Professional Documents
Culture Documents
ABCs
Human Brain
Physiology Guide
Understand
•Nerve impulses
•Touch & Pain
•Hearing & Vision
•Movement
•Speech & EEG
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Contents of this book include revisions of Human Brain Function: An Introduction to Clinical Neurophysiology by Robert Lavine , 1996, ISBN
0536598223. This book was previously published by: Pearson Education, Inc.
EBOOK TABLE OF CONTENTS
(Supplement to Contents with page numbers from print edition)
Preface
Contents
1. Structure and Function
Motoneurons and Nerve Photos
Neurons in a Spinal Reflex Diagram
Brain Connections with Spinal Neurons Diagram
Giant Axons from the Giant Squid
2. The Nerve Impulse
Voltage-Gated Sodium Channels
3. Sensory Reception
4. Synaptic Reception
5. Somatic Sensation
6. Auditory and Vestibular Function
7. Vision
8. Chemical Senses
9. Posture and Movement
Neurological Examination outline
Brain Connections for Movement
10. Autonomic Function
11. Cerebral and Higher Function
Reading and the Brain Revised
Mental Status Examination outline
Glossary
References
Quiz Yourself—Part I
Quiz Yourself—Part II
Copyright Acknowledgments
Supplementary Updates
Brain Basics
Neurotransmitters Overview
Preface
Neuroscience ABCs: Human and brief, with diagrams and review
Brain Physiology Guide is for questions included.
students of medicine, neuroscience,
health sciences, psychology, and biology, Each chapter contains a concise and
and all other readers who want to illustrated discussion of a particular
understand the basics of how the human topic. The aim is to be clear about
brain and nervous system work. essentials rather than encyclopedic in
Neuroscience has made major advances scope. Topics are
recently, while it’s an important
Structure and function (design and
university program required for a wide activity)
range of subjects. This book can
The nerve impulse (how signals are
accompany a neuroscience,
sent)
neurobiology, medical or physician’s
assistant class, or to help to review Sensory reception (from the
environment to the brain)
them, and interested readers from other
backgrounds can find that it helps Synaptic transmission (from one cell
understand how the nervous system to another, with neurotransmitters)
works in neurological and mental health Somatic sensation (touch, pain,
and disease. temperature)
Auditory and vestibular function
It provides an understanding of how the (hearing and balance)
brain works to transmit signals, feel Vision (how the eye and brain work
bodily sensations, hear, see, control together to detect objects, shapes,
movements, think, and remember. colors, faces, and words)
Chemical senses (taste and smell)
Neuroscience can be difficult, so I’ve
Posture and movement (standing,
made a special effort make it readable reflexes, voluntary movement)
Autonomic function (sympathetic neurotransmitters are highlighted at the
and parasympathetic systems) end of the book. For smartphone
Cerebral physiology and higher readers, you may try holding the phone
functions (attention, sleep, speech, horizontally if the text seems too small.
memory, EEG)
7. Vision How the eye works and how the visual 129
system converts light into sensed visual
images
9. Posture and Movement How motor systems control the skeletal 165
muscles
Glossary 233
References 242
Quiz. Yourself 244
Index 251
1. Structure and Function
The Cellular Structure of the Nervous microscopic gap at a junction called a synapse.
System The axon terminal is typically enlarged and
contains chemical compounds in numerous
Neurons membrane packages called vesicles. When the
axon terminal forms a synapse upon the cell
The building blocks of the nervous system are body (soma), dendrite, or axon of another
the nerve cells or neurons. Neurons are special- neuron, the synapses are termed axosomatic,
ized to generate and carry information by axodendritic, or axoaxonal, respectively. The
means of electrical and chemical signals. A axon may also terminate on a muscle cell in a
typical neuron, as shown in figure 1-1, consists synapse called a neuromuscular junction, or on a
of an enlarged area called the cell body (or gland cell.
soma), which contains the nucleus and much
of the metabolic machinery of the cell; the Receptors
dendrites, several short, thin extensions that
divide into branches like a tree and receive Sensory receptors are structures that convey
information from other neurons; and the axon, information about the outside world and the
a specialized thin extension that transmits interior of the body to the spinal cord and brain
neural signals for a relatively long distance, (figure 1-2). Anatomically, they include a
after which it divides into several branches that wide variety of types: free nerve endings; nerve
end on other cells. The axon shown in figure endings associated with capsules, hair follicles,
1-1 is covered by segments of an insulating muscle fibers, or other cells; and highly mod
lipid material, the myelin sheath, and arises from ified nerve cells, such as the rods and cones of
the cell body at the axon hillock, a cone-shaped the retina. Each sensory receptor is specialized
region, followed by the initial segment, a short to receive a particular type of stimulus (touch,
length of bare axon before the myelin-covered light, sound, chemicals, or others) and trans
length of axon begins. late it into an electrical signal. After modifica
Variations on this basic scheme can be tion, these signals are carried into the spinal
found. For example, the axon may lack a cord and brain by sensory nerve axons as
myelin sheath; the axon may be short; or the sociated with the receptors. Thus, receptor
cell body (in spinal sensory neurons) may be activity is the initial input stage of the nervous
placed to one side of the axon. system.
Synapses Effectors
The axon of one neuron (or its branch) Moving, talking, and facial expression are
communicates with another neuron across a controlled by muscle fibers, one form of effector
2 Structure and Function
Figure 1-1. (A) Diagram of a typical neuron, show General Plan of the Nervous System
ing cell body, dendrites, synapse, axon with myelin
sheath and axon terminal, and a neuromuscular
The two main divisions of the nervous system
junction. (B) Photomicrograph of an alpha
motoneuron in ventral ham of cat spinal cord. Four are the peripheral nervous system (P.NS) and the
dendrites emanate from the cell body. Arrow points central nervous system (CNS) (figure 1-3).
to the axon, at the junction of the initial segment
with the myelinated portion. (C) Photomicrograph of
a bundle (or fascicle) of myelinated axons, forming
a small peripheral nerve, shown m cross-section. The Peripheral Nervous System
Dark borders around each axon are myelin sheath.
