FOUNDATION BIOLOGY 1

BIO091

CHAPTER 5
ENERGY
PHOTOSYNTHESIS
1

CONTENTS
Chap 5: Energy
5.1 : Autotrophic and heterotrophic nutrition

5.2 : Photosynthesis

5.2.1 : Absorption spectrum of photosynthetic pigments

5.2.2 : Light reaction
5.2.3 : Dark reaction
5.2.4 : C3, C4 and CAM plant
2

LEARNING OUTCOMES
1.

Differentiate between autotrophy and heterotrophy based on their energy and

carbon sources.
2.

Discuss the relationship between an action spectrum and an absorption

spectrum

3.

Describe the photosynthetic pigments including the structure of a chloroplast.

4.

Explain the photoactivation of chlorophyll that converts light energy into ATP
and NADP+.

5.

Trace the movement of electrons in linear electron flow and cyclic electron

flow.

LEARNING OUTCOMES
5.

Describe Calvin cycle that involves three stages.

6.

Discuss the role of ATP and NADPH.

7.

Explain carbon fixation in C3, C4 and CAM plants.

8.

Discuss the major consequences of photorespiration.

9.

Describe two important photosynthetic adaptations that minimize

photorespiration.
4

INTRODUCTION

Life on earth is solar powered.

The chloroplast in plants & other photosynthetic organisms capture light

energy from the sun & convert it to chemical energy.

This chemical energy is stored in sugar & other organic molecules.

This conversion process is called photosynthesis.

INTRODUCTION

Energy flows into an ecosystem as

sunlight and leaves as heat.

Photosynthesis,
generates O2 & organic

molecules, which are used in

cellular respiration.

Cells use chemical energy stored in

organic molecules,
to generate ATP during cellular
respiration.
ATP is used to power cellular
works.

INTRODUCTION

Photosynthesis is a biochemical process that is very important to life.

Nearly all life depends on it for organic nutrients

An organism acquires organic compounds (used for energy & carbon

skeleton) by 1 of 2 major modes:
1. Autotrophic nutrition
2. Heterotrophic nutrition
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AUTOTROPHS

(auto = self ; trophos = feed)

are self-feeders

They sustain themselves without eating anything derived from other

organisms.

Are producers of biosphere,

producing organic molecules from CO2 & other inorganic raw

materials.

These organic molecules produced are the source of organic

compounds for all non-autotrophic organisms.

AUTOTROPHS

Almost all plants (green plants) are photoautotrophs

using energy from sunlight to synthesize organic molecules from
H2O & CO2.

Photosynthesis also occurs in

Algae,
Certain unicellular eukaryotes,
Some prokaryotes.

Photoautotrophs

Multicellular algae

Terrestrial plants

predominant producers of food.

Photoautotrophs
Some unicellular
protists (unicellular
eukaryotes) Euglena

Photosynthetic
prokaryotes purple
sulphur bacteria
Cyanobacteria

(prokaryotes)

HETEROTROPHS
(hetero = other)

Unable to make their own food,

.: obtain their organic materials produced by other organisms.

They are the biospheres consumers

All most all heterotrophs, including humans, depend on

photoautotrophs for food & O2.

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HETEROTROPHS
Decomposers:

are heterotrophs

consumes the remains of dead organisms by decomposing organic

litter:

carcasses

feces

fallen leaves

Most fungi & many types of prokaryotes are decomposers.

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PHOTOSYNTHESIS
Definition of Photosynthesis:

Photosynthesis = The conversion of light energy to chemical energy that is

stored in sugars or other organic compounds.

(Source: Campbell 10th edition)

The simplified equation for Photosynthesis:

Photosynthesis is the process where - autotrophic organisms use light energy

to make organic molecules & oxygen gas from carbon dioxide & water.

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PHOTOSYNTHESIS

The process of photosynthesis most likely originated in a group of bacteria that had

infolded regions of the plasma membrane.

In existing photosynthetic bacteria, infolded photosynthetic membranes function

similarly to the internal membrane of chloroplast (a eukaryotic organelle).

