larger islands 50 km (31 miles) to
the north.[33]
The islands that surrounded the Solnhofen lagoon were low lying, semi-arid and sub-tropical with a
long dry season and little rain.[34] The closest modern analogue for the Solnhofn conditions is said to be
Orca Basin in the northern Gulf of Mexico, though that is much deeper than the Solnhofn lagoons.[32] The
flora of these islands was adapted to these dry conditions and consisted mostly of low (3 m [10 ft]) shrubs.
[35] Contrary to reconstructions of Archaeopteryx climbing large trees, these seem to have been mostly
absent from the islands; few trunks have been found in the sediments and fossilized tree pollen is also
absent.
The lifestyle of Archaeopteryx is difficult to reconstruct and there are several theories regarding it.
Some researchers suggest that it was primarily adapted to life on the ground,[36] while other researchers
suggest that it was principally arboreal.[37] The absence of trees does not preclude Archaeopteryx from an
arboreal lifestyle; several species of extant bird live exclusively in low shrubs. Various aspects of the
morphology of Archaeopteryx point to either an arboreal or ground existence, the length of its legs, the
elongation in its feet; and some authorities consider it likely to have been a generalist capable of feeding in
both shrubs, open ground and even alongside the shores of the lagoon.[33] It most likely hunted small prey,
seizing it with its jaws if it was small enough or with its claws if it was larger.

[edit] History of discovery

 Timeline of Archaeopteryx discoveries. (Click to enlarge)
Over the years, ten body fossil specimens of Archaeopteryx and a feather that may belong to it have
been found. All of the fossils come from the limestone deposits, quarried for centuries, near Solnhofen,
Germany.[30][38]
 The famous, single feather
The initial discovery, a single feather, was unearthed in 1860 and described a year later by Christian
Erich Hermann von Meyer. It is currently located at the Humboldt Museum für Naturkunde in Berlin. This is
generally assigned to Archaeopteryx and was the initial holotype, but whether it actually is a feather of this
species or another, as yet undiscovered, proto-bird is unknown. There are some indications it is indeed not
from the same animal as most of the skeletons (the "typical" A. lithographica).[39]
Soon after, the first skeleton, known as the London Specimen (BMNH 37001)[40] was unearthed in
1861 near Langenaltheim, Germany and given to a local physician Karl Häberlein in return for medical
services. He then sold it to the Natural History Museum in London, where it remains.[30] Missing most of its
head and neck, it was described in 1863 by Richard Owen as Archaeopteryx macrura, who assumed it did
not belong to the same species as the feather. In the subsequent 4th edition of his On the Origin of Species,
[41] Charles Darwin described how some authors had maintained "that the whole class of birds came
suddenly into existence during the eocene period; but now we know, on the authority of Professor Owen, that
a bird certainly lived during the deposition of the upper greensand; and still more recently, that strange bird,
the Archeopteryx, with a long lizard-like tail, bearing a pair of feathers on each joint, and with its wings
furnished with two free claws, has been discovered in the oolitic slates of Solnhofen. Hardly any recent
discovery shows more forcibly than this how little we as yet know of the former inhabitants of the world."[42]
 Replica of the London Archaeopteryx.
The Greek term "pteryx" (πτερυξ) primarily means "wing", but can also designate merely "feather".
Von Meyer suggested this in his description. At first he referred to a single feather which appeared like a
modern bird's remex (wing feather), but he had heard of and been shown a rough sketch of the London
specimen, to which he referred as a "Skelet eines mit Federn bedeckten Thiers " ("skeleton of an animal
covered in feathers"). In German, this ambiguity is resolved by the term Schwinge which does not necessarily
mean a wing used for flying. Urschwinge was the favored translation of Archaeopteryx among German
scholars in the late 19th century. In English, "ancient pinion" offers a rough approximation.
Since then ten specimens have been recovered:
The Berlin Specimen (HMN 1880) was discovered in 1876 or 1877 on the Blumenberg near
Eichstätt, Germany, by Jakob Niemeyer. He exchanged this precious fossil for a cow, with Johann Dörr.
Placed on sale in 1881, with potential buyers including O.C. Marsh of Yale University's Peabody Museum, it
was bought by the Humboldt Museum für Naturkunde, where it is now displayed. The transaction was
financed by Ernst Werner von Siemens, founder of the famous company that bears his name.[30] Described
in 1884 by Wilhelm Dames, it is the most complete specimen, and the first with a complete head. Once
classified as a new species, A. siemensii, a recent evaluation supports the A. siemensii species definition.
[23]
 Cast of the Maxberg Specimen
Composed of a torso, the Maxberg Specimen (S5) was discovered in 1956 or 1958 near
Langenaltheim and described in 1959 by Heller. It is currently missing, though it was once exhibited at the
Maxberg Museum in Solnhofen. It belonged to Eduard Opitsch, who loaned it to the museum. After his death
in 1991, the specimen was discovered to be missing and may have been stolen or sold. The specimen is
missing its head and tail, although the rest of the skeleton is mostly intact.
 Slab of the Haarlem Specimen
The Haarlem Specimen (TM 6428, also known as the Teyler Specimen) was discovered in 1855 near
Riedenburg, Germany and described as a Pterodactylus crassipes in 1875 by von Meyer. It was reclassified
in 1970 by John Ostrom and is currently located at the Teylers Museum in Haarlem, the Netherlands. It was
the very first specimen, despite the classification error. It is also one of the least complete specimens,
consisting mostly of limb bones and isolated cervical vertebrae and ribs.
The Eichstätt Specimen (JM 2257) was discovered in 1951 or 1955 near Workerszell, Germany and
described by Peter Wellnhofer in 1974. Currently located at the Jura Museum in Eichstätt, Germany, it is the
smallest specimen and has the second best head. It is possibly a separate genus ( Jurapteryx recurva) or
species (A. recurva).
 The Solnhofen Specimen (BSP 1999) was discovered in the 1960s near Eichstätt, Germany and
described in 1988 by Wellnhofer. Currently located at the Bürgermeister-Müller-Museum in Solnhofen, it was
originally classified as Compsognathus by an amateur collector. It is the largest specimen known and may
belong to a separate genus and species, Wellnhoferia grandis. It is missing only portions of the neck, tail,
backbone, and head.

The Munich Specimen

The Munich Specimen (S6, formerly known as the Solnhofen-Aktien-Verein Specimen) was
discovered in 1991 near Langenaltheim and described in 1993 by Wellnhofer. It is currently located at the
Paläontologisches Museum München in Munich. What was initially believed to be a bony sternum turned out
to be part of the coracoid,[43] but a cartilaginous sternum may have been present. Only the front of its face is
missing. It may be a new species, A. bavarica.
 Daiting Specimen
An eighth, fragmentary specimen was discovered in the late 1980s, not in Solnhofen limestone, but
in somewhat younger sediments at Daiting, Suevia. It is therefore known as the Daiting Specimen, and has
been known since 1996 only from a cast, briefly shown at the Naturkundemuseum in Bamberg. Long having
been missing and therefore dubbed the 'Phantom', it was purchased by palaeontologist Raimund
Albertsdörfer in 2009.[44] It was on display for the first time with six other original fossils of Archaeopteryx at
the Munich Mineral Show in October 2009.[45] A first, quick look by scientists indicate that this specimen
might represent a new species of Archaeopteryx.[46] It was found in a limestone bed that was a few hundred
thousand years younger than the other finds.[44]
 Bürgermeister-Müller ("chicken wing") Specimen
Another fragmentary fossil was found in 2000. It is in private possession and on loan to the
Bürgermeister-Müller Museum in Solenhofen, so it is called the Bürgermeister-Müller Specimen. As the
fragment represents the remains of a single wing of Archaeopteryx, the popular name of this fossil is "chicken
wing".
Long in a private collection, the Thermopolis Specimen (WDC CSG 100) was discovered in Germany
and described in 2005 by Mayr, Pohl, and Peters. Donated to the Wyoming Dinosaur Center in Thermopolis,
Wyoming, it has the best-preserved head and feet; most of the neck and the lower jaw have not been
preserved. The "Thermopolis" specimen was described in the December 2, 2005 Science journal article as
"A well-preserved Archaeopteryx specimen with theropod features"; it shows that the Archaeopteryx lacked a
reversed toe—a universal feature of birds—limiting its ability to perch on branches and implying a terrestrial or
trunk-climbing lifestyle.[47] This has been interpreted as evidence of theropod ancestry. The specimen also
has a hyperextendible second toe. "Until now, the feature was thought to belong only to the species' close
relatives, the deinonychosaurs."[48] In 1988, Gregory S. Paul claimed to have found evidence of a
hyperextensible toe, but this was not verified and accepted by other scientists until the Thermopolis
specimen was described.[49]

The Thermopolis Specimen

This tenth and latest specimen was assigned to Archaeopteryx siemensii in 2007.[50] The specimen
itself, currently on loan to the Royal Tyrrell Museum, Drumheller, Alberta, Canada, is considered the most
complete and well preserved Archaeopteryx remains yet.[50]
[edit] Taxonomy
Today, the fossils are usually assigned to a single species A. lithographica, but the taxonomic history
is complicated. Dozens of names have been published for the handful of specimens, most of which are
simply spelling errors (lapsus). Originally, the name A. lithographica only referred to the single feather
described by von Meyer. In 1960, Swinton proposed that the name Archaeopteryx lithographica be officially
transferred from the feather to the London specimen.[51] The ICZN did suppress the plethora of alternative
names initially proposed for the first skeleton specimens,[52] which mainly resulted from the acrimonious
dispute between von Meyer and his opponent Johann Andreas Wagner (whose Griphosaurus problematicus
—"problematic riddle-lizard"—was a vitriolic sneer at von Meyer's Archaeopteryx).[53] In addition, descriptions
of Archaeopteryx fossils as pterosaurs before their true nature was realized were also suppressed.[3][54]

Eichstätt Specimen, once considered a distinct genus, Jurapteryx

 The relationships of the specimens are problematic. Most subsequent specimens have been given
their own species at one point or another. The Berlin specimen has been designated as Archaeornis
siemensii, the Eichstätt specimen as Jurapteryx recurva, the Munich specimen as Archaeopteryx bavarica
and the Solnhofen specimen was designated as Wellnhoferia grandis.
 The Solnhofen Specimen, by some considered as belonging to the genus Wellnhoferia
Recently, it has been argued that all the specimens belong to the same species.[55] However,
significant differences exist among the specimens. In particular, the Munich, Eichstätt, Solnhofen and
Thermopolis specimens differ from the London, Berlin, and Haarlem specimens in being smaller or much
larger, having different finger proportions, having more slender snouts, lined with forward-pointing teeth and
possible presence of a sternum. These differences are as large as or larger than the differences seen today
between adults of different bird species. However, it is also possible that these differences could be
explained by different ages of the living birds.
Finally, it is worth noting that the feather, the first specimen of Archaeopteryx described, does not
agree well with the flight-related feathers of Archaeopteryx. It certainly is a flight feather of a contemporary
species, but its size and proportions indicate that it may belong to another, smaller species of feathered
theropod, of which only this feather is so far known.[39] As the feather was the original type specimen, this
has created significant nomenclatorial confusion.