Whiu circle in center of bundle is a blood vessel The peripheral nervous system (PNS) consists
The nerve shown is cranial nerve IV (trochlear) of
o f the peripheral nerves, bundles of axons (nerve
the rat, which supplies eye muscle fibers; it contains
an average of 270 axons, the largest of which are fibers) that function to carry signals back and
10 n in diameter. Courtesy of Dr. James M. Kems. forth between peripheral organs (receptors and
effectors) and the central nervous system.
(figure 1-2). In addition, the internal environ These nerve fibers are subdivided according to
ment of the body, including blood pressure, the peripheral organs on which they end:
metabolic rate, and body temperature, is con somatic nerve fibers supply skin, skeletal mus
trolled not only by muscle fibers but also by the cles, tendons, and joints, and visceral nerve
secretions of glands, another form of effector. fibers supply the gut and other visceral organs.
The effectors are the means by which the Classified according to function, afferent nerve
nervous system exerts an effect on both the fibers carry sensory information to the central
outside and inside environments. Their activity nervous system (input), while efferent nerve
represents the final output of the nervous fibers carry motor-command signals from the
system. central nervous system to the effectors (out-
A
3
4 Structure and Function
Figure 1-2. Several types of sensory receptors and an 4. Visceral efferent, carrying motor-command
effector. The three receptors on top are associated signals to visceral effectors (smooth muscle,
with skin; the rod is in the retina; and the muscle cardiac muscle, and glands); these nerves are
stretch receptor is associated with specialized (in
also called the visceral motor or autonomic
trafusal) muscle fibers in skeletal muscle. A larger
(extrafusal) muscle fiber that acts as an effector is nervous system, and are further subdivided
shown at lower right. Arrows indicate direction of into sympathetic and parasympathetic
neural signals. systems.
Dorsal column
D O R S At
Dorsal root
Dorsal horn (Gray matter)
White matter
Ventral root
sory cortex of brain; descending fiber shown carries white matter, dorsal column is shown. Dorsal root
motor information from motor cortex of brain to contains afferent fibers, and ventral root contains ef
spinal motoneuron. (B) A segment of spinal cord, ferent fibers originating in motoneurons; dashed lines
corresponding to section within dotted lines in A. indicate individual fibers corresponding to those
White matter contains axons, gray matter contains shown in the reflex arc in A. Enlarged area (gan
cell bodies, dendrites, and synapses. Within gray glion) of dorsal root contains cell bodies of afferent fibers.
matter, dorsal and ventral horns are shown; within Dorsal and ventral roots join to form peripheral nerves.
5
Neurons in a Spinal Reflex
Muscle
Sensory Axon
Receptor
Reflex arc
Muscle
Effector Synapse
Motor Axon
Cell Body
This simplified cartoon shows a muscle receptor responding to stretch, exciting a motor
neuron that causes the muscle effector to contract. Color coding show sensory axons in blue,
and motor axons in red. Together, they form a reflex arc.
3
Brain Connections
with Spinal Neurons
Brain
Ascending &
Descending
Tracts
Muscle
Receptors Sensory Axons
Spinal
Reflex arc Cord
Muscle
Effector
Motor Axon
CNS
In addition to the reflex arc, sensory tracts ascend to a sensory area in the brain, and motor areas
send descending tracts to control spinal motor neurons.
6 Structure and Function
I Olfactory Smell
II Optic Vision
III Oculomotor Eye movement
Constriction of pupil
Accommodation of lens
IV Trochlear • Eye movement
V Trigeminal Somatic sensation from part of Middle-ear reflex (via tensor tympani
face and head muscle)
Chewing (mastication)
VI Abducens Eye movement
VII Facial Taste from anterior two-thirds Middle-ear reflex (via stapedius
of tongue muscle)
Facial expression
Tearing (lacrimation)
Salivation
The peripheral nerve fibers enter and exit through S3. The peripheral nerves directly
the spinal cord via a series of roots: the dorsal connected to the brain are cranial nerves of
roots carry afferent nerve fibers into the cord which there are twelve pairs. Their names and
and the ventral roots carry efferent nerve fibers major functions are listed in table 1—1.
from the cord. The roots are labeled according
to associated segments of the vertebral column:
The Central Nervous System
cervical (C), thoracic (T), lumbar (L), sacral
(S), and coccygeal (Co). They are numbered The central nervous system (CNS) consists of
within each segment (C l through 8; T1 the brain and spinal cord (figure 1-3A ), each
through 12, LI through 5, SI through 5, and of which is enclosed in a bony covering, the
Co-1). The arm is innervated by spinal nerves brain within the skull and the spinal cord
C4 through T1 and the leg by nerves L2 within the vertebral column. Both are divided
7 Structure and Function
station for information traveling up to the the temporal lobe is the amygdala, contain
cerebral cortex from other areas. The hy ing nuclei that can influence such behaviors
pothalamus (ventral to the thalamus) is a as fighting and eating; the amygdala is part
control center for visceral functions, includ of the limbic system, a group of primitive
ing, for example, changes in cardiovascular structures within the cerebrum.
function during anger and changes in thy
roid gland secretion during cold exposure.
Neural Signals
4. The cerebrum, divided at the midline into
two cerebral hemispheres connected by a thick The nervous system uses signals to transmit
band of axons called the corpus callosum, is information. Information is transmitted from
the largest and most visible part of the the outside world into the nervous system (by
human brain. Comparison of the human means of sensory processes), from one part of
cerebrum with that of lower vertebrates the nervous system to another, and from the
suggests a great increase in size and function nervous system back to the outside world (by
in the course of evolution. The outer surface means of motor processes). The signals are
or cerebral cortex is known as the gray matter carried by the nerve cells, including their
because of its layers of nerve cells. The extensions, the dendrites and axons. In every
surface area of the cortex is increased by its day life, signals include traffic lights, Morse
numerous folds (gyri), separated by grooves code, and the electronic pulses within com
(sulci and fissures). Beneath the cortex is puters. In the nervous system, signals can be
the white matter, consisting of nerve fibers divided into action potentials, graded receptor
traveling to and from the cells in various potentials, graded synaptic potentials, and
parts of the cortex. Each cerebral hemis chemical transmitter release.