According to the Endosymbiont theory,

the original chloroplast was believed to have been a photosynthetic
prokaryote that lived inside an ancestor of eukaryotic cells.

Chloroplast are present in a variety of photosynthesizing organisms.

15

PHOTOSYNTHESIS

Prochloron, a photosynthetic bacterium have infoldings of the cell

membrane that form thylakoids inside the cell's cytoplasm
16

The Importance of Photosynthesis

The energy entering chloroplasts as sunlight gets stored as chemical

energy in organic compounds.

Sugar made in the chloroplasts supplies chemical energy and carbon skeletons to

synthesize the organic molecules of cells.

Produces the building blocks for complex compounds in plants (e.g. cellulose & lignin).

Provides the energy needed for chemical changes in the form of ATP.

Plants store excess sugar as starch in structures such as roots, tubers, seeds, and fruits

Provides food to primary consumers (directly) & secondary (& higher) consumers
(indirectly).
17

Helps reduce excess CO2 & increase O2 concentrations in the air.

PHOTOSYNTHESIS

18

The Site of Photosynthesis:

THE LEAF
The leaves are the major site of photosynthesis for most plants.
It is adapted to:
receive energy from the sun
allow diffusion of gasses & water vapor
remove the products of photosynthesis.
The leaves have large surface area & are held to
receive maximum illumination

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The Site of Photosynthesis:

THE LEAF

Chloroplasts are found mainly in the

cells of the Mesophyll tissue

in the interior of the leaf.

A typical mesophyll cell has 3040

chloroplasts.

Stomata (microscopic pores in the

epidermis) allows CO2 enters & O2
exits the leaf. ( gas exchange )

Veins
deliver water from roots to the leaves.

carry off sugar from the mesophyll cells to non-photosynthetic areas of the plant.

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The Site of Photosynthesis:

CHLOROPLASTS

Photosynthesis in eukaryotes takes place in

chloroplast.

Chloroplasts have important role absorbtion of

light energy during photosynthesis.

A chloroplast has an envelope of TWO

membranes surrounding a dense fluid called

stroma.

Chloroplast contains a system of interconnecting

membranous flattened sacs called thylakoids

Thylakoid sacs are stacked in columns called

grana. (singular = granum)

Thylakoid membrane encloses a fluid-filled interior

space called thylakoid space.

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The Site of Photosynthesis:

CHLOROPLASTS

Chlorophyll is the green pigments that gives leaves their

green color.

Thylakoid membranes & grana increase the surface area for attachment of
chlorophyll molecules,
accessory molecules &

involved in the light reactions.

electron carriers

Light energy absorbed by chlorophyll drives the synthesis of organic

molecules in the chloroplast.

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23

Photosynthesis: Chemical Equation

The overall chemical reactions in photosynthesis is summarized as:

Chloroplasts split H2O into hydrogen and oxygen, incorporating the

electrons of hydrogen into sugar molecules

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Photosynthesis is a Redox Reaction

Photosynthesis involve redox reactions.

Water is split, and electrons are transferred along with

hydrogen ions from water to carbon dioxide, reducing CO2 to sugar.

Photosynthesis is endergonic (require energy):

Light energy is captured by chlorophyll molecules to excites electrons

Light energy is converted to chemical energy, &

Chemical energy is stored in the chemical bonds of sugars .

25

The Nature of Sunlight

Light is a form of electromagnetic energy = electromagnetic radiation.

Electromagnetic energy travels in rhythmic waves.

Wavelength,
is the distance between the crests of electromagnetic waves.

ranges from less than a nanometer (gamma rays) to more than a kilometer
(radio waves)

26

The Electromagnatic Spectrum

Electromagnetic spectrum:
is the entire range of radiation (electromagnetic energy)
The sun radiates the full spectrum of electromagnetic
energy,
but the atmosphere allows
only visible light to pass through &
screen out a substantial amount of other radiation.
27

Visible light
Visible light:

Consists of wavelengths from the electromagnetic

spectrum that is most important to life.

Ranges from 380nm to 750nm in

wavelength.