[edit] Synonyms
If two names are given, the first denotes the original describer of the "species", the second the author
on whom the given name combination is based. As always in zoological nomenclature, putting an author's
name in parentheses denotes that the taxon was originally described in a different genus.
• Pterodactylus crassipes Meyer, 1857 [suppressed in favor of A. lithographica 1977 per ICZN
Opinion 1070]
 • Rhamphorhynchus crassipes (Meyer, 1857) (as Pterodactylus (Rhamphorhynchus)
crassipes) [suppressed in favor of A. lithographica 1977 per ICZN Opinion 1070]
• Archaeopteryx lithographica Meyer, 1861 [nomen conservandum]
• Scaphognathus crassipes (Meyer, 1857) Wagner, 1861 [suppressed in favor of A.
lithographica 1977 per ICZN Opinion 1070]
• Archaeopterix lithographica Anon., 1861 [lapsus]
• Griphosaurus problematicus Wagner, 1861 [nomen oblitum 1961 per ICZN Opinion 607]
• Griphornis longicaudatus Woodward, 1862 [nomen oblitum 1961 per ICZN Opinion 607]
• Griphosaurus longicaudatum (Woodward, 1862) [lapsus]
• Griphosaurus longicaudatus (Owen, 1862) [nomen oblitum 1961 per ICZN Opinion 607]
• Archaeopteryx macrura Owen, 1862 [nomen oblitum 1961 per ICZN Opinion 607]
• Archaeopterix macrura Owen, 1862 [lapsus]
• Archaeopterix macrurus Egerton, 1862 [lapsus]
• Archeopteryx macrurus Owen, 1863 [unjustified emendation]
• Archaeopteryx macroura Vogt, 1879 [lapsus]
• Archaeopteryx siemensii Dames, 1897
• Archaeopteryx siemensi Dames, 1897 [lapsus]
• Archaeornis siemensii (Dames, 1897) Petronievics, 1917[50]
• Archaeopteryx oweni Petronievics, 1917 [nomen oblitum 1961 per ICZN Opinion 607]
• Gryphornis longicaudatus Lambrecht, 1933 [lapsus]
• Gryphosaurus problematicus Lambrecht, 1933 [lapsus]
• Archaeopteryx macrourus Owen, 1862 fide Lambrecht, 1933 [lapsus]
 • Archaeornis siemensi (Dames, 1897) fide Lambrecht, 1933? [lapsus]
• Archeopteryx macrura Ostrom, 1970 [lapsus]
• Archaeopteryx crassipes (Meyer, 1857) Ostrom, 1972 [suppressed in favor of A.
lithographica 1977 per ICZN Opinion 1070]
• Archaeopterix lithographica di Gregorio, 1984 [lapsus]
• Archaeopteryx recurva Howgate, 1984
• Jurapteryx recurva (Howgate, 1984) Howgate, 1985
• Archaeopteryx bavarica Wellnhofer, 1993
• Wellnhoferia grandis Elżanowski, 2001
The last 4 taxa may be valid genera and species.
"Archaeopteryx" vicensensis (Anon. fide Lambrecht, 1933) is a nomen nudum for what appears to be
an undescribed pterosaur.

[edit] Controversies
[edit] Authenticity
Beginning in 1985, a group including astronomer Fred Hoyle and physicist Lee Spetner published a
series of papers claiming that the feathers on the Berlin and London specimens of Archaeopteryx were
forged.[56][57][58][59] Their claims were repudiated by Alan J. Charig and others at the British Museum
(Natural History).[60] Most of their evidence for a forgery was based on unfamiliarity with the processes of
lithification; for example, they proposed that based on the difference in texture associated with the feathers,
feather impressions were applied to a thin layer of cement,[57] without realizing that feathers themselves
would have caused a textural difference.[60] They also expressed disbelief that slabs would split so
smoothly, or that one half of a slab containing fossils would have good preservation, but not the counterslab.
[56][58] These, though, are common properties of Solnhofen fossils because the dead animals would fall
onto hardened surfaces which would form a natural plane for the future slabs to split along, leaving the bulk
of the fossil on one side and little on the other.[60] They also misinterpreted the fossils, claiming that the tail
was forged as one large feather,[57] when this is visibly not the case.[60] In addition, they claimed that the
other specimens of Archaeopteryx known at the time did not have feathers,[56][57] which is untrue; the
Maxberg and Eichstätt specimens have obvious feathers.[60] Finally, the motives they suggested for a
forgery are not strong, and contradictory; one is that Richard Owen wanted to forge evidence in support of
Charles Darwin's theory of evolution, which is unlikely given Owen's views toward Darwin and his theory. The
other is that Owen wanted to set a trap for Darwin, hoping the latter would support the fossils so Owen could
discredit him with the forgery; this is unlikely because Owen himself wrote a detailed paper on the London
specimen, so such an action would certainly backfire.[61]
Charig et al. pointed to the presence of hairline cracks in the slabs running through both rock and
fossil impressions, and mineral growth over the slabs that had occurred before discovery and preparation, as
evidence that the feathers were original.[60] Spetner et al. then attempted to show that the cracks would
have naturally propagated through their postulated cement layer,[62] but neglected to account for the fact
that the cracks were old and had been filled with calcite, and thus were not able to propagate.[61] They also
attempted to show the presence of cement on the London specimen through X-ray spectroscopy, and did find
something that was not rock.[62] However, it was not cement, either, and is most probably from a fragment of
silicone rubber left behind when molds were made of the specimen.[61] Their suggestions have not been
taken seriously by palaeontologists, as their evidence was largely based on misunderstandings of geology,
and they never discussed the other feather-bearing specimens, which have increased in number since then.
Charig et al. reported a discoloration: a dark band between two layers of limestone – however, they say it is
the product of sedimentation.[60] It is natural for limestone to take on the color of its surroundings and most
limestones are colored (if not color banded) to some degree[63] – the darkness was attributed to such
impurities. They also mention that a complete absence of air bubbles in the rock slabs is further proof that the
specimen is authentic.[60]

[edit] Archaeopteryx and Protoavis

In 1984, Sankar Chatterjee discovered fossils which he claimed in 1991 belonged to a fossil bird far
older than Archaeopteryx. These fossils, believed to be around 210 to 225 million years old, have been
assigned the name Protoavis.[64] The fossils are too badly preserved to allow an estimate of flying ability;
although Chatterjee's reconstructions usually show feathers, many paleontologists, including Paul (2002) and
Witmer (2002) have rejected the claims that Protoavis was an earlier bird (or, alternately, that it existed at
all).[35][65] The fossils were found disarticulated, and were collected from different locations. Because the
fossils are in poor condition, Archaeopteryx remains the earliest universally recognized bird.[66]

[edit] Phylogenetic position

 Growth depictions for Archaeopteryx used to compare the growth of Archaeopteryx to dinosaurs and
birds
Modern paleontology has consistently placed Archaeopteryx as the most primitive bird. It is not
thought to be a true ancestor of modern birds but, rather, a close relative of that ancestor (see Avialae and
Aves).[67]

Skeletal elements from the largest specimen showing growth lines that were used to compare the
growth of Archaeopteryx to dinosaurs and birds
Nonetheless, Archaeopteryx is so often used as a model of the true ancestral bird that it has seemed
almost heretical to suggest otherwise. Several authors have done so.[65] Lowe (1935)[68] and Thulborn
(1984)[69] questioned whether Archaeopteryx truly was the first bird. They suggested that Archaeopteryx
was a dinosaur that was no more closely related to birds than were other dinosaur groups. Kurzanov (1987)
suggested that Avimimus was more likely to be the ancestor of all birds than Archaeopteryx.[70] Barsbold
(1983)[71] and Zweers and Van den Berge (1997)[72] noted that many maniraptoran lineages are extremely
birdlike, and suggested that different groups of birds may have descended from different dinosaur ancestors.
A histological study from 2009 showed that Archaeopteryx grew slowly like dinosaurs, whereas the
fast growth of modern birds was achieved later in bird evolution.[73]

[edit] In popular culture

Archaeopteryx is the best known early bird and has thus received widespread attention. Its easily
recognizable appearance and the intense public interest in dinosaurs have caused Archaeopteryx to become
a feature of worldwide popular culture. For example, the second book in the Time Machine series, Search for
Dinosaurs, takes the reader on a journey to the Mesozoic to find and photograph an Archaeopteryx.[74] In
one of the "strangest" appearances of Archaeopteryx in popular culture, Alfred Jarry's play Ubu cocu, ou
l'Archéopteryx ('Ubu cuckolded, or the Archaeopteryx') includes an Archaeopteryx as an important character.
[75] A main belt asteroid discovered in 1991, 9860 Archaeopteryx, was named in honour of the genus.[76]
[77]

[edit] See also

Paleontology portal

Dinosaurs portal
 • Dinosaur
• Origin of birds
• Feathered dinosaurs
• Temporal paradox (paleontology)