phere is divided into four lobes (figure 1-5):
the occipital (at the back of the head), the
Action Potentials
parietal (top), the temporal (sides), and the
frontal (at the front of the head, behind the Like one of the voltaic cells in a battery, the
forehead). The cortex contains motor areas nerve cell at rest is electrically charged across
that govern voluntary movement and keep its membrane. The charge gives rise to an
reflex movements under control, sensory electrical potential difference, a membrane po
areas that register incoming information tential, measured in millivolts (thousandths of a
from sensory receptors (with separate areas volt, abbreviated mV). The magnitude of this
for vision, hearing, body sensation, and so potential difference ranges from 55 mV to 90
on), and association areas that associate mV, with the inside of the cell negative
information from several sources and carry compared with the outside. This membrane
out such higher functions as the understand potential is altered by the action potential, a
ing of speech and the recognition of objects. brief reversal of polarity that sweeps along the
Within the white matter are gray areas that nerve axon (figure 1-6).
contain clusters of cells and synapses and are The action potential is triggered by electrical
divided into nuclei. Between the sides of the or other stimuli above a certain strength.
thalamus and the cerebral cortex are the When neural signals need to be transmitted
basal ganglia, containing several nuclear over long distances, such as from one end of an
structures that help to regulate posture and axon to another, the signals used are action
movement. Below the surface of the tip of potentials.
9 Structure and Function
central
sulcus
parietooccipital
fissure
preoccipital
anterior ram us notch
premotor
area motor area
visual
vestibular area
taste area
area
Figure 1-5. Lateral views of the left cerebral hemis Graded Receptor Potentials
phere. (A) Division into bbes; sulcus and fissure
are terms for grooves in the cortex; the lateral fissure
has three rami or branches. (B) Subdivision into In neurons specialized to receive sensory stimuli
areas based on function (auditory, visual, and so (receptors), stimuli can produce a change in
on) and on the structure of cell layers, or cytoar- membrane potential called a receptor potential.
chitecture, with numbers given by Brodmann. From The change is graded in size, depending on the
Barr, M .L The Human Nervous System: An
Anatomic Viewpoint (3rd ed.). Hagerstown, Md.:
stimulus. For example, in receptors sensitive to
Harper & Row Publishers, Inc., 1979. Reprinted pressure on the skin, the greater the pressure
with permission. the greater the potential change (figure 1-6).
Functional Magnetic
Resonance Imaging (fMRI)
Functional Magnetic Resonance Imaging, or
fMRI, shows brain scans based on the
observation that when neurons in a brain
region become active, they use more oxygen
for metabolic activity. This leads to a local
increase in oxygenated blood to that region.
Within a special, strong magnetic field, the
localized increase in oxygenated blood
produces an image that can change over time,
while a person is lying still or performing a
mental task, from which we can infer localized
changes in neural activity.
Figure 1-6. Neural signals. S = stimulus (arrows The Timing of Neural Activity
below graph indicate stimuli of three intensities),
R .P . = receptor potential, A.P. = action potential, Since neurophysiology concerns sequences of
S.P. = synaptic potential. activity in which information is transmitted by
precise electrical and chemical signals, time is
an important dimension. Time is generally
Graded Synaptic Potentials measured in milliseconds (thousandths of a
Synapses are specialized regions where two second, abbreviated msec). There are four basic
neurons communicate across a microscopic parameters that can be measured: duration,
gap. At synapses, neurons respond to signals velocity, intervals, and frequency.
arriving from other nerve cells by generating
synaptic potentials. Like receptor potentials, Duration
these are graded in size according to the
strength of the stimulus. Another type of The duration of an action potential is easy to
synapse is the junction between nerve and remember— about 1 msec. During this brief
muscle cells. time, the membrane state changes from resting
Graded potentials, whether in receptors or to active and then returns to resting. Graded
synapses, carry information only over very potentials generally last longer; synaptic poten
short distances, such as within a nerve cell tials in the spinal cord, for example, have
body. durations of approximately 12 msec.
The synaptic gap between neurons is generally The action potential is conducted along axons
crossed by chemical transmitters (for example, at velocities ranging from about 1 to 100 m/sec,
acetylcholine or norepinephrine). When an depending on the structure and function of the
action potential reaches the end of a nerve axon. For example, among different fibers in
axon, it causes the release of a chemical peripheral nerves, those conducting at a veloc
transmitter stored in the axon ending, which ity of about 100 m/sec may carry muscle-length
diffuses across the synaptic gap to the next information; those conducting at about 50
nerve cell. Chemical transmitters, therefore, m/sec may carry touch and pressure informa
carry information from cell to cell. tion; and those conducting at about 1 m/sec
11 Structure and Function
Stim. to Spinal
Fnger-»-Wrist (MN) * cord » Cortex
l I____________________ I_____________________ I
0 3 12 25
Tme (msec)
Figure 1-7. The time sequence of neural responses stimulus generally causes a shorter interval
along a sensory pathway. MN = median nerve. between action potentials.
(Neural responses at cervical spinal cord and somatic
sensory cortex are averaged evoked potentials, ob
tained by computer processing of electrical signals re Frequency
corded with disc electrodes on die skin).
Frequency can be defined as the number of
events per second. In the case of action
may carry pain and temperature information. potentials in a single nerve cell, the frequency
These velocities are slowed down in certain is the number of action potentials per second
diseases and therefore can be useful in clinical and is the inverse of the mean interval between
diagnosis. action potentials; the frequency is high when
the intervals are short. Thus, a stronger
Intervals stimulus generally causes a higher frequency of
action potentials. Another way of approaching
Time intervals in neural transmission can be this is to say that the frequency of action
precisely measured, even from outside the potentials generally acts as a code, transmitting
body. For example, recording electrodes on the information about the strength of a stimulus.
skin can be used to measure the intervals (Inhibition and other factors modify this code
between an electrical pulse applied to the in ways that will be discussed in later chapters.)
fingertip and the action potentials at several In the case of electrical waves recorded from
stages of the sensory pathway. The action the brain as part of the electroencephalogram
potential is first conducted from the digital (EEG), the frequency is the number of waves
nerves in the finger to the median nerve, from or cycles per second (c/s). A well-known type
which it can be recorded at the wrist after an of EEG pattern consists of alpha waves, occur
interval of about 3 msec (figure 1-7). After ring at about 10 c/s. Different levels of sleep or
about 25 msec, a neural response can be wakefulness are associated with higher or lower
recorded over the appropriate area of the EEG frequencies.
cerebral cortex. When the time interval and The components described thus far can be
the distance between two points along the further organized according to three overall
peripheral nerve are determined, distance can processes: sensory, motor, and higher.
be divided by time to calculate the conduction
velocity. For the pathway to the cerebral
cortex, the time interval includes synaptic
Sensory Processes
delays as well as nerve-conduction time.