Can be detected as various colors by the human

eye.

Drives photosynthesis.

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The Electromagnetic Spectrum

White light is a mixture of all wavelength of visible light.

29

Energy Vs. Wavelength

Light behaves as though it consists of discrete particles called
photons

Each photon has a fixed quantity of energy.

The amount of energy is inversely related to the wavelength of the light.
the shorter the wavelength, the greater the energy of each
photon of that light.
E.g.: a photon of a violet light (380 nm) has ~twice as much
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energy as a photon of red light (750 nm).

Photosynthetic Pigments:
The Light Receptors

When lights meets matter, it may reflected, transmitted or absorbed.

Pigments are substances that absorb visible light.

Chloroplast pigments:
Chlorophyll a the main photosynthetic pigment,
Chlorophyll b accessory pigment
Carotenoids a group of accessory pigments.

broaden the spectrum

used for photosynthesis.

Different pigments absorb light of different wavelengths.

Wavelength that are not absorbed are reflected or transmitted

If a pigment absorbs all wavelengths it appears black.

31

Photosynthetic Pigments:
The Light Receptors
Leaves appear green because
chlorophyll,
absorbs violet-blue & red

light (color most effective on

driving photosynthesis)
&
reflects & transmits green
light.

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Photosynthetic Pigments:
Spectrophotometer
Spectrophotometer:
an instrument that can
measure the ability of a

pigment to absorb various

wavelengths of light.

The machine directs beam

of light of different
wavelength through a

solution of the pigments

and measures the fraction

of the light transmitted at

each wavelength.

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Absorption Spectrum of Photosynthesis

Absorption spectrum:

The graph obtained when a pigments light absorption is

plotted against wavelength.

Light can perform work in chloroplasts only if it is absorbed.

Therefore, the absorption spectra of chloroplast pigments,

tells the effectiveness of different wavelengths for driving
photosynthesis

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Absorption Spectrum of Photosynthesis

The 3 curves show the wavelengths

of light best absorbed by three types

of chloroplast pigments.

The spectrum of chlorophyll a,

The absorption of violet blue &

red is maximum.
Green is the least effective

color.

.: the wavelength that are optimally

absorbed by green plant as a source

of energy are:
Violet-blue & red light. 35

Action Spectrum
Action spectrum

The graph obtained when the

rate of photosynthesis is plotted

against wavelength.

It shows that photosynthesis is:

most efficient in blue-

violet light & red light, &

least efficient in
green light.

36

Action Spectrum Vs. Absorption

Spectrum

In comparing between the 2

spectrum:
The action spectrum for
photosynthesis does not

exactly match the absorption

spectrum for chlorophyll a.

The action spectrum for

photosynthesis is much

broader than absorption

spectrum of chlorophyll a.

This is due to the presence of

37

accessory pigments.

Accessory Pigments
Accessory pigments:
are photosynthetically important in chloroplasts.
have different absorption spectra e.g.
carotenoids &
chlorophyll b

.: accessory pigments broaden the spectrum of

colors that can be used for photosynthesis
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Accessory Pigment: Carotenoids

Carotenoids:

are hydrocarbons.

have various shades of yellow & orange,

because they absorb violet & blue-green light, &
reflect yellow & orange light.

broaden the spectrum of colors that can drive photosynthesis.

Important function of carotenoid photoprotection

absorb & dissipate excessive light energy that would damage chlorophyll.
have antioxidant properties that are good for human health benefits

39

Structure Of Chlorophyll Molecule

Porphyrin ring: Light-absorbing head of molecule

Magnesium atom at the center.

A slight structural difference

between the 2 chlorophylls,
cause them to absorb at

slightly different wavelengths.

Hydrocarbon tail:
Interact with hydrophobic
regions of proteins inside

.: under visible light:

Chlorophyll a appears bluegreen.

thylakoid membranes of
chloroplast.

Chlorophyll b appear olive

green (it absorbs blue 40
&
orange; reflect yellow-green)

Excitation of Chlorophyll by Light

A pigment molecule is said to be in its

ground state, if its electron is in its

normal orbital.