[edit] References
1. ^ Yalden D.W. (1984). "What size was Archaeopteryx?". Zoological Journal of the Linnean
Society 82 (1-2): 177–188. doi:10.1111/j.1096-3642.1984.tb00541.x.
http://www3.interscience.wiley.com/journal/120801944/abstract.
2. ^ Chiappe L.M., Witme L.M. (2002). University od California Press Berkeley and Los
Angeles, California. ed. Mesozoic birds: above the heads of dinosaurs. University od California
Press. pp. 151. ISBN 0-520-20094-2. http://books.google.com/?id=2MQeh1KCp7sC.
3. ^ a b Archaeopteryx: An Early Bird - University of California, Berkeley Museum of
Paleontology. Retrieved 2006-OCT-18
4. ^ Archaeopteryx lithographica - Nick Longrich, University of Calgary. Discusses how many
wings an Archaeopteryx had and other questions.
5. ^ Wellnhofer P (2004). "The Plumage of Archaeopteryx". in Currie PJ, Koppelhus EB, Shugar
MA, Wright JL. Feathered Dragons. Indiana University Press. pp. 282–300. ISBN 0-253-34373-9.
6. ^ Schweigert, G. (2007). "Ammonite biostratigraphy as a tool for dating Upper Jurassic
lithographic limestones from South Germany – first results and open questions." Neues Jahrbuch für
Geologie und Paläontologie - Abhandlungen, 245(1): 117-125. doi:10.1127/0077-7749/2007/0245-
0117
7. ^ a b c d Lambert, David (1993). The Ultimate Dinosaur Book. New York: Dorling Kindersley.
pp. 38–81. ISBN 1-56458-304-X.
8. ^ Kennedy, Elaine (2000). Solnhofen Limestone: Home of Archaeopteryx. Geoscience
Reports. 30: 1–4. Retrieved 2006-10-18.
9. ^ Holtz, Thomas, Jr. (1995). "Archaeopteryxs Relationship With Modern Birds". Journal of
Dinosaur Paleontology. http://www.dinosauria.com/jdp/archie/archie.htm. Retrieved 2007-03-01.
10.^ Palaeogeography, Palaeoclimatology, Palaeoecology 130 (1997) 275-292
11.^ Bühler, P. & Bock, W.J. (2002). Zur Archaeopteryx-Nomenklatur: Missverständnisse und
Lösung. Journal of Ornithology. 143(3): 269–286. [Article in German, English abstract]
doi:10.1046/j.1439-0361.2002.02006.x (HTML abstract)
12.^ Huxley T.H. (1868). On the animals which are most nearly intermediate between birds and
reptiles. Geol. Mag. 5, 357–65; Annals & Magazine of Nat Hist 2, 66–75; Scientific Memoirs 3, 3–13.
13.^ Huxley T.H. (1868) Remarks upon Archaeopteryx lithographica. Proc Roy Soc 16, 243–48;
Sci Memoirs 3, 340-45.
14.^ Huxley T.H. (1870) Further evidence of the affinity between the dinosaurian reptiles and
birds. Quart J Geol Soc 26, 32–50; Sci Mem 3, 487–509.
15.^ Nedin, C. (1999). All About Archaeopteryx. talk.origins archive. Version of June 10, 2002;
retrieved 2006-10-18.
16.^ Olson, S.L. & Feduccia, A. (1979). Flight capability and the pectoral girdle of
Archaeopteryx. Nature. 278(5701). 247–248. doi:10.1038/278247a0 (HTML abstract)
17.^ Ostrom, J.H. (1976). Archaeopteryx and the origin of birds. Biol. J. Linn. Soc.. 8: 91–182.
 18.^ Ostrom, J.H. (1985). Introduction to Archaeopteryx. In: Hecht, M.K.O.; Ostrom, J.H.; Viohl,
G. & Wellnhofer, P. (eds.) The Beginnings of Birds: Proceedings of the International Archaeopteryx
Conference: 9–20. Eichstätt, Freunde des Jura-Museums Eichstätt.
19.^ Owen, R. (1863). On the Archaeopteryx of Von Meyer, with a description of the fossil
remains of a long-tailed species from the lithographic stone of Solnhofen . Phil. Trans. Roy. Soc.
London. 153: 33–47.
20.^ Feduccia, A. & Tordoff, H.B. (1979). Feathers of Archaeopteryx: Asymmetric vanes
indicate aerodynamic function. Science. 203(4384): 1021–1022.
21.^ a b Christensen P, Bonde N. (2004). Body plumage in Archaeopteryx: a review, and new
evidence from the Berlin specimen. Comptes Rendus Palevol. 3: 99–118. PDF fulltext
22.^ a b Longrich N. (2006): Structure and function of hindlimb feathers in Archaeopteryx
lithographica. Paleobiology. 32(3): 417–431. doi:10.1666/04014.1 (HTML abstract)
23.^ a b c Elżanowski A. (2002): Archaeopterygidae (Upper Jurassic of Germany). In: Chiappe,
L. M. & Witmer, L. M (eds.), Mesozoic Birds: Above the Heads of Dinosaurs : 129–159. University of
California Press, Berkeley.
24.^ Senter, P. (2006). Scapular orientation in theropods and basal birds and the origin of
flapping flight. Acta Palaeontologica Polonica. 51(2): 305–313. PDF fulltext
25.^ Thomson and Speakman (1994).
26.^ Nudds, Robert L. & Dyke, Gareth J. (May 14, 2010). "Narrow Primary Feather Rachises in
Confuciusornis and Archaeopteryx Suggest Poor Flight Ability". Science 328 (5980): 887–889.
doi:10.1126/science.1188895. PMID 20466930.
27.^ Science Magazine News: "Did First Feathers Prevent Early Flight?", 13-5-2010.
28.^ Witmer, L. M. (2004). Palaeontology: Inside the oldest bird brain . Nature. 430(7000): 619–
620. PMID 15295579 doi:10.1038/430619a
 29.^ Alonso, P. D., Milner, A. C., Ketcham, R. A., Cookson, M. J. & Rowe, T. B. (2004). The
avian nature of the brain and inner ear of Archaeopteryx. Nature. 430(7000): 666–669. PMID
15295597. doi:10.1038/nature02706. PDF fulltext Supplementary info
30.^ a b c d Chiappe, Luis M. (2007). Glorified Dinosaurs. Sydney: UNSW Press. pp. 118–146.
ISBN 0-471-24723-5.
31.^ Davis, P.; and Briggs, D. (1998). "The impact of decay and disarticulation on the
preservation of fossil birds". PALAIOS (PALAIOS, Vol. 13, No. 1) 13 (1): 3–13. doi:10.2307/3515277.
http://palaios.sepmonline.org/cgi/content/abstract/13/1/3. Retrieved 2007-03-25.
32.^ a b Bartell K.W., Swinburne N.H.M. and Conway-Morris S. 1990. Solnhofen: a study in
Mesozoic palaeontology. Cambridge (transl. and revised from Bartel K.W. 1978. Ein Blick in die
Erdgeschichte. Ott.
33.^ a b Paul, Gregory S. (2002). Dinosaurs of the Air: the Evolution and Loss of Flight in
Dinosaurs and Birds. Baltimore: Johns Hopkins University Press. ISBN 0-8018-6763-0.
34.^ Buisonje, P.H. de (1985). "Climatological conditions during deposition of the Solnhofen
limestones". in Hecht, M.K.; Ostrom, J.H.; Viohl, G.; and Wellnhofer, P. (eds.). The beginnings of
Birds: Proceedings of the International Archaeopteryx Conference, Eichstatt, 1984. Eichstätt:
Freunde des Jura-Museums Eichstätt. pp. 45–65. ISBN 978-3980117807.
35.^ a b Paul, Gregory S. (2002). Dinosaurs of the Air: The Evolution and Loss of Flight in
Dinosaurs and Birds. Baltimore: Johns Hopkins University Press. ISBN 0801867630.
36.^ Ostrom, J.H. (1976). "Archaeopteryx and the origin of birds". Biological Journal of the
Linnean Society 8: 91–182. doi:10.1111/j.1095-8312.1976.tb00244.x.
37.^ Feduccia, A. (1993). "Evidence from claw geometry indicating arboreal habits of
Archaeopteryx." Science. 259(5096): 790–793.
 38.^ National Geographic News- Earliest Bird Had Feet Like Dinosaur, Fossil Shows - Nicholas
Bakalar, December 1, 2005, Page 1. Retrieved 2006-10-18.
39.^ a b Griffiths, P. J. (1996). The Isolated Archaeopteryx Feather. Archaeopteryx 14: 1–26.
40.^ British Museum of Natural History - 'BMNH 37001' - the type specimen
41.^ Darwin, Origin of Species, Chapter 9, p. 367
42.^ Darwin, Charles (1859). On the Origin of Species. John Murray.
http://www.readprint.com/chapter-2217/Charles-Darwin. . Please note Darwin's spelling:
'Archeopteryx', not 'Archaeopteryx'.
43.^ Wellnhofer, P. & Tischlinger, H. (2004). Das "Brustbein" von Archaeopteryx bavarica
Wellnhofer 1993 - eine Revision. Archaeopteryx. 22: 3–15. [Article in German]
44.^ a b Archäologischer Sensationsfund in Daiting, (German) Augsburger Allgemeine -
Donauwörth edition, published: 28 November 2009, accessed: 23 December 2009
45.^ Sammler und Forscher - ein schwieriges Verhältnis (German), Sueddeutsche Zeitung,
published: 25 October 2009, accessed: 25 December 2009
46.^ Wiedergefundener Archaeopteryx ist wohl neue Art (German) Die Zeit, accessed: 25
December 2009
47.^ Mayr G, Pohl B & Peters DS. (2005). A well-preserved Archaeopteryx specimen with
theropod features. Science. 310(5753): 1483–1486. doi:10.1126/science.1120331 See commentary
on article
48.^ National Geographic News- Earliest Bird Had Feet Like Dinosaur, Fossil Shows - Nicholas
Bakalar, December 1, 2005, Page 2. Retrieved 2006-10-18.
49.^ Paul, G.S. (1988). Predatory Dinosaurs of the World, a Complete Illustrated Guide . New
York: Simon and Schuster. 464 p.
 50.^ a b c Mayr, G., Phol, B., Hartman, S. & Peters, D.S. (2007). The tenth skeletal specimen of
Archaeopteryx. Zoological Journal of the Linnean Society, 149, 97–116.
51.^ Swinton, W. E. (1960). Opinion 1084, Proposed addition of the generic name
Archaeopteryx VON MEYER, 1861 and the specific name Lithographica, VON MEYER, 1861, as
published in the binomen Archaeopteryx Lithographica to the official lists (Class Aves). Bulletin of
Zoological Nomenclature 17(6–8): 224–226.
52.^ ICZN. (1961). Opinion 607, Archaeopteryx VON MEYER, 1861 (Aves); Addition to the
Official list. Bulletin of Zoological Nomenclature 18(4): 260–261.
53.^ Wagner A (1861) Über ein neues, angeblich mit Vogelfedern versehenes Reptil aus dem
Solnhofener lithographischen Schiefer. Sitzungberichte der Bayerischen Akademie der
Wissenschaften, mathematisch-physikalisch Classe 146–154
54.^ ICZN. (1977). Opinion 1070. Conservation of Archaeopteryx lithographica VON MEYER
1861 (Aves). Bulletin of Zoological Nomenclature 33: 165–166.
55.^ Archaeopteryx turns out to be singular bird of a feather. New Scientist 2443:17. 17 April
2004. See commentary on article.
56.^ a b c Hoyle, F.; Wickramasinghe, N.C.; and Watkins, R.S. (1985). " Archaeopteryx". British
Journal of Photography 132: 693–694.
57.^ a b c d Watkins, R.S.; Hoyle, F.; Wickrmasinghe, N.C.; Watkins, J.; Rabilizirov, R.; and
Spetner, L.M. (1985). "Archaeopteryx - a photographic study". British Journal of Photography 132:
264–266.
58.^ a b Watkins, R.S.; Hoyle, F.; Wickrmasinghe, N.C.; Watkins, J.; Rabilizirov, R.; and Spetner,
L.M. (1985). "Archaeopteryx - a further comment". British Journal of Photography 132: 358–359, 367.
59.^ Watkins, R.S.; Hoyle, F.; Wickrmasinghe, N.C.; Watkins, J.; Rabilizirov, R.; and Spetner,
L.M. (1985). "Archaeopteryx - more evidence". British Journal of Photography 132: 468–470.
 60.^ a b c d e f g h Charig, A.J.; Greenaway, F.; Milner, A.N.; Walker, C.A.; and Whybrow, P.J.
(1986). "Archaeopteryx is not a forgery". Science 232 (4750): 622–626.
doi:10.1126/science.232.4750.622. PMID 17781413.
61.^ a b c Nedin, Chris (2007-12-15). "On Archaeopteryx, Astronomers, and Forgery".
http://www.talkorigins.org/faqs/archaeopteryx/forgery.html. Retrieved 2007-03-17.
62.^ a b Spetner, L.M.; Hoyle, F.; Wickramasinghe, N.C.; and Magaritz, M. (1988).
"Archaeopteryx - more evidence for a forgery". The British Journal of Photography 135: 14–17.
63.^ http://encarta.msn.com/encyclopedia_761565838/limestone_(mineral).html as at 13-08-09.
Archived 2009-10-31.
64.^ Chatterjee, Sankar (1991). "Cranial anatomy and relationships of a new Triassic bird from
Texas". Phil. Trans. R. Soc. B 332 (1265): 277–342. doi:10.1098/rstb.1991.0056.
65.^ a b Witmer, Lawrence M. (2002). "The debate on avian ancestry". in Witmer, L.; Chiappe,
L.. Mesozoic Birds: Above the Heads of Dinosaurs. Berkeley: University of California Press. pp. 3–30.
ISBN 0520200942.
66.^ Ostrom, J. H. (1996). "The questionable validity of Protoavis". Archaeopteryx 14: 39–42.
67.^ Clarke, Julia. A.; Norell, Mark. A. (2002). "The Morphology and Phylogenetic Position of
Apsaravis ukhaana from the Late Cretaceous of Mongolia". American Museum Novitates 3387 (1): 1–
46. doi:10.1206/0003-0082(2002)387<0001:TMAPPO>2.0.CO;2.
68.^ Lowe, P. R. (1935). "On the relationship of the Struthiones to the dinosaurs and to the rest
of the avian class, with special reference to the position of Archaeopteryx". Ibis 5 (2): 398–432.
doi:10.1111/j.1474-919X.1935.tb02979.x.
69.^ Thulborn, R. A. (1984). "The avian relationships of Archaeopteryx, and the origin of birds".
Zoological Journal of the Linnean Society 82 (1-2): 119–158. doi:10.1111/j.1096-
3642.1984.tb00539.x.
 70.^ Kurzanov, S. M. (1987). "Avimimidae and the problem of the origin of birds". Transactions
of the joint Soviet-Mongolian Paleontological Expedition 31: 31–94. ISSN 0320-2305.
71.^ Barsbold, Rhinchen (1983). "Carnivorous dinosaurs from the Cretaceous of Mongolia".
Transactions of the joint Soviet-Mongolian Paleontological Expedition 19: 5–119. ISSN 0320-2305.
72.^ Zweers, G. A.; Van den Berge, J.C. (1997). "Evolutionary patterns of avian trophic
diversification". Zoology: Analysis of Complex Systems 100: 25–57. ISSN 0944-2006.
73.^ Erickson, Gregory M.; Rauhut, Oliver W. M., Zhou, Zhonghe, Turner, Alan H, Inouye, Brian
D. Hu, Dongyu, Norell, Mark A. (2009). "Was Dinosaurian Physiology Inherited by Birds? Reconciling
Slow Growth in Archaeopteryx". PLoS One.
doi:10.1371/journal.pone.0007390;jsessionid=F7E462DE00439EEA45BCC1AF96012EE0.
http://www.plosone.org/article/info%3Adoi
%2F10.1371%2Fjournal.pone.0007390;jsessionid=F7E462DE00439EEA45BCC1AF96012EE0.
Retrieved 2009-10-25.
74.^ Bischoff, David; Search for Dinosaurs (Time Machine, No. 2); published 1984 by Bantam
Books; introduction
75.^ Buffetaut, E. (1985). The strangest interpretation of Archaeopteryx In: Hecht, M.K.O.;
Ostrom, J.H.; Viohl, G. & Wellnhofer, P. (eds.) The Beginnings of Birds: Proceedings of the
International Archaeopteryx Conference: 369-370. Eichstätt, Freunde des Jura-Museums Eichstätt.
76.^ "JPL Small-Body Database Browser: 9860 Archaeopteryx (1991 PW9)". NASA.
http://ssd.jpl.nasa.gov/sbdb.cgi?sstr=9860. Retrieved 2007-03-01.
77.^ Williams, Gareth. "Minor Planet Names: Alphabetical List". Smithsonian Astrophysical
Observatory. http://www.cfa.harvard.edu/iau/lists/MPNames.html. Retrieved 2007-03-01.
[edit] Further reading
• de Beer, G.R. (1954). Archaeopteryx lithographica: a study based upon the British Museum
specimen. Trustees of the British Museum, London.
• Chambers, P. (2002). Bones of Contention: The Fossil that Shook Science. John Murray,
London. ISBN 0-7195-6059-4.
• Feduccia, A. (1996). The Origin and Evolution of Birds. Yale University Press, New Haven.
ISBN 0-300-06460-8.
• Heilmann, G. (1926). The Origin of Birds. Witherby, London.
• Huxley T.H. (1871). Manual of the anatomy of vertebrate animals. London.
• von Meyer, H. (1861). Archaeopteryx litographica (Vogel-Feder) und Pterodactylus von
Solenhofen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 1861:
678–679, plate V [Article in German] Fulltext at Google Books.
• Shipman, P. (1998). Taking Wing: Archaeopteryx and the Evolution of Bird Flight . Weidenfeld
& Nicolson, London. ISBN 0-297-84156-4.
• Wellnhofer, P. (2008). Archaeopteryx. Der Urvogel von Solnhofen (in German). Verlag
Friedrich Pfeil, Munich. ISBN 978-389937076-8