In a single nerve cell, the time intervals A primitive animal creeping along the ocean
between action potentials depend on two fac floor has a reasonable chance of survival if it
tors: the properties of the nerve cell and the can escape when brushed by a larger predator,
strength of the stimulus affecting it. A stronger and a greater chance if it can sense the shadows
yer
A Giant Squid
(ClipArtist.info, Public Domain)
Outside
Inside
Figure 1-9. Specificity of receptors. The information arriving over these and
other channels is integrated by the spinal
fibers and by the motor systems in the CNS motoneuron (figure 1 -8 ) before it produces one
that govern these nerve fibers. or more action potentials as an output signal to
Consider some of the motor processes in the muscle effector.
volved in picking up a book. A convenient
structure on which to focus attention is one of Conduction
the spinal motoneurons supplying the arm
The action potentials are then conducted along
muscles.
the motoneuron axon (a somatic efferent fiber)
to the arm muscle it supplies. Each axon
Integration supplies a number of separate muscle fibers
The motoneuron receives information from within the muscle. At the nerve ending a
chemical transmitter is released, crosses the
numerous sources over thousands of synapses.
The sources include the muscle stretch recep neuromuscular junction, and alters the electri
cal potential of the muscle membrane.
tors, which provide information about the
length of the muscle, and descending fibers
from various motor centers in the brain (figure Transduction
secretions of the digestive tract, is regulated can be found during coma and epilepsy, as well
largely by activity in smooth muscles and as other conditions. In addition, computer
exocrine glands. These effectors are governed processing of the EEG can provide recordings
by the visceral efferent fibers in the autonomic that reflect the brain’s electrical response to a
nervous system. sensory stimulus, such as a click or a visual
The autonomic system is divided into a pattern. These responses, called averaged evoked
sympathetic division, important in protecting potentials, can be altered by physiological,
against danger and cold, and a parasympathetic psychological, and pathological processes.
division, partly involved in digestion and other
relaxing activities. Since both systems are
Language and Left-Right Cerebral Differences
important for regulation of the internal envi
ronment of the body, an imbalance, such as People communicate with each other by means
excessive sympathetic activation due to a of language, both spoken and written. The
tumor in the adrenal medulla gland, can seri neural basis of language has been demonstrated
ously impair this regulation. Visceral efferent mostly by the study of patients with some sort
activity is commonly considered involuntary, of language disorder. Language problems can
but under certain conditions people can learn result from deafness (a sensory disorder) or
to affect variables such as their heart rate. paralysis of the tongue and throat (a motor
disorder), but in addition, there are language
disorders, called aphasias, that are due to
Higher Processes and Cerebral Activity disturbances in higher processes. In the major
A number of psychological processes are not ity of such patients, lesions have been found in
purely sensory or motor and are often called the left cerebral hemisphere, leading to the
higher functions or higher processes because of conclusion that the left hemisphere is domi
their complexity and the belief that they are nant for language in the majority of the
better developed in humans than in lower population (over 90%). Particular areas within
animals. These processes include consciousness the parietal, temporal, and frontal lobes are
(ranging from deep sleep to alertness), lan related to different forms of aphasia; one form
guage, and memory. Their partial association involves an inability to understand speech,
with cerebral activity can be shown by elec another involves an inability to produce it.
troencephalography and other physiological Studies of patients with severed connections
techniques as well as by their impairment in between cerebral hemispheres (split-brain pa
disorders affecting the cerebrum. tients), methods of activating and recording
from each hemisphere separately, subtle struc
tural differences between left and right areas of
Consciousness and the Electroencephalogram
cerebral cortex, and a clinical test in which left
The electroencephalogram (EEG) can be re and right hemispheres can be anesthetized
corded with metal disc electrodes placed on the separately for a few seconds are extending our
scalp and connected to electronic devices that understanding of the cerebral control of
amplify and display the minute electrical waves language.
emanating from the cerebral cortex. The fre
quency of these waves is related roughly to
Memory
levels of consciousness or alertness, from about
2 c/s during a deep sleep up to about 20 c/s To students of the biomedical sciences who
during excitement. Clinically, EEG changes must remember enormous amounts of material,
15 Structure and Function
memory has more than theoretical interest, havior due to experience, but on more
How much can it hold, and how long does it complex and advanced level.
last? One experimental approach is to present a
list of items (words or numbers) and ask
someone to repeat them in the correct order
immediately afterward. For example, you might
read each of these lists of numbers, one digit at
a time, and ask someone to repeat them
immediately after each list:
List 1: 6 4 1 9
List 2: 18 2 5 6 4
List 3: 2 1 8 6 4 9 3 2 8 9 7
You've used neurons in your occipital lobe (to see the shapes of the letters), temporal lobe (for
recognizing the shapes as letters, words, and their meaning), and frontal lobe (for scanning the
screen and holding the reading device).
After each chapter, you'll see review questions, but you can skip through them unless they help
prepare for a class or test. And some missing page numbers that came when the pages were
formatted for an eBook.
Next, travel through the nervous system to what happens at the synapses, how we sense the
world, prepare for action, move around, speak, sleep, and so on. Have a good trip.
The Author
21 Structure and Function
Review Questions
1. Diagram a neuron, labeling the cell body, dendrites, axon, and synapses.
3. List and define the major components of the peripheral nervous system and the
central nervous system.
6. Compare action potentials with graded potentials. Compare their roles in neu
ronal communication.
7. List approximate values for. (1) the duration of an action potential, (2) the con
duction velocity of a typical somatic afferent axon responding to touch, and (3)
the time interval between a stimulus applied to the fingertips and a response in
the cerebral cortex.
10. Give examples of how levels of consciousness, language, and memory can be
studied scientifically and related to cerebral function.