.: absorption of a photon of light

causes a transition of the chlorophyll

molecule from its ground state to its

excited state.

The photon boosts an electron (of the

chlorophyll) to an orbital where its has

more potential energy.

The excited state is unstable

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Excitation of Chlorophyll by Light

The excited electrons drop

back down to the ground-

state orbital, releasing

excess energy as:
i. Heat This
converts light energy to

heat.
ii. afterglow called
fluorescence this is

due to photons that are

given off.

42

Excitation of Chlorophyll by Light

.: an isolated solution of chlorophyll that

is excited (i.e. illuminated) with

ultraviolet-light will:
fluoresces with a red-orange glow
(i.e. release photon) &

release heat.

43

A Preview of Photosynthesis Stage

The 2 stages of photosynthesis:

1.

2.

The LIGHT REACTIONS (Light Dependent Reactions)

The photo part of photosynthesis.

Takes place in the thylakoid membranes

Solar energy is converted to chemical energy.

The CALVIN CYCLE (Light Independent Reactions)

The synthesis part of photosynthesis.

Takes place in the stroma of chloroplast.

The chemical energy is used to make organic molecules of food.

44

A Preview of Photosynthesis Stage:

The Light Reactions
During the Light Reaction:

Solar energy is converted to chemical energy

Water is split to provide the source of:

Electrons and protons(H+ ions) that reduce NADP+ to NADPH.

NADP+ (Nicotinamide adenine dinucleotide phosphate) is
an electron acceptor that accept a pair of electrons and
an H+ ions from water.

Release O2 as by-product.

Generate ATP through photophosphorylation,

i.e. the addition of phosphate group to ADP during chemiosmosis.

45

A Preview of Photosynthesis Stage:

The Light Reactions
Therefore the chemical energy produced during the Light
Reactions are:
NADPH(Nicotinamide adenine dinucleotide phosphate)
a source of electrons
ii. ATP the energy currency of
i.

cell.
NADPH & ATP produced will be used during the Calvin
Cycle.
46

A Preview of Photosyntheis Stage:

The Calvin Cycle

Also known as the Dark Reaction or The Light Independent Reaction.

because it does not require light energy directly.

The cycle:

makes sugar (carbohydrate) from CO2 using ATP and NADPH

begins with carbon fixation incorporating CO2 into organic molecules in

the chloroplast.

occurs during daylight because the cycle

47

requires NADPH & ATP that was produced during the Light Reaction.

An Overview of Photosyntheis
The thylakoid membrane:
-

The sites of the Light

Reaction.

The stroma:
-

The site of the Calvin cycle

The Light Reaction:

-

use solar energy to make

ATP & NADPH, & supply

them to the Calvin cycle.

The Calvin Cycle:

- Incorporates CO2 into organic
molecules & converted48
it to
sugar.

1. LIGHT DEPENDENT
REACTION

49

PHOTOSYSTEM
In Light Dependent Reaction
Light reaction involves PHOTOSYSTEM, which is a cluster
of chlorophyll molecules, small organic molecules and
proteins that are organized and embedded in the

thylakoid membrane.
A photosystem is composed of (i) a reaction-center
complex surrounded by several (ii) light-harvesting
complexes.

50

PHOTOSYSTEM
In Light Dependent Reaction

i.

A reaction-center complex:

Is a type of protein complex.

Holds a special pair of chlorophyll a molecules.

Also contain primary electron acceptor

51

PHOTOSYSTEM
In Light Dependent Reaction

ii.

Light-harvesting complexes:

Each complex consists of various pigment molecules(chlorophyll a, chlorophyll b &

carotenoids) bound to proteins.

These variety of pigment molecules allows a photosystem to harvest light energy over:
1.

a larger surface area

2.

a larger portion of spectrum this increases the range of wavelengths for52

plants to

obtain light energy.