[edit] External links

Wikispecies has information related to: Archaeopteryx
 Wikimedia Commons has media related to: Archaeopteryx

• Journal of Dinosaur Paleontology – With many articles on dinosaur-bird links.

• All About Archaeopteryx from Talk.Origins.
• Use of SSRL X-ray takes 'transformative glimpse' – A look at chemicals linking birds and
dinosaurs.

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Birds (class: Aves)

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Hybrids

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Superorder: Palaeognathae
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 Struthioniformes (ratites) • Tinamiformes (tinamous)

Superorder: Neognathae

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Retrieved from "http://en.wikipedia.org/wiki/Archaeopteryx"

Categories: Archaeopterygidae | Dinosaurs of Europe | Genera of birds | Jurassic birds | Jurassic

dinosaurs | Solnhofen fauna | Transitional fossil | Greek loanwords | Prehistoric birds of Europe | Animals
described in 1861

Birds (class: Aves)

Anatomy Bird anatomy • Flight • Eggs • Feathers • Plumage • Beak • Vision • Dactyly •
Preen gland

Behaviour Singing • Intelligence • Migration • Incubation • Brood parasites • Nesting •

Hybrids
 Evolution Evolution of birds • Origin of birds • Darwin's finches • Seabirds

Superorder: Palaeognathae

Struthioniformes (ratites) • Tinamiformes (tinamous)

Superorder: Neognathae

Accipitriformes • Anseriformes (waterfowl) • Apodiformes (swifts and

Subclass: hummingbirds) • Caprimulgiformes (nightjars and relatives) • Cariamae (seriemas and
Neornithes relatives) • Charadriiformes (gulls and relatives) • Ciconiiformes (storks) • Coliiformes
(29 orders of (mousebirds) • Columbiformes (doves and pigeons) • Coraciiformes (kingfishers and
modern birds) relatives) • Cuculiformes (cuckoos and relatives) • Falconiformes (falcons and
relatives) • Galliformes (gamebirds) • Gaviiformes (loons or divers) • Gruiformes
(cranes and relatives) • Passeriformes (perching birds) • Pelecaniformes (pelicans
and relatives) • Phaethontiformes (tropicbirds) • Phoenicopteriformes (flamingos) •
Piciformes (woodpeckers and relatives) • Podicipediformes (grebes) •
Procellariiformes (albatrosses and petrels) • Psittaciformes (parrots) • Pteroclidiformes
(sandgrouses) • Sphenisciformes (penguins) • Strigiformes (owls) • Trogoniformes
(trogons and quetzals)