11. Explain die terms in italics in the following passage from die instructor in a
childbirth preparation class: "During labor and delivery, a general anesthetic
reduces die level of consciousness by affecting synaptic transmission in die CMS,
particularly in the reticular form ation; on the other hand, a local anesthetic can be
injected to block conduction in the spinal cord, roots or peripheral nerves at a sacral
level."
2. The Nerve Impulse
The nerve cell is the building block of the potential is a sudden, large electrical change
nervous system, and its role is to carry signals. from this resting potential.
The first signal to consider is the nerve impulse
and its electrical manifestation, the action poten
The Nerve Membrane
tial. The action potential, like the other signals
of nerve cells, depends on electrical and chem The nerve membrane (figure 2 -2 ) separates the
ical changes across the nerve membranes. Un internal solution (axoplasm) from the external
derstanding these electrical-chemical changes solution (extracellular fluid). According to
may be difficult, but is important in interpret classic concepts, it is composed of a lipid Mayer,
ing (1) normal nerve activity, (2) abnormal in which the nonpolar chains face toward the
nerve activity (as in epilepsy or multiple center of the membrane and the polar ends face
sclerosis), and (3) the action of drugs (such as outward. On the sides of the membrane are
local anesthetics). The same concepts are molecules of protein. There are thought to be
useful in the study of physiology of heart and aqueous channels or pores, lined with protein,
skeletal muscle. through which water and certain small ions can
To begin with, consider a single neuron; for travel. Since only certain ions can pass through
example, a motoneuron, which has its cell it, the membrane is called semipermeable.
body in the spinal cord and sends its axon via The solutions inside and outside the cell
the sciatic nerve to the leg (figure 2 -1 ). The contain about 155 mEq/liter of cations (posi
axon of this neuron, like other axons or nerve tively charged ions) and the same concentra
fibers, is specialized to communicate informa tion of anions (negatively charged ions).
tion from one place to another— in this case, However, their composition differs.
from the spinal cord to the lower leg, a distance There is a high concentration of potassium
of about one meter. This information is sent by ions (associated with large, organic anions)
means of nerve impulses, which can be electri inside and a high concentration of sodium ions
cally recorded as action potentials. Nerve im (associated with chloride ions) outside. We will
pulses are electrochemical changes across the see that this unequal distribution of ionic
nerve membrane. These changes affect only a species leads to an electrical voltage (the
small section of the axon at any instant but resting potential) across the membrane, and
sweep down the axon from one end to another that a change in this voltage gives rise to the
at rates of up to 200 mph. When the nerve nerve impulse.
membrane is in the resting state, an electrical The unequal distribution of sodium and
voltage difference, called the resting potential, potassium ions is maintained by a Na+- K +
exists across the nerve membrane. The action pump in the membrane. This pump uses
25
26 The Nerve Impulse
Figure 2-1. A motoneuron. (A) Motoneuron cell ferent fibers have degenerated). From Eccles, J.C.
body, axon (nerve fiber), and terminals (motor end- The Understanding of the Brain. New York:
pious) on muscle fibers. (B) A peripheral nerve in McGraw-Hill Book Co., 1973. Reprinted with per-
cross-section, showing individual motor axons (the of- mission.
2 7 The Nerve Impulse
Figure 2-5. Diagram of the distribution across nerve brane during neural activity. We will start by
membrane of ionic species that contribute to the rest describing a simple circuit— that of a flash
ing potential. Concentrations are approximate only,
and only a few ions are shown. light. A simple flashlight circuit involves a
battery, a conducting path (wire), and a light
bulb (figure 2-6A ). Positive charge flows from
Changes in K+, or K +„ lead to predictable
change in EK. Increasing K +0 to 15 mM, for positive to negative poles of the battery, con
example, would mean that stituting a current (this is equivalent to saying
that negative charge flows from negative to
150 positive poles). The wire conductor can be cut
Ek = -61 log — = -61 log 10 = -6 1 mV.
and both cut ends inserted into an ionic
solution (figure 2-6B ). Current will continue
Electrical Properties o f the Membrane to flow through the solution, carried by the
dissolved ions: this flow is called volume conduc
We have already mentioned voltage, but other tion. A circuit diagram represents this flow
basic electrical concepts are also important in (figure 2-6C ), and the standard symbols are
understanding what happens across the mem given in table 2 -1 .
V or E Voltage or elec Volts (V) The amount of electrical energy that potentially can be ex
trical potential erted on a charged particle such as an electron or an ion
difference
I Current Amps (A) The flow of charged particles through a conductive medium
such as a metal wire or an electrolyte solution
R Resistance Ohms (Cl) The resistance to the flow of charged particles through a
conductive medium or across a membrane
G or g Conductance Mhos The ease with which charged particles can flow through a
conductive medium; the inverse of resistance (G = 1/R)b
C Capacitance Farads (F) The ability of two conductors in the same vicinity to store
charged particles on both sides of an insulating medium
such as air or the non-porous area of a membrane
“Prefixes to units include: p. or micro-, one millionth; m or milli-, one thousanth; K or Kil(o)-, one thousand, M
or meg(a)-, one million.
bThe ability of a membrane to allow K ions to cross it is its potassium conductance, GK or gK, which is directly
related to its potassium permeability, PK; and similarly for other ions
30 The Nerve Impulse
Figure 2-6. Simple electrical circuits. (A) Battery the battery charge only until the capacitor is
and light bulb, connected with wire conductor. Cur completely charged with positive charge on one
rent (I) is shown as the flow of positive charge; this side and negative charge on the other. The
is identical to negative charge flowing in the opposite capacitor is then able to store the charge on its
direction. (B) Same as A, except that current flows
through a chamber filled with salt solution. (C) plates, separated by the nonconducting ele
Schematic circuit diagram of B, showing a voltage ment in between.
source (V) and a resistance (R). (D) Same as C, These symbols allow us to draw an electrical
with the addition of a capacitor (C). model of a segment of nerve membrane (figure
2 -7 ). In the model, the concentration differ
ence of K + across the membrane is the source
Electric current can be compared with the of electrical pressure, which can be represented
flow of water: the electrical potential (or by a battery whose voltage is Ek, the Nemst
voltage) difference produced by the battery can equilibrium potential for K + (Ek — -9 0 mV).
be compared to the pressure difference pro
duced by a pump, the electrical conductor to a
pipe, and the electric current in the conductor Figure 2—7. Electrical model of a segment of resting
to the flow of water through the pipe. Electrical nerve membrane. Vm is the voltage (potential) dif
resistance is increased by such factors as a ference across the membrane, or
thinner wire conductor or a more dilute ionic Ek is the sodium equilibrium potential, which is the
source of the voltage difference. Gk is the potassium
solution, just as resistance in a hydraulic system conductance, which can be attributed to membrane
is increased by a narrower pipe. channels that allow potassium ions to pass through.