PHOTOSYSTEM
In Light Dependent Reaction

Within a light-harvesting complex,

a pigment molecule absorbs a photon and the energy is transferred from

pigment molecule to pigment molecule until it reaches chlorophyll a
molecules in the reaction center complex

In the reaction-center complex, the pair of chlorophyll a molecules use the

light energy (photons) to boost one electrons to a higher energy

level and transfer the excited electron to the primary electron acceptor.
The primary electron acceptor accepted the electron & become reduced

this is redox reaction.

This is the first step of the light reaction.
This process is called photoactivation of chlorophyll.

53

PHOTOSYSTEM
In Light Dependent Reaction
a

54

PHOTOSYSTEM
In Light Dependent Reaction

Two types of photosystems in the thylakoid membrane, each has a

characteristic reaction-center complex:

1. Photosystem II (PS II)

has chlorophyll a at reaction-center complex called P680

P680 best at absorbing a wavelength of 680 nm

2. Photosystem I (PS I)

has chlorophyll a at reaction-center complex called P700

P700 best at absorbing a wavelength of 700 nm

** The number of the 2 photosystems reflect the order of their discover.

** PS II function first in the light reaction.

55

Routes for Electron Flow in the Light

Dependent Reaction

During the light reactions, there are 2 possible routes for electrons flow:

1. Linear electron flow

also known as non-cyclic photophosphorylation

Involves PS I & PS II

Produces both ATP & NADPH

2. Cyclic electron flow

also known as cyclic photophosphorylation

Involves PS I

Only generates ATP

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Linear Electron Flow

1.

A photon hits a pigment (in the light-

harvesting complex) of PS II.

The energy is transferred from one

pigment molecule to another until it

reaches the P680 pair of chlorophyll a

molecules (in the PS II reaction-center

complex).

The P680 becomes photoactivated

and excites an electron to a higher

energy level.
2.

The excited electron from P680 is

transferred to the primary electron
acceptor.

57

Linear Electron Flow

3.

P680+ (P680 that is missing an electron) is a very

strong oxidizing agent; its electron hole must be
filled.

A H2O molecule is split by an enzyme into:

2 Electrons

2 Hydrogen ions, H+

1 Oxygen atom

The electron are transferred one by one to the P680+,

thus reducing it to P680.

the H+ are released into the thylakoid space.

The oxygen atom immediately combines with an

oxygen atom (from another splitting of water)
58

forming O2 as by-product.

Linear Electron Flow

4.

Each photoexcited electron passes

from the primary electron acceptor of
PS II to PS I via an electron

transport chain (ETC).

5.

The exergonic fall of electrons to a

lower level provides energy for the

chemiosmosis synthesis of ATP.

The energy released is used to

pumped H+ from
stroma into the thylakoid space.

This creates a proton gradient

59
across the thylakoid membrane.

Electron Transport Chain Couples with

Chemiosmosis
Electron falls down the electron
transport chain couples with

chemiosmosis to synthesize ATP.

Free energy drops as electron

falls down the chain and energy is

released.

The energy creates a photon

gradient across the thylakoid
membrane.
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Electron Transport Chain Couples with

Chemiosmosis

The energy transfers (pumps) the H+ from the

stroma into the thylakoid membrane

The H+ build up inside the thylakoid space.

The H+ then diffuse down the concentration

gradient from the thylakoid space into the stroma

through an enzyme ATP synthase.

The enzyme catalyzes the reaction where ADP

and inorganic phosphate (Pi) produce ATP.
**phosphorylation of ADP to ATP

This process is called chemiosmosis

61

Linear Electron Flow

6.

Meanwhile, the PS I reaction-center complex receives the light energy.

The light energy excites an electron of the P700 pair of chlorophyll a in the
reaction-center.

The photoexcited electron is transferred to PS IS primary electron acceptor.

P700 becomes P700+ and act as electron acceptor.

P700+ accepts an electron that reaches the bottom of ETC from PS II.

62

Routes for Electron Flow in the Light

Dependent Reaction
a

7. Each electron falls down an ETC from the primary

electron acceptor of PS I through protein ferredoxin (Fd).

This chain does not create a proton gradient, and thus no ATP
produced.