Fossil birds Archaeopteryx • Enantiornithes • Hesperornithes

 Birds and Ringing • Ornithology • Bird collections • Birdwatching • Bird feeding •
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Kingdom: Animalia · Phylum: Chordata · Class: Sauropsida · Subclass: Diapsida

Primitive Euparkeriidae • Erythrosuchidae • Proterochampsidae •

Archosauromorphs Proterosuchidae • Choristodera • Prolacertiformes • Rhynchosauria •
Trilophosauria • Vancleavea
 Crurotarsi Ornithosuchidae • Aetosauria • Phytosauria • Rauisuchia •
Archosaurs Crocodylomorpha • Crocodilia

Avemetatarsalia and Scleromochlus • Pterosauria • Dinosauromorpha • Dinosauria •

Ornithodira Archosaurs Ornithischia • Saurischia • Aves

Avian Archosaurs Avialae • Archaeopteryx • Confuciusornis • Ichthyornis •

Enantiornithes • Hesperornithes • Neornithes • Palaeognathae • Neognathae
 W000

Évolution du flagelle
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

La compréhension de l´évolution du flagelle est de première importance pour les biologistes puisque
les trois différents flagelles eucaryotes et procaryotes sont des structures cellulaires sophistiquées et
complexes permettant aux cellules de se mouvoir dans leur milieu. Il est important de préciser que les
flagelles bactériens et eucaryotes ne sont pas analogues et n´ont aucun lien de parenté.
Sommaire
[masquer]
• 1 Le flagelle eucaryote
• 1.1 Le modèle symbiotique
• 1.2 Le modèle endogène
• 2 Le flagelle bactérien
• 3 Le flagelle des Archées
• 4 Notes et références
• 5 Voir aussi
Le flagelle eucaryote [modifier]

flagelle chez un eucaryote.

Il existe deux modèles distincts concernant l´origine du cil et du flagelle eucaryote.
Le modèle symbiotique [modifier]
Ce modèle, également appelé endosymbiotique ou exogène, stipule qu´une bactérie spirochaetes,
en symbiose avec une cellule eucaryote ou une Archée primitive, serait à l´origine du cil et du flagelle
eucaryote[1]. Ainsi, le symbionte servirait d'organèle mobile. Cette hypothèse est l'oeuvre de Lynn Margulis,
microbiologiste américaine réconnue pour avoir émis et popularisé l´origine symbiotique des mitochondries
et des chloroplastes. Cependant, l´originie symbiotique du flagelle eucaryote est très critiquée par de
nombreux experts, dont Thomas Cavalier-Smith, qui sont plus favorables à une origine endogène[2].
Le principal argument en faveur de la théorie symbiotique est qu´il existe des espèces eucaryotes
unicellulaires résidant dans les intestins de termite tel que les Mixotricha et les Trichonympha qui possédent
un symbionte spirochaetes assurant leur mobilité. Malgré ces exemples de symbiose, l´absence d´homologie
entre les flagelles eucaryotes et les spirochètes est la principale critique contre cette hypothèse. De plus,
contrairement aux mitochondries et aux chloroplastes, les cils et les flagelles eucaryotes n´ont pas de
génome propre.

Le modèle endogène [modifier]

Ce modèle stipule que le cil et le flagelle eucaryote sont issus du cytosquelète eucaryote pré-existant
essentiel pour la formation du fuseau mitotique[3]. Les protéines de la famille des tubulines et dynéines sont
à la fois importantes pour la formation du fuseau mitotique et du flagelle eucaryote. Par ailleurs, dans
certaines espèces unicellulaires eucaryotes, le corps basal d´un flagelle eucaryote focalise et organise le
fuseau mitotique.
Soutenant l´hypothèse endogène du cil, l´étude de la structure en 3 dimensions de la tubuline a montré une
conformation similaire à celle de la protéine du cytosquelette bactérien FtsZ confirmant ainsi une vieille
filiation initialement basée sur de faibles mais significatives homologies de séquence[4]. Cette découverte
est essentielle car le cytosquelette des bactéries spirochaetes ne possède pas de protéine FtsZ et suggère
ainsi que les bactéries spirochaetes ne sont pas à l´origine du cil.
Néanmoins, il existe de nombreux cas d´évolution parallèle, également appelé évolution convergente, où
deux molécules ou bien deux structures sont apparues indépendamment l´une de l´autre. De ce fait, la
structure de la tubuline pourrait être le résultat d´une évolution parallèle[5].
Le flagelle bactérien [modifier]

Flagelle d'une bactérie Gram-

 L´origine du flagelle bactérien est plus étudiée que le flagelle eucaryote. Il semble que les
composants du flagelle bactérien aient évolué à partir du système de transport de type III qui permet la
sécrétion de protéine bactérienne[6]. La sécrétion de protéine dans le milieu extérieur est complexe et
implique de nombreux complexes moléculaires permettant le ciblage et le transport à travers la paroi
bactérienne et la membrane bactérienne. Dans le cas des gram négatif qui possèdent deux membranes, il
existe six systèmes de sécrétion appelés système de transport de type I à VI. Le système de transport de
type III est impliqué dans le transport et la sécrétion de toxines. Par exemple, le bacille Yersinia pestis, agent
pathogène responsable de la peste, injecte dans les cellules eucaryotes une toxine grâce au système de
transport de type III[7]. Le flagelle bactérien et le système de transport III dérive d´une structure ancestrale
commune probablement requise pour le transport des protéines. Les deux systèmes ont en commun neuf
proteines homologues. Par exemple, la proteine ATPase Fli1 du flagelle est homologue à la proteine T3SS
ATPase (qui signifie Type 3 Secretion System ATPase). Récemment, des études comparatives de
séquences et de structures crystallographiques ont mis en évidence une forte homologie entre la proteine
FliG, un composant du moteur du flagelle situé dans le disque C, et le transporteur d´ions Mg2+ MgtE[8].
Notons pour conclure, que la structure des flagelles bactériens diffère entre les bactéries Gram négatives et
positives. En effet, alors que les bactéries Gram négatives possèdent deux membranes, les bactéries Gram
positives ne possèdent qu´une seule membrane. Par conséquent, ayant une paroi bactérienne différente et
qu´une seule membrane, les bactéries gram positives n´ont pas les complexes constituant du flagelle
nommés disque P et disque L qui chez les bactéries Gram négatives intéragit respectivement avec la paroi
bactérienne et la membrane externe.

Le flagelle des Archées [modifier]

L´analyse des séquences protéiques et génétiques révèle que le flagelle des Archées est apparenté
au pilus de type IV[9]. Le flagelle des Archées est similaire mais ne présent pas d´homologie avec celui des
bactéries. En plus de l´absence d´homologie de séquence entre les protéines des deux systèmes, le flagelle
des Archées croît par l'addition de monomère de flagellines à la base du flagelle alors que l´addition de
flagellines s´effectue dans la partie distale du flagelle chez les bactéries. Par ailleurs, le flagelle des Archées
à un plus petit diamètre car il n´a pas de canal central.
Les pili, des appendices extracellulaires, peuvent se rétracter et participer à la mobilité bactérienne
par un mécanisme de glissade social (appelé social gliding ou bien twitching en anglais) en interagissant
avec une cellule hôte ou une autre bactérie. Le pilus de type IV est assemblé via le système de transport de
type II. Pour l´instant, aucune bactérie n´utilise de pili de type IV pour nager.

Notes et références [modifier]

1. ↑ Margulis, L. Chapman, M. Guerrero, R. Hall J. (2006) The last eukaryotic common
ancestor (LECA): acquisition of cytoskeletal motility from aerotolerant spirochetes in the Proterozoic
Eon Proc Natl Acad Sci U S A. Aug 29;103(35):13080-5
2. ↑ Cavalier-smith, T.(2006) Cell evolution and Earth history: stasis and revolution Philos
Trans R Soc Lond B Biol Sci. Jun 29;361(1470):969-1006
3. ↑ Bornens, M. Azimzadeh, J. (2007) Origin and evolution of the centrosome Adv Exp Med
Biol. 607:119-29. Review
4. ↑ Erickson, H.P. (2007 ) Evolution of the cytoskeleton Bioessays. Jul;29(7):668-77. Review
5. ↑ Edward H. Egelman (1998) Tubulin family: Kinship of key proteins across phylogenetic
domains, Current Biology, Vol8, Issue 8, Pages R288-R290
6. ↑ Pallen, J.M.;Penn, C.W.;Chaudhuri, R.R.(2005). Bacterial flagellar diversity in the post-
genomic era, Trends in Microbiology, Vol.13,n4,143-149
 7. ↑ Shao, F. Biochemical functions of Yersinia type III effectors Curr Opin Microbiol. 2008
Feb;11(1):21-9
8. ↑ Snyder, L.A.S. Loman, N.J. Fütterer, K. Pallen, M.J. Bacterial flagellar diversity and
evolution: seek simplicity and distrust it? Trends in microbiology vol.10(1):1-5 2009
9. ↑ Faguy, DM. Jarrell, KF. Kuzio, J Kalmokoff, M.L. (1994) Molecular analysis of archael
flagellins: similarity to the type IV pilin-transport superfamily widespread in bacteria , Can J Microbiol,
Jan;40(1); 67-71 Review

Voir aussi [modifier]

• Cell evolution and earth history
• Flagellum Evolution at The pandas thumb.
• Evolution of the bacterial flagella

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique Dérive génétique · Mutation génétique · Recombinaison

évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens
 Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des
Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

• Portail de la microbiologie
Ce document provient de « http://fr.wikipedia.org/wiki/%C3%89volution_du_flagelle ».