The amount of current, I, flowing through Qn is the membrane capacitance, attributed to the
the electrical circuit is directly proportional to lipid layers of die membrane which separate ions.
the potential difference, V, and inversely pro
portional to the resistance, R; that is, I = V/R
(Ohm’s Law). Conductance, symbolized by G
or g, is simply the inverse of resistance (G =
1/R); a concentrated salt solution in figure 2 -
6B might have a low resistance and a high
conductance, meaning that it conducted ionic
current easily.
Finally, a capacitor— typically two thin
metal plates separated by a nonconducting gap
or sheet of paper— can be inserted into the
circuit (figure 2-6D ). Then current flows from
31 The Nerve Impulse
This concentration difference, represented by or potential during the resting state is called the
Eg, can be visualized as pushing potassium ions resting potential. As we shall see, the resting
out through channels through the membrane. potential is a baseline from which various
These channels offer a resistance, R, or, looked changes— including the action potential—
at in a more positive light, provide a conduc take off.
tance, Gic, to the flow of potassium ions. The
potassium ions that pass through the potassium
channels may be thought of as huddling against Changes in Membrane Potential
the outside wall of the nerve membrane, Changes in membrane potential (from the
attracted there by the large negative ions they resting-potential level) can be measured by
have left behind on the inside. The ability of inserting small-tipped electrodes into the cell.
the membrane to hold opposite charges across These microelectrodes are made from thin glass
it is its capacitance, Cm, and is a property of the capillary tubing. The tubing is heated and
lipid layers that do not allow ions to cross. pulled to form a micropipette or microelectrode
Once a certain number of potassium ions have with an extremely small but open tip. The
collected on the outside of the nerve mem microelectrode is filled with an ionic solution
brane, they repel the outward movement of and connected to a voltage-recording device to
their positively charged brethren because of measure the voltage in a neuron. The recording
simple electrostatic principles; this standoff device measures the membrane potential, in
represents an equilibrium situation, which to a millivolts, between the microelectrode inside
large extent determines the resting potential. the cell and a reference electrode outside the
The effect of the potassium ion concentration cell in the extracellular fluid (figure 2-8).
difference is to deposit positive charges on the Originally, when both microelectrode tip
outside surface of the membrane capacitance and reference electrode are outside the nerve
and to leave the inside negatively charged. fiber, they record the same electrical potential,
Only a few charges, charging the membrane in so the potential difference between them is
this way, are enough to cause a resting poten zero. Once the microelectrode pierces the
tial; they are too few to change the concentra nerve membrane and enters the axoplasm, the
tion difference across the membrane. recorded potential difference goes suddenly
Actually, the membrane is somewhat perme negative to about - 7 0 mV, the resting poten
able to other ions as well as K +. Thus, the tial. In other words, the resting membrane is
membrane potential at rest is not exactly equal polarized to about - 7 0 mV, inside negative.
to Ek (-9 0 mV); it is somewhat less negative Now, electrical stimuli can cause the mem
(-7 0 mV to -8 5 mV).2 This electrical voltage brane voltage to change slightly, .so that it is
either slightly depolarized (less negative on the
2The contribution of the other ions to the membrane inside, for example, -6 5 mV); or hyperpolarized
potential is expressed by the Goldman equation: (more negative on the inside, for example, -8 0
mV). A weak stimulus from an electrical
PK (K +,) + PN,(N a+f) + Pa (Cl-„)
Vro = -61 lo g ---------------------------------------------------- stimulator, as shown in figure 2 -9 , produces a
Pk(K +„) + PN.(N a \ ) + Pcl(C l-) slight depolarization in the membrane poten
Pk. Pn». and P a are the membrane permeabilities of the
tial, as registered by a nearby microelectrode.
different ions. The contribution of each ion to this Other microelectrodes 1 mm and 2 mm away
equation depends on its permeability. A potassium permea record similar but much reduced voltage
bility much greater than that of the other ions would result
in a membrane potential close to the Nemst equilibrium
changes: the depolarization decays with dis
potential for potassium. tance (figure 2 -9 ). A little farther along the
3 2 The Nerve Impulse
when recorded some distance away; this makes The voltage never actually reaches this point
it a good signal for transmitting information. but stops at about + 3 0 mV to + 4 0 mV when
Also, as long as the depolarizing stimulus is repolarizing processes take over.
over a threshold value, further changes in the In the electrical model, the increase in G n3
stimulus have no effect on the height or shape during the depolarizing phase of the spike can
of the action potential. This characteristic is be represented by adding another section (fig
known as the all-or-none law: the action poten ure 2 -1 1 ). The sodium section is drawn heavier
tial is either full size or it does not occur at than the potassium section because the conduct
all. ance to sodium ion is (briefly) much greater than
the conductance to potassium ion during the
rising phase of the spike. (While at rest GNa =
about 1/20 G k; at the peak of the action
The Ionic Basis for the Action Potential potential G n3 has increased more than 100
times its resting value in a very short time).