63

Routes for Electron Flow in the Light

Dependent Reaction
a

8. The enzyme NADP+ reductase catalyzes the transfer the electrons from

Fd to NADP+.
2 electrons are required for the reduction of NADP + to NADPH.
This process also uses H+ from stroma

64

The electrons of NADPH are available for the reactions of Calvin Cycle

Cyclic Electron Flow

Cyclic Electron flow:

is an alternative path that uses Photosytem I only.

Also know as cyclic photophosphorylation.
It is a short circuit:

the electron cycle back from ferredoxin (Fd) to the cytochrome complex and
return to P700 in the PS I reaction-center complex.

Produces ATP, but not NADPH and no ATP released.

Benefits:
Cyclic electron flow generates surplus ATP (via chemiosmosis), satisfying the

higher demand in the Calvin cycle

Generate extra ATP for C4 plants (in regions with intense light) as an adaptation

of photorespiration . The bundle sheath cell carry out cyclic electron flow to
generate ATP (will be explain more of C4 plants later)
The cyclic electron flow may also be photoprotective.

65

Cyclic Electron Flow

66

Comparison of Liner Electron Flow Vs. Cyclic

Electron Flow
LINEAR ELECTRON FLOW

CYCLIC ELECTRON FLOW

(Non-cyclic photophosphorylation)

(Cyclic photophosphorylation)

Involved PS I & PS II

Involve PS I only

Electron flow is linear (non-cyclic)

Electron flow is cyclic

Photolysis of water occurs

No photolysis of water

Water is the first electron donor

Electron starts from PS I (P700)

Last electron acceptor is NADP+

Electron is transferred back to PS I

(P700)

Produce ATP, NADPH & O2

Produce ATP

- The ATP & NADPH produced are used in the light independent reaction in the
stroma.
67
- O2 is released as a by products.

Overall Process of
Light Dependent Reaction
a

68

2. CALVIN CYCLE
(LIGHT INDEPENDENT
REACTION)
69

The Calvin Cycle

The Calvin cycle, like the citric acid cycle, regenerates its

starting material after molecules enter and leave the cycle.

The Calvin cycle:
carried out in the Stroma
is anabolic building carbohydrates from smaller molecules
and consuming energy.
Use ATP as energy source and NADPH as the reducing power
of high-energy electrons.
70

The Calvin Cycle

Carbon enters the cycle as CO2 and leaves as a sugar named glyceraldehyde-3-

phospate (G3P)
The carbohydrates produced directly from Calvin cycle is actually not glucose,
but a three carbon sugar (G3P)

For net synthesis of 1 G3P molecule,

the cycle must take place 3 times, fixing 3CO2 molecules.

(one per turn of the cycle)

The 3 phases of Calvin Cycle:

1. Carbon Fixation (catalyzed by Rubisco)
2. Reduction

71

Phase 1: Carbon Fixation

The Calvin cycle in corporates each CO2 molecule,one at a time by

attaching it to a ribulose bisphosphate

(RuBP).
RuBP is a 5 carbon sugar & acceptor of CO2.

The reaction is catalyzed by Rubisco / RuBP carboxylase-oxygenase

The product of this reaction is an unstable six carbon intermediate,

which immediately split into two molecules of 3- phosphoglycerate (three
carbon each).

The carbon that was originally part of CO2 molecule is now part of a carbon
72
skeleton ; the carbon has been fixed.

Phase 1: Carbon Fixation

73

Phase 2: Reduction

Each molecule of 3-phosphoglycerate received an additional phosphate group from ATP to

become:
1,3-biphosphoglycerate.

1,3-biphosphoglycerate:
is reduced by a pair of electrons donated from NADPH & it loses a phosphate group &
become:

Glyceraldehyde-3-phosphate (G3P) - a 3C sugar - stores more potential energy.

For every 3 molecules of CO2 that enters the cycle, 6 molecules of G3P are formed.

1 of these 6 molecules of G3P, is counted as a net gain of carbohydrate,

& exits the cycle to be used by the plant to make organic compounds such as glucose.
74

Phase 2: Reduction

75

Phase 3: Regeneration of the CO2

acceptor (RuBP)
In a complex series of reactions,
the carbon skeleton of 5 molecules of G3P (3C sugar) are
rearranged into 3 molecules of RuBP (5C
sugar).