Catégorie : Histoire évolutive | [+]

 W000

Céphalisation
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Cet article est une ébauche concernant la biologie.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
La céphalisation désigne la réunion des organes sensoriels vers l'avant d'un animal en même temps
qu'une concentration de son système nerveux, ce qui forme alors une tête.
Dans le monde animal [modifier]
La céphalisation se retrouve chez bon nombre d'animaux bilatériens : ils possèdent en effet une
symétrie bilatérale, et ont par conséquent trois axes de polarité : gauche et droite, dos et ventre et avant et
arrière. À l'avant s'ouvre la bouche, à l'arrière l'anus. La tête, quand elle est présente, est à l'avant et inclut la
bouche.
La plupart de ces bilatériens possèdent une tête, plus ou moins différenciée. Certains, comme les
bivalves, n'ont pas de tête centralisée. Le système nerveux n'est pas toujours concentré : il est souvent
simple et un vrai cerveau est plus rare, on en trouve en particulier chez les vertébrés ainsi que chez les
céphalopodes (comme le poulpe).
Chez les crâniates (dont font partie les vertébrés) la tête est protégée par un crâne. Chez les
gnathostomes (vertébrés à mâchoire) une mâchoire se différencie et est incluse dans la tête.

Chez les arthropodes [modifier]

Article détaillé : tête de l'insecte.
La tête est individualisée chez les hexapodes.
Elle est fusionnée avec le thorax pour former le céphalothorax, chez les crustacés et arachnides.

Chez l'homme [modifier]

Article détaillé : tête humaine.
 • Portail de la biologie
Ce document provient de « http://fr.wikipedia.org/wiki/C%C3%A9phalisation ».

Catégorie : Histoire évolutive | [+]

 w000

Histoire évolutive des gastéropodes

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Les premiers gastéropodes connus sont exclusivement marins, les plus anciens représentants du
groupe, les genres Chippewaella et Strepsodiscus, ont été datés de la fin du Cambrien.[réf. nécessaire]
Leurs parents les plus proches vivant seraient les céphalopodes.
Les conchifères forment un groupe d'animaux bien distinct des mollusques qui ne regroupaient à
l'époque que les animaux à corps mous sans coquille. Les formes de mollusque à coquille du début du
Cambrien, comme Helcionella et Scenella ne sont pas considérées comme des gastéropodes. Aldanella ,
qui produit pourtant une coquille, n'est quant à elle probablement même pas un mollusque.
Durant l'Ordovicien, les genres de gastéropodes découverts sont très variés mais tous toujours
aquatiques. En général, les fossiles de gastéropodes découverts dans les roches du début du paléozoïque
sont trop mal conservés pour permettre une identification précise, ils sont en outre, moins présents que ceux
des bivalves. Pourtant, quinze espèces du Silurien, du genre Poleumita, ont pu être identifiées.
La plupart des gastéropodes de l'ère paléozoïque appartiennent à des groupes dits "primitifs", dont
quelques-uns survivent encore aujourd'hui. Durant le Carbonifère, la plupart des formes qui sont aujourd'hui
contemporaines, apparaissent. Mais en dépit des similitudes d'apparence, la majorité de ces anciens
groupes, ne sont pas directement liés aux formes actuelles. Ce n'est qu'au cours du mésozoïque, que sont
apparu la plupart des gastéropodes que nous pouvont observer dans la nature.[réf. nécessaire]
Une radiation évolutive s'est produite au début du cénozoïque.

Les gastéropodes terrestres [modifier]

Les premiers Pulmonata, c'est-à-dire les gastéropodes terrestres, sont des escargots du genre
Maturipupa. Leurs fossiles ont été datés du Carbonifère. Ils ont été découverts en Europe. Les escargots
terrestres proches des escargots modernes sont rares avant le Crétacé, où le genre Helix est apparu. Les
limaces sont également issues de ce groupe.
Secondairement, certains d'entre eux sont (re)devenus aquatiques, colonisant les eaux douces
continentales (Ce sont essentiellement les lymnées et les planorbes).
Histoire de cette théorie [modifier]
La classification traditionnelle était basée sur des observations morphologiques. Une nouvelle
classification, basée sur les études génétiques, a été proposée en 1997 par Ponder & Lindberg. Une autre,
plus complète, l'a été par Bouchet & Rocroi en 2005.

[Enrouler]
v·d·m
Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation
 • Portail de l’origine et de l’évolution du vivant

• Portail de la zoologie
Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_gast
%C3%A9ropodes ».

Catégories : Gastéropode | Histoire évolutive | [+]

 w000

Histoire évolutive des primates

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Cet article ne cite pas suffisamment ses sources (juin 2008).
Si vous connaissez le thème traité, merci d'indiquer les passages à sourcer avec
{{Référence souhaitée}} ou, mieux, incluez les références utiles en les liant aux notes de
bas de page. (Modifier l'article)
Cet article est une ébauche concernant l'origine et évolution de la vie.
Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les
recommandations des projets correspondants.
 Alors que les mammifères subissent une radiation évolutive dès le début de l'ère tertiaire : de
nombreuses espèces différentes apparaissent et occupent des milieux variés : les airs, les milieux
aquatiques... Il se produit l'apparition des mammifères arboricoles : les primates. Parmi eux, un grand
nombre sont frugivores et c'est d'ailleurs parallèlement à l'acquisition de ce régime frugivore que devient
possible la vision en couleur, pour le repérage des fruits mûrs (les chiens ou les ruminants ont une
perception des couleurs bien inférieure). Parmi eux les singes (simiens) puis les grands singes, se seraient
diversifiés, ces derniers n'ayant pas de queue préhensiles.

Sommaire
[masquer]
• 1 Les singes de l'Ancien Monde (ou Catarrhiniens)
• 2 Les singes du Nouveau Monde (ou Platyrrhiniens)
• 3 Lignée humaine
• 4 Comparaison génétique entre humains et chimpanzés
• 4.1 Au niveau chromosomique
• 5 Notes et références
• 6 Voir aussi
• 6.1 Articles connexes
Les singes de l'Ancien Monde (ou Catarrhiniens) [modifier]

Les singes du Nouveau Monde (ou Platyrrhiniens) [modifier]

L’Amérique du Sud eut une position insulaire durant presque toute l’ère tertiaire. Une faune unique
s’y développa pendant plusieurs dizaines de millions d’années. À plusieurs reprises lors des périodes de
sécheresse, le bloc de la forêt humide se fragmenta en poches géographiques, créant des biotopes séparés
appelés « refuges », qui entrainèrent à leur tour une modification des préférences écologiques et un
isolement reproductif, deux facteurs-clés dans le développement de nouvelles espèces. Durant le tertiaire,
les seuls grands prédateurs d’Amérique du Sud étaient les borhyènes et les thylacosmilus, des marsupiaux
carnivores trop lents pour constituer un danger pour les primates.
Au début du pléistocène, lorsqu’une connexion fut rétablie entre les deux continents au niveau de Panama,
la faune nord-américaine déferla vers le sud où régnaient de meilleures conditions climatiques. Tous les
marsupiaux (hormis les opossums) furent évincés par les nouveaux arrivants. Des carnivores plus agressifs
firent leur apparition, comme le smilodon (« tigre » à dents de sabre) ou le cerdocyon (« renard » des
prairies, omnivore). Pourtant, aucun des prédateurs venus du nord ne put mettre en danger la moindre
espèce de primates. De nos jours, l’ocelot et le chat-tigre constituent de réelles menaces pour les petits
singes, mais ces derniers ripostent par la vélocité, la vigilance et une communication évoluée.
Clairement, en débarquant il y a au moins 30 millions d’années dans le Nouveau Monde, les singes ont
trouvé une terre d’accueil éminemment propice à leur épanouissement et à leur diversification. Les seuls
mammifères occupant des niches écologiques voisines sont les paresseux, les opossums, les porcs-épics et
les écureuils. De nos jours, 30 % des espèces de primates recensées vivent sur ce seul continent et d’autres
restent à découvrir dans les régions inexplorées d’Amazonie. L’on est aujourd’hui à peu près certains que les
singes ont colonisé l’Amérique du Sud depuis l’Afrique en dérivant sur des radeaux à une époque où de
nombreuses îles émergeaient encore de l’océan Atlantique. Tous les singes modernes sud-américains
descendraient d’un stock unique, mais cet ancêtre commun n’a pu encore être identifié. Les plus vieux
fossiles exhumés (espèces des genres Branisella et Dolichocebus) datent d’environ 25 millions d’années.

Lignée humaine [modifier]

Dans le groupe des primates, Homo sapiens fait partie[1] des :
• Haplorhiniens (plus de rhinarium, mais un nez), au même titre que les tarsiers ;
• Simiiformes (arrière des orbites crâniennes fermées), au même titre que les ouistitis ;
• Catarhiniens (narines dirigées vers le bas), au même titre que les colobes ou les babouins ;
• Hominoïdes (pas de queue), au même titre que les gibbons ou singes hurleurs ;
• Hominoïdés, au même titre que l'orang-outan ;
• Hominidés, au même titre que les gorilles ;
• Homininés, au même titre que les chimpanzés et bonobos ;
• Hominines, au même titre que les Australopithèques (éteints), Ardipithèques (éteints) et
Paranthropus (éteints) ;
• Le genre Homo, au même titre que l'homme de Néandertal ou l'homme de Flores.
 La classification phylogénétique permet aujourd'hui de positionner de façon précise la lignée
humaine par rapport aux huit genres non-éteints de singes qui forment la super-famille des Hominoïdes :
• le regroupement des diverses espèces d'hommes (genre Homo, à 46 chromosomes) avec
les diverses espèces d'australopithèques forme la tribu des Hominines,
• le regroupement de la tribu des Hominines avec celle des Panines (genre Pan, à 48
chromosomes) forme la sous-famille des Homininés,
• le regroupement de la sous-famille des Homininés avec celle des Gorillinés (genre Gorilla, à
48 chromosomes) forme la famille des Hominidés,
• le regroupement de la famille des Hominidés avec celles des Pongidés (famille maintenant
limitée au seul genre Pongo, à 48 chromosomes, comprenant l'espèce orang outan) [2] et des
Hylobatidés (quatre genres : le genre Hylobates, à 44 chromosomes, qui comprend les espèces de
gibbons, le genre Hoolock, à 38 chromosomes, le genre Nomascus, à 52 chromosomes, et le genre
Symphalangus, à 50 chromosomes) forme la super-famille des Hominoïdes ;
À un degré plus élevé dans la phylogénie :
• le regroupement de la super-famille des Hominoïdes avec celle des Cercopithécoïdes forme
le groupe zoologique des Catarhiniens (singes du Vieux-Monde),
• le regroupement des Catarhiniens avec les Platyrhiniens (singes du Nouveau-Monde) forme
l'infra-ordre des Simiiformes (singes),
• ensuite, le regroupement des Simiiformes avec les Tarsiformes forme le sous-ordre des
Haplorhiniens,
• le regroupement du sous-ordre des Haplorhiniens avec celui des Strepsirhiniens (qui
comprend les infra-ordres des Lorisiformes et des Lémuriformes) forme l'ordre des Primates,
 Comparaison génétique entre humains et chimpanzés [modifier]
La comparaison génétique entre humains et chimpanzés permet de démontrer l'étroite parenté entre
l'espèce humaine moderne actuelle et l'espèce la plus proche : les chimpanzés.