The action potential occurs because, at Changes in ionic conductance can be
threshold depolarization, N a+ permeability
graphed on the same time axis as the action
channels in the membrane suddenly open, and
potential (figure 2-12):
the membrane quickly becomes much more
permeable to Na+ . The sodium permeability,
Pxja, and thus the sodium conductance, G n3, 1. Sodium conductance rises quickly, causing
increase rapidly. Na+ then rushes into the cell, rapid depolarization.
driven by its concentration gradient (Na+„ is 2. At the peak of the action potential, the
greater than Na+,) and by the voltage difference inside becomes positive— the degree to
across the membrane. If Na+ concentration which it becomes positive is called the
alone were to determine membrane potential, overshoot.
this potential would be given by the Nemst 3. Only a very small number of Na+ ions move
equation: inward during each action potential (about
34 The Nerve Impulse
OUTSOE
Figure 2-11. Electrical model of a segment of nerve only if the second stimulus is unusually strong.
membrane during the depolarization phase of an ac A main cause of refractoriness is that, once the
tion potential. Identical to model of resting mem
sodium channels have opened and then closed,
brane (figure 2-7), with the addition of a significant
sodium conductance (G \J, due to the opening of it is difficult to get them to open again until a
sodium channels, which allows the sodium equilib
rium potential (E^J to contribute a large part of the
membrane potential. Positive ions pass inward ACTION POTENTIAL
through these channels and are separated from nega
tive charges on the outside by the membrane capaci
tance. The result is a reversal of membrane polarity.
OUTSOE
Figure 2-11. Electrical model of a segment of nerve only if the second stimulus is unusually strong.
membrane during the depolarization phase of an ac A main cause of refractoriness is that, once the
tion potential. Identical to model of resting mem
sodium channels have opened and then closed,
brane (figure 2-7), with the addition of a significant
sodium conductance (G \J, due to the opening of it is difficult to get them to open again until a
sodium channels, which allows the sodium equilib
rium potential (E^J to contribute a large part of the
membrane potential. Positive ions pass inward ACTION POTENTIAL
through these channels and are separated from nega
tive charges on the outside by the membrane capaci
tance. The result is a reversal of membrane polarity.
depolarization and excitation. Since the cur diameter, the lower the internal longitudinal
rent jumps ffom one node to the next, conduc resistance, the greater the current flow, and the
tion in myelinated axons is called saltatory, higher the conduction velocity. An additional
after the Latin for jump, saltare. factor increasing conduction velocity in my
DemyeUnation occurs in multiple sclerosis and elinated fibers is the greater spacing between
other diseases. The myelin sheath around the nodes found in larger-diameter axons. Experi
intemode degenerates, so that some of the ments show that in myelinated axons the
stimulating current passes through the inter conduction velocity (in m/sec) increases with
node and only the current left over passes diameter, and equals approximately 6 X the
through the next node. The current left over diameter (in p.). For example, in myelinated
may be so small that conduction may be slowed fibers; an axon 2 p. in diameter conducts at 6 x
or blocked. Ordinarily, the stimulating current 2 = 12 m/sec; an axon 5 p. in diameter
passing through the node is more than enough conducts at 6 x 5 = 30 m/sec; and an axon 10
to depolarize the node to threshold and excite p. in diameter conducts at 6 X 10 = 60 m/sec.
an action potential. That is, the ratio of This relationship is graphed in figure 2-15. We
stimulating current to threshold current, called will see in chapter 5 (Somatic Sensation) that
the safety factor, is greater than one. In demy- fibers of different size and velocity have differ
elination, it may be reduced to less than one. ent functions. In addition, the conduction
In both unmyelinated and myelinated nerve velocity of fibers in the human body can be
fibers, currents must travel longitudinally along measured, and is used as an index of nerve
the inside of the axon. The larger the axon damage and recovery.
Conduction
velocity
(m/sec)
Diameter (microns)
37 The Nerve Impulse
cell membrane. If the electrode is internal to a changes due to conduction in a large number of
single neuron, it records a positive-going spike axons at the same time. As more fibers become
(figure 2-16A ). However, if the electrode is active (with stronger stimulation) the response
external to a single neuron, it records a nega becomes larger. The response has several peaks
tive-going spike (figure 2-16B ), since the representing fibers that conduct at different
outside of the membrane briefly becomes more speeds. The first peak represents the largest-
negative during the spike. This record is ob diameter, fastest-conducting fibers; the next
tained when the reference electrode is placed peak represents smaller-diameter, slower-
over a damaged section of nerve that does not conducting fibers; and so on. Since the com
conduct an action potential itself. pound action potential can change in height
If the electrodes are placed outside a whole, and form, it does not look like an action
compound nerve, consisting of many individual potential from a single neuron and does not
axons, a compound action potential is obtained obey the all-or-none law.
(figure 2-16C ). The compound action poten
tial represents a combination of the voltage
A Giant Squid
(ClipArtist.info, Public Domain)
Outside
Inside
Review Questions
2. State concentrations for sodium, potassium, and chloride ions within mam
malian nerve cells and in the extracellular fluid.
3. Calculate the equilibrium potential for potassium and sodium using the Nemst
equation.
4. Define and give units of measurement for electrical potential, resistance, cur
rent, capacitance.
41 The Nerve Impulse
Review Questions
2. State concentrations for sodium, potassium, and chloride ions within mam
malian nerve cells and in the extracellular fluid.
3. Calculate the equilibrium potential for potassium and sodium using the Nemst
equation.
4. Define and give units of measurement for electrical potential, resistance, cur
rent, capacitance.
41 The Nerve Impulse
Review Questions
2. State concentrations for sodium, potassium, and chloride ions within mam
malian nerve cells and in the extracellular fluid.
3. Calculate the equilibrium potential for potassium and sodium using the Nemst
equation.
4. Define and give units of measurement for electrical potential, resistance, cur
rent, capacitance.
42 The Nerve Impulse
5. Define conductance of the membrane and relate this to membrane resistance and
ionic permeability.
7. Define resting potential and state a typical value for this in a neuron.
10. Draw a diagram to show the time-course of the membrane potential changes in
an axon when a sufficient stimulus is applied to elicit an action potential.
11. On such a diagram mark the depolarization phase, the spike potential, the over
shoot, and the repolarization phase.
13. Describe the concept of sodium and potassium channels and the effect of
tetrodotoxin.
14. When the fluid bathing an isolated axon was changed so that one-half of the
external Na+ was replaced with the larger, organic ion choline, the size of the
overshoot was reduced. Explain why this occurred.
16. "The absolute refractory period imposes an upper limit on the frequency of nerve
impulses." Explain this statement as it would apply to a motor neuron with an
absolute refractory period of 4 msec (1/250) sec. What would be the highest pos
sible frequency of nerve impulses in spikes per second?
45 The Nerve Impulse
17. D raw a diagram to illustrate the local current flow that accompanies the passage
of a nerve impulse along an axon.