To accomplish this, the cycle spends 3 molecules of ATP.

The RuBP is now ready to receive CO2 again & the cycle
continues.

76

77

The Calvin Cycle

For the net synthesis of 1 G3P molecule,the Calvin cycle consumes:

3 molecules of CO2
3 times cycle
9 molecules of ATP

Produced during light reaction

6 molecules of NADPH

The G3P from the Calvin cycle becomes the starting material for metabolic
pathways that synthesize other organic compounds.

including glucose (formed by combining 2 G3P) & other carbohydrates.

78

79

80

Alternative Mechanisms of
Carbon Fixation in Hot and
Arid Climates
81

Photorespiration

Plants have special adaptation to deal with the problem of dehydration through
metabolic adaptations.

Plants have to compromise between photosynthesis & the prevention of

excessive water loss from the plant.

On a hot, dry day most plants close their stomata, to:

reduce transpiration (i.e. the evaporative loss of water from leaves) &
conserves water within the plant.

This response reduces photosynthetic yield, by limiting access to CO2.

The CO2 concentration begins to decrease in the air spaces within the leaf, &
The O2 concentration released form the light reactions begins to increase

This conditions within the leaf leads to: PHOTORESPIRATION a wasteful 82

process
for plants.

83

C3 Plants and Photorespiration

C3 plants are plants that, initially fix carbon in the Calvin cycle by

adding CO2 to RuBP with the help of Rubisco

This initial fixation of carbon occurs in most plants.

These plants are called C3 plants because the 1st organic product of
carbon fixation is a 3 carbon compound (3-phosphoglycerate).

E.g.: Rice, Wheat, and Soybean important in

84

agriculture.

C3 Plants and Photorespiration

On hot dry days, C3 plants close partially their stomata:

producing less sugar due to the declining level of CO2 in the leaf that
starves the Calvin cycle.

Rubisco can bind O2 in place of CO2.

As CO2 become scarce within the air spaces of the leaf & O2 increase,
Rubisco adds O2 in the Calvin cycle instead of CO2.
The product splits & a 2 carbon compound that leaves the chloroplast.

Mitochondria & peroxisomes breaks the 2 carbon molecules & release CO2.
This process is called Photorespiration

85

C3 Plants and Photorespiration

Photorespiration:
occurs in the light (photo),
consumes O2 & producing CO2 (respiration),
generates no ATP , but it consumes ATP
(this differs photorespiration from the normal cellular respiration).

produces no sugar & decreases photosynthetic output.

drains away as much as 50% of carbon fixed by the Calvin cycle.
this is a problem in many plants.
86

C3 Plants and Photorespiration

Photorespiration is naturally viewed as wasteful, because

it makes plants produces less foodespecially for
heterotrophs.

If photorespiration could be reduced in certain plants species,

crop yields & food supplies might increase.

In some plants species that lives in hot & arid climates,

they have evolved an alternate modes of carbon fixation that,
minimize photorespiration &
optimize the Calvin cycle.

87

Adaptation to Photorespiration
The 2 most important photosynthetic

adaptations to photorespiration:
1. C4 photosynthesis
2. CAM

88

C4 PLANTS
C4 plants are so named because they preface the Calvin
cycle with an alternate mode of carbon fixation,
that forms a four carbon compunds as its
first product.

Among the C4 plants important to agriculture:

sugarcane and corn, members of the grass family.
89

C4 PLANTS
In the anatomy of C4 plants leaves, there are 2
types of photosynthetic cells:
1.

Bundle-sheath cells

are arranged into tightly packed sheaths

around the veins of the leaf.
Calvin Cycle occurs here.

2.

Mesophyll cells
are arranged more closely between the
bundle sheath & the leaf surface.
Where CO2 is incorporated into PEP

prior to the Calvin cycle.

Kranz anatomy: is the arrangement pattern of these 2 chloroplast-rich
layers around vascular bundle.