Au niveau chromosomique [modifier]

La séparation de la lignée humaine d'avec les autres genres de singes se serait faite
progressivement, la dernière en date ayant été celle d'avec le genre Pan (bifurcation des Homininés en
Hominines et Panines, par fusion des deux paires de chromosomes {2p, 2q} de l'ancêtre commun aux
humains, aux chimpanzés et aux bonobos en la paire de chromosomes {2} du genre Homo qui a par suite 46
chromosomes, le genre Pan ayant conservé les deux paires de chromosomes {2p, 2q} de l'ancêtre commun
et donc 48 chromosomes), il y a moins de 6,3 millions d'années selon des travaux récents menés sous la
direction de David Reich de la Harvard Medical School à Boston et publiés dans la revue Nature en mai
2006. Toutefois, ces travaux indiquent également que cette séparation ne semble pas avoir été brutale, car
la comparaison des chromosomes X de l'homme moderne et du chimpanzé moderne montre des similitudes
qui semblent refléter une longue "ré-hybridation" entre le genre Homo et le genre Pan ; par suite, bien qu'un
métissage entre l'homme moderne et le chimpanzé moderne soit impossible (en plus des chromosomes
sexuels {X, Y} qui sont très voisins, il subsiste 13 autres paires de chromosomes {3, 6, 7, 8, 10, 11, 12, 14,
16, 19, 20, 21, 22} qui semblent pratiquement identiques entre les deux espèces, ainsi que 6 paires qui sont
restées proches à la suite d'insertions {1}, d'inversions {4, 5, 17} et de délétions {13, 18} relativement simples
et facilement identifiables, mais deux paires de chromosomes {9, 15} ont été plus profondément différentiées
à la suite de mutations complexes probablement échelonnées dans le temps au sein des genres Homo et/ou
Pan), un métissage significatif entre au moins une espèce de chimpanzé et des espèces d'australopithèques
et probablement des espèces d'homminidés, conduisant à des échanges de gènes entre les deux genres
(98,8 % des gènes sont communs entre les hommes modernes et les chimpanzés modernes), a dû exister
pendant peut-être 4 millions d'années, durant (?)[réf. nécessaire]

Notes et références [modifier]

1. ↑ Classification phylogénétique de Guillaume Lecointre et Hervé Le Guyader, Belin
2. ↑ Le groupement de ces deux familles, qui possèdent cinq chromosomes {6, 19, 21, 22, X}
pratiquement identiques, étant parfois appelé Hominoïdés

Voir aussi [modifier]

Articles connexes [modifier]
• Histoire évolutive des mammifères
• Primates (classification phylogénétique)
• Liste détaillée des familles et genres de primates
• Liste alphabétique des genres de primates
• Liste alphabétique des noms vernaculaires de primates
• Orientation bibliographique en mammalogie

[Enrouler]
v·d·m
 Évolution biologique
[Dérouler]Mécanismes

Coévolution · Évolution insulaire · Évolution parallèle · Microévolution et

Évolution
macroévolution

Génétique
Dérive génétique · Mutation génétique · Recombinaison
évolutive

Sélection Pression de sélection · Sélection artificielle · Sélection de parentèle ·

naturelle Sélection écologique · Sélection sexuelle

Adaptation · Convergence évolutive · Exaptation · Extinction des

Spéciation
espèces · Isolement reproductif · Radiation évolutive

Théories de Équilibre ponctué · Saltationnisme · Théorie neutraliste · Théorie

l'évolution synthétique · Transmission des caractères acquis

[Dérouler]Branches évolutives

Histoire évolutive du vivant · Origines de la vie

Végétaux Histoire évolutive des végétaux

 Protostomiens Histoire évolutive des gastéropodes

Histoire évolutive des cétacés · Histoire évolutive des équidés · Histoire

évolutive des hippopotamidés · Histoire évolutive des homininés · Histoire
Mammifères
évolutive des mammifères · Histoire évolutive des primates · Histoire évolutive
des siréniens

Autre Histoire évolutive des poissons cartilagineux · Histoire évolutive des

Deutérostomiens dinosaures · Histoire évolutive des oiseaux

[Dérouler]Caractères adaptatifs

Histoire évolutive des dents · Évolution du flagelle · Histoire évolutive de l'oeil · Céphalisation

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Histoire_%C3%A9volutive_des_primates ».

Catégories : Histoire évolutive | Anthropologie physique et biologique | Zoologie | [+]

 w000

Évolution de la pluricellularité
Un article de Wikipédia, l'encyclopédie libre.

Aller à : Navigation, rechercher

Les organismes pluricellulaires sont les êtres vivants possédant plusieurs cellules, proches
spatiallement et interagissant fortement. Ils sont présents dans l'ensemble du monde vivant, et sont
particulièrement nombreux chez les Eucaryotes (Végétaux, Métazoaires, etc.). L'apparition des premiers
organismes pluricellulaires date d'au moins 2,1 milliards d'années, et pourrait remonter jusqu'à 3,5 milliards
d'années[1]. La pluricellularité est cependant apparue au moins 25 fois au cours de l'évolution[2], par des
mécanismes différents, probablement en raison des avantages sélectifs qu'elle confère, comme la possibilité
d'une augmentation de la taille de l'organisme ou d'une spécialisation des différentes cellules. La
pluricellularité entraîne cependant l'apparition de conflits génomiques et de lignées cellulaires égoïstes, telles
que les tumeurs. Ces conflits peuvent se résoudre par l'apparition de mécanismes de prévention contre ces
cellules tricheuses, ou encore par un retour à l'unicellularité.
Sommaire
[masquer]
• 1 La pluricellularité
• 1.1 Définition
• 1.2 Différents types
• 2 L'étude de l'évolution de la pluricellularité
• 2.1 Histoire
• 2.2 Données disponibles
• 2.2.1 Les fossiles pluricellulaires
• 2.2.2 Les organismes contemporains
pluricellulaires
• 2.2.3 Les organismes unicellulaires à
vestiges pluricellulaires
• 2.3 Méthodes d'études
• 2.3.1 Biologie comparative
• 2.3.2 Expérimentation
• 3 Les différentes apparitions de la pluricellularité
• 3.1 Chez les Eubactéries
• 3.2 Ches les Archées
• 3.3 Chez les Eucaryotes
• 4 Mécanismes évolutifs
La pluricellularité [modifier]
Définition [modifier]
Un organisme cellulaire est un ensemble de cellules dont l'activité est coordonnée, et qui sont en
contact physique, ou suffisamment proches spatialement pour interagir fortement[3]. Ainsi, une colonie
cellulaire ne peut être considéré comme un organisme pluricellulaire dans la mesure ou les cellules ne sont
pas ou peu coordinées entre elles.
Différents types [modifier]

L'étude de l'évolution de la pluricellularité [modifier]

Histoire [modifier]

Données disponibles [modifier]

Les fossiles pluricellulaires [modifier]

Les organismes contemporains pluricellulaires [modifier]

Les organismes unicellulaires à vestiges pluricellulaires [modifier]

Méthodes d'études [modifier]

Biologie comparative [modifier]

Expérimentation [modifier]
[4]
Les différentes apparitions de la pluricellularité [modifier]

Chez les Eubactéries [modifier]

La pluricellularité est apparue au moins trois fois de manières indépendantes chez les Eubactéries:
chez les Myxobacteria, chez les Actinobacteria et chez les Cyanobacteria[5]. La plupart des espèces de ces
trois groupes possèdent au moins une phase de vie pluricellulaire. Par ailleurs, certains auteurs considèrent
qu'un grand nombre de colonies bactériennes peuvent être considérées comme pluricellulaires, compte tenu
de l'importance de la communication entre les différentes cellules et la coordination de leurs actions[6].

Ches les Archées [modifier]

On trouve certaines espèces d'Archées, du genre Methanosarcina, formant des groupes de cellules
se liant entre elles après la division. Ces espèces, anaérobies obligatoires, pourraient bénéficier de la
pluricellularité en protégeant les cellules les plus au centre d'un trop fort taux de dioxygène dans le milieu
extérieur.

Chez les Eucaryotes [modifier]

Articles détaillés : Évolution de la pluricellularité chez les Métazoaires et Évolution de la
pluricellularité chez les Embryophytes.
La pluricellularité est apparue de nombreuses fois de manières indépendantes chez les Eucaryotes.
Si la pluricellularité des Métazoaires semble monophylétique, ce n'est probablement pas le cas de celle des
plantes ou des champignons. Ainsi, on trouve des organismes pluricellulaires chez les Opisthokonta
(Metazoa, certains Mycota et certains Choanomonada), chez les Amoebozoa, chez les Archaeplastida
(Embryophytes, certains autres Chlorophyta et certains Rhodophyta), chez les Stramenopila (certains
Oomycota, certains Phaeophyceae et de rares formes pluricellulaires chez les Bacillariophyta), chez certains
Excavata et chez les Alveolata (rares formes pluricellulaires chez les Ciliophora). On ne retrouve pas de
forme coloniale ou pluricellulaire chez les Rhizaria et les Excavata.