90

The C4 Pathway
1.

In mesophyll cell,

PEP carboxylase adds CO2 to

phosphoenolpyruvate (PEP) A CO2

acceptor forming oxaloacetate (a 4

carbon product).
PEP carboxylase:

Enzyme that present only in mesophyll cell.

Has a higher affinity for CO2 than does
Rubisco.
Has no affinity for O2.

Can fix carbon efficiently when Rubisco

cannot (that is when CO2 concentration are
91

low and O2 increase)

The C4 Pathway

2. Oxaloacetate is converted to
Malate (4C).

Mesophyll cells export malate to

Bundle-sheath cells through
plasmodesmata.
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The C4 Pathway
3.

In the bundle-sheath cells, malate will release

CO2 & regenerates pyruvate

i.

CO2 regenerated:

increases the CO2 concentration in the

stroma of chloroplast.
enters into the normal Calvin cycle to
undergo fixation by rubisco, producing
organic molecules (G3P).

ii.

Pyruvate regenerated is transported back to

mesophyll cells
In the mesophyll cell, ATP is used to

convert pyruvate to PEP, allowing

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the reaction cycle to continues.

The C4 Pathway
Notes:

To generate these
extra ATP, bundle-

sheath cell carry

out cyclic electron
tranfer

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Summary:
Adaptation of C4 Plants

Physiological adaptation of C4 plants is advantageous to carry out photosynthesis in

hot regions with intense sunlight.

Where stomata partially closed during the day but decrease the rate of CO2
intake.

The physiological adaptation of C4 plants allows these plants to have mesophyll cells to:
i.

absorbs CO2 efficiently by having enzyme PEP Carboxylase ; has a higher affinity
for CO2 .

ii.

then pump the CO2 into bundle-sheath cells

keeping the CO2 concentration in these cells high enough for Rubisco to bind
CO2 rather than O2.

This is considered as a CO2concentrating pump that is powered by ATP.

.: C4 photosynthesis spends ATP energy to minimize photorespiration and enhance

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sugar production.

CRASSULACEAN ACID METABOLISM

(CAM) PATHWAY

A second photosynthetic adaptation to arid conditions has evolved in succulent

(water-storing) plants, many cacti, pineapple & other several families.

The mode of carbon fixation is called crassulacean acid metabolism, or
CAM

Named after the plant family Crassulaceae in which the process was 1st
discovered.
These plants are called CAM plants

CAM plants open their stomata during the night & close them during the
day

Closing stomata during the day helps desert plants conserve water, but it also
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prevents CO2 from entering the leaves

CRASSULACEAN ACID METABOLISM

(CAM) PATHWAY

During the night, when stomata are open,

these plants take up CO2 & incorporate it into a variety of organic acids.

The mesophyll cells of CAM plants store the organic

acids they make during the night in their vacuoles until the morning, when the
stomata close.

During the day (stomata closed), when the light reactions can
supply ATP & NADPH for the Calvin cycle,
CO2 is released from the organic acids made the night before, to become
incorporated into sugar (during Calvin cycle) in the chloroplasts.

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CRASSULACEAN ACID METABOLISM

(CAM) PATHWAY

Night Stomata open: CO2 fixed as malic acid &

stored as malate in vacuole.

Day Stomata closed : malic acid from vacuole

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converted to CO2 for starch synthesis.

C4 & CAM photosynthesis compared

The CAM pathway is similar to the C4 pathway,

in that CO2 is 1st incorporated into organic intermediate before it enters the Calvin
cycle

The difference is that:

1. In C4 plants,
the initial steps of carbon fixation are separated structurally from the Calvin

cycle (mesophyll cells, bundle-sheath cells)

2. In CAM plants,
the 2 steps occur at separate times (night, day), but within the same cell
(mesophyll cells).

Note: CAM, C4 & C3 plants eventually use Calvin cycle to make sugar from CO2. 99

C4 & CAM photosynthesis compared

100
The C4 & CAM pathways are evolutionary solutions to the problem of maintaining photosynthesis
with stomata closed on hot, dry days.

THE END

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