Mécanismes évolutifs [modifier]

[7]

Bénéfices [modifier]

Avantages liés à la taille [modifier]

Diminution du risque de prédation [modifier]

[8]

Spécialisation fonctionnelle [modifier]

[9]
 Nutrition plus efficace [modifier]

Dispersion plus efficace [modifier]

Diminution des interactions non coopératives [modifier]

Adaptations nécessaires à la pluricellularité [modifier]

Différenciation cellulaire [modifier]

Communication intercellulaire [modifier]

Adhesion intercellulaire [modifier]

Contraintes [modifier]

Mécanismes moléculaires et cellulaires [modifier]

Conséquences évolutives de la pluricellularité [modifier]

Conflits génomiques [modifier]

Complexification [modifier]

Retours à l'unicellularité [modifier]

• Mammifères:
• [10]
• [11]
• Bacteria:
• [12]

Notes et références [modifier]

1. ↑ Stéphane Foucart, « Paléontologie : une nouvelle histoire de la vie [archive] », Le Monde.
Mis en ligne le 2 jullet 2010, consulté le 3 juillet 2010
2. ↑ (en) Richard K. Grosberg et Richard R. Strathmann, « The evolution of multicellularity: a
minor major transition », dans Annual review of ecology evolution and systematics, vol. 38, 2007,
p. 621-654 (ISSN 1543-592X) [ texte intégral [archive], lien DOI [archive] ]
3. ↑ (en) Dale Kaiser, « Building a multicellular organism », dans Annual Review of Genetics,
vol. 35, 2001, p. 103-23 (ISSN 0066-4197) [ résumé [archive] ]
4. ↑ (en) Martin E. Boraas, Dianne B. Seale et Joseph E. Boxhorn, « Phagotrophy by a
flagellate selects for colonial prey: A possible origin of multicellularity », dans Evolutionary Ecology,
vol. 12, no 2, Février 1998, p. 153-164 (ISSN 0269-7653) [ résumé [archive], lien DOI [archive] ]
 5. ↑ (en) John Tyler Bonner, « The Origins of Multicellularity », dans Integrative biology,
vol. 1, no 1, 1998, p. 27-36 [ résumé [archive] ]
6. ↑ (en) James A. Shapiro, « Thinking about bacterial populations as multicellular
organisms », dans Annual review of microbiology, vol. 52, Octobre 1998, p. 81-104 (ISSN 0066-
4227) [ texte intégral [archive], lien DOI [archive] ]
7. ↑ (en) Antonis Rokas, « The Origins of multicellularity and the early history of the genetic
toolkit for animal development », dans Annual review of genetics, vol. 42, 2008, p. 235-251
(ISSN 0066-4197) [ texte intégral [archive], lien DOI [archive] ]
8. ↑ (en) Steven M. Stanley, « An Ecological Theory for the Sudden Origin of Multicellular
Life in the Late Precambrian », dans PNAS, vol. 70, no 5, Mai 1973, p. 1486-1489 (ISSN 0027-8424)
[ texte intégral [archive] ]
9. ↑ (en) Richard E. Michod, « Evolution of individuality during the transition from unicellular
to multicellular life », dans PNAS, vol. 104, no S1, 15 Mai 2007, p. 8613-8618 (ISSN 0027-8424)
[ texte intégral [archive], lien DOI [archive] ]
10.↑ (en) Claudio Murgia, Jonathan K. Pritchard, Su Yeon Kim, Ariberto Fassati et Robin A.
Weiss, « Clonal origin and evolution of a transmissible cancer », dans Cell, vol. 126, no 3, 11 Août
2006, p. 477-487 (ISSN 0092-8674) [ texte intégral [archive], lien DOI [archive] ]
11.↑ (en) A.-M. Pearse et K. Swift, « Allograft theory: Transmission of devil facial-tumour
disease : An uncanny similarity in the karyotype of these malignant tumours means that they could
be infective », dans Nature, vol. 439, no 7076, 2 Février 2006, p. 549 (ISSN 0028-0836)
[ résumé [archive], lien DOI [archive] ]
 12.↑ (en) Gregory J. Velicer, Lee Kroos et Richard E. Lenski, « Loss of social behaviors by
Myxococcus xanthus during evolution in an unstructured habitat », dans PNAS, vol. 95, no 21, 13
Octobre 1998, p. 12376-12380 (ISSN 0027-8424) [ texte intégral [archive] ]

Annexes [modifier]
Articles connexes [modifier]

Liens externes [modifier]

Bibliographie [modifier]
Ce document provient de « http://fr.wikipedia.org/wiki/%C3%89volution_de_la_pluricellularit
%C3%A9 ».

Catégorie : Histoire évolutive

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Explosion cambrienne
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L’explosion cambrienne (il y a entre 542 et 530 Ma) décrit l'apparition soudaine - à l'échelle
géologique - des organismes pluricellulaires.
Le nombre d'individus et leur diversité sont faibles, contrairement à la disparité[1], leur organisation
anatomique. Dans les années 1970 (Whittington), de nouveaux embranchements, inclassables dans la
classification classique, ont été mis au jour grâce à de nouvelles études de la faune de Burgess.
 Cette période est marquée par une nette transition dans le contenu fossilifère des couches
géologiques correspondantes, ainsi que par la formation de la plupart des grands embranchements actuels
de Métazoaires (animaux pluricellulaires)[2].
L'apparition de ces organismes a été qualifiée d'« explosive », d'une part en raison du changement
évolutif, en effet extrêmement rapide, d'autre part à cause des limites technologiques lors de leur
découverte. En effet, les microscopes d'alors ne permettaient pas d'analyser avec précision et de différencier
tous les micro-fossiles des périodes antérieures au Cambrien. L'amélioration des microscopes a peu à peu
révélé un éventail de micro-fossiles plus anciens.
La découverte en 1909 de la faune de Burgess révéla alors une quantité importante de fossiles
animaux alors inconnus, qui fut reliée à une période géologique désormais désuète, le « pré-cambrien ».
Des découvertes plus récentes de micro-fossiles ont montré que le « pré-cambrien » ne recélait pas
seulement des animaux unicellulaires (protistes et diploblastique). En 1994, des animaux triploblastiques
(constitués de plus de deux couches cellulaires) ont été découverts, préservés sous forme d'embryons
phosphatisés dans des roches du sud de la Chine[3]. Ces fossiles sont estimés avoir 570 millions d'années
et sont donc plus vieux que la faune de l'Édiacarien, trouvée dans des strates plus jeunes de 10 millions
d'années.
Sommaire
[masquer]
• 1 Fossiles [4]
• 2 Causes de l'explosion cambrienne
• 3 Voir aussi
• 3.1 Liens externes
• 4 Références

Fossiles [4] [modifier]

Cette période de l'histoire évolutive est à l'origine des fossiles parmi les plus étonnants jamais
découverts. Le seul site des schistes de Burgess a fourni une énorme quantité d'informations sur cette
période marquée d'importants changements évolutifs, pendant laquelle se sont formés la plupart des
embranchements animaux actuels. De nombreuses configurations cellulaires aujourd'hui disparues sont
également apparues à cette période, et restent aujourd'hui énigmatiques.
Avant l'explosion, les traces fossiles étaient dominées par des organismes unicellulaires. Seule la
faune d'Ediacara montre la présence d'animaux au corps mou, ainsi que certains micro-fossiles
pluricellulaires, attestant ainsi que la pluricellularité était déjà apparue 30 millions d'années plus tôt.
Lors de l'explosion cambrienne, on voit apparaître les premières coquilles ainsi que d'autres
structures morphologiques rigides. Les squelettes étant mieux préservés que les corps mous lors de la
sédimentation, les fossiles deviennent beaucoup plus abondants à cette période, ce qui facilite l'étude de la
faune post-cambrienne, et explique le changement drastique dans la composition fossilifère des couches
datant du Cambrien.

Causes de l'explosion cambrienne [modifier]

L'explosion cambrienne aurait pu être précipitée par une série de changements environnementaux
durant et avant cette période. Tout d'abord, la glaciation de Varangia a conduit à la glaciation de la plupart
des océans. Un réchauffement planétaire et une déglaciation extrêmement rapides suivent alors, précédant
la radiation évolutive.
Dans les environnements arctiques actuels, on trouve des organismes unicellulaires étendus sous
les couches de glace afin de maximiser leur exposition au soleil. Il est possible que des adaptations utiles à
la survie de colonies de ce type aient également contribué à la formation des premiers animaux
triploblastiques, il y a 570 millions d'années. De plus, l'environnement glacial de la terre n'aurait permis le
développement que de quelques niches écologiques, et la libération de nombreuses niches lors de la
déglaciation explique la diversification extrêmement rapide des animaux du Cambrien.

Voir aussi [modifier]

Liens externes [modifier]
• Eric F.-Bellot, Le Schiste de Burgess , 15/06/2002
• L’explosion cambrienne: Une collection de monstres marins
Références [modifier]
1. ↑ la disparité caractérise le nombre de plans anatomiques différents
2. ↑ D.Y.C. Wang, S. Kumar & S. B. Hedges, « "Divergence time estimates for the early history
of animal phyla and the origin of plants, animals and fungi." [archive] », dans Proceedings of the
Royal Society of London, Series B, Biological Sciences , vol. 266, 1999, p. 163-171
3. ↑ S. Xiao, Y. Zang & A. Knoll, « "Three-dimensional preservation of algae and animal
embryos in a Neoproterozoic phosphorite." [archive] », dans Nature, vol. 391, 1998, p. 553-558
4. ↑ A.G. Collins, « Metazoa: Fossil record [archive] ». Consulté le 14 Décembre 2005

• Portail de l’origine et de l’évolution du vivant

Ce document provient de « http://fr.wikipedia.org/wiki/Explosion_cambrienne ».

Catégories : Paléozoïque | Explosion radiative | Histoire évolutive | [+]

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