20. Describe the relationship between the diameter of a myelinated nerve fiber and
its conduction velocity.
45 The Nerve Impulse
17. D raw a diagram to illustrate the local current flow that accompanies the passage
of a nerve impulse along an axon.
20. Describe the relationship between the diameter of a myelinated nerve fiber and
its conduction velocity.
46 The Nerve Impulse
21. Describe a compound action potential and how it differs from a single-axon
action potential
23. In the same patient described in question 19, sensory nerve impulses were stud
ied by electrically stimulating the digital nerve in the middle finger with ring elec
trodes and a special stimulating circuit, while compound action potentials were
recorded over the median curve at tire w rist A n intern assisting in this test
noticed that as the size of the stimulus increased, so did the size of the recorded
potential. He asked whether the equipment was faulty because he expected the
all-or-nothing law to apply. Can you answer?
46 The Nerve Impulse
21. Describe a compound action potential and how it differs from a single-axon
action potential
23. In the same patient described in question 19, sensory nerve impulses were stud
ied by electrically stimulating the digital nerve in the middle finger with ring elec
trodes and a special stimulating circuit, while compound action potentials were
recorded over the median curve at tire w rist A n intern assisting in this test
noticed that as the size of the stimulus increased, so did the size of the recorded
potential. He asked whether the equipment was faulty because he expected the
all-or-nothing law to apply. Can you answer?
3. Sensory Reception
As you read this page, the words act on you by found in subcutaneous tissue in the hand
means of light energy. Music and speech act on (figure 3 -1 ). Under a microscope, it is an oval
you by means of sound energy. Another per structure consisting of many layers (lamina),
son’s handshake presses on your hand, com rather like an onion. Within the core of this
municating by means of mechanical energy. structure is a nerve ending. While the ending
Within the nervous system, information is of the nerve fiber is bare of myelin, a myelin
transmitted by means of electrochemical signals sheath surrounds the fiber as it exits the
called nerve impulses. Thus, the pressure from corpuscle. The nerve fiber and the specialized
a handshake must somehow be converted into tissue surrounding it together constitute a re
nerve impulses in the hand, which are then ceptor. If you press quickly on someone’s hand,
transmitted up the median, ulnar, and radial the mechanical energy is transmitted through
nerves in the arm to the spinal cord and, the skin to the pacinian corpuscles. What
finally, to a sensory area of the cerebral cortex. happens next was shown in a remarkable series
How is the mechanical pressure first converted of experiments using pacinian corpuscles
into nerve impulses in the hand? How do these isolated from animals, and poked by an elec
nerve impulses carry information about the trically driven probe (figure 3-2A ).
type and amount of pressure? How do similar
processes occur for vision, hearing, and the Receptor Potential. A mechanical indentation of
other senses? These questions are taken up in the outer surface of the corpuscle is transmitted
this chapter. through the corpuscle to the membrane of the
nerve ending at the core of the corpuscle. The
nerve membrane converts or transduces the
Transduction in Mechanoreceptors mechanical indentation into an electrical de
The process of converting one form of energy polarization. In this depolarization, the inside
(in this case mechanical energy) into another of the nerve ending, at a resting potential of
form of energy (in this case the electrochemical about -7 0 mV, becomes a few millivolts less
energy of the nerve impulse) is called transduc negative, for a few milliseconds. This depolari
tion. This transduction occurs in specialized zation is called a receptor potential (because it
cells in the skin and subcutaneous tissue called takes place in a receptor) or a generator potential
receptors, or more specifically, mechanorecep (because if large enough it can generate an
tors, since they respond best to mechanical action potential). The use of pharmacological
energy. substances that block action potentials has
permitted the study of receptor potentials in
Example 1: The Pacinian Corpuscle isolation.
Consider the pacinian corpuscle, a receptor The receptor potential of the pacinian cot-
47
48 Sensory Reception
Stimulus
intensity
-, + 4 0
0 Membrane
potential
- Th C">V)
- -7 0
49 Sensory Reception
Receptor potential
/ ^
51 Sensory Reception
stretch) of adequate intensity, the frequency of tory period is-2 msec (1/500 sec), 500 spikes/sec
the action potentials is greatest at the onset of is the absolute upper limit of spike frequency for
the stretch, and there is then an adaptation that neuron.
(decline of the response) to a lower level, but Human skeletal muscles are supplied with
this adaptation is slow and some response is stretch receptors that behave in many ways
maintained throughout the stimulus (figure 3— similar to those of crustaceans, but in human
5D). Thus the stretch receptor is useful for stretch receptors the cell bodies are distant
detecting steady stretch of the muscle as well as from the sensory endings. The sensory nerve
changes in stretch. In general, slowly adapting endings are attached to muscle fibers in such a
(tonic) receptors are useful for detecting steady way that a stretch of the muscle deforms the
stimuli as well as changing stimuli, while nerve endings. A depolarizing receptor poten
rapidly adapting (phasic) receptors are useful tial follows, which, if above threshold, leads to
only for detecting changing stimuli. a series of nerve impulses. The timing and
Like other sensory receptors, the crustacean frequency of the impulses inform the CNS
stretch receptor signals the central nervous about the change in length of the muscles,
system by means of the timing and frequency of information used in controlling muscle con
nerve impulses. In the stretch receptor, the traction.
onset of a series of impulses tells the nervous
system approximately when a muscle stretch Other Slou/ly Adapting (Tonic) Mechanoreceptors.
begins. During the stretch, the frequency of On the fingertips, there are slowly adapting
impulses (in spikes/sec) tells the nervous system receptors with disc-shaped endings (Merkel’s
the degree of stretch; a greater stimulus intensity, discs) that transmit information about the
over a wide range, produces a greater receptor- degree of pressure exerted on the fingers. In the
potential amplitude, which produces a higher joint capsules, there are slowly adapting joint
spike frequency (figure 3—6). receptors that signal the angle of the limb at
A higher spike frequency means that spikes the joint, as well as the direction and rate of
come closer together in time, up to a limiting limb movement.
value. Certainly, the interval between two
spikes cannot be shorter than the absolute
refractory period. Thus if the absolute refrac General Classification and Function of
Sensory Receptors
Figure 3 -6 . Response of a slowly adapting receptor There are other types of mechanoreceptors in
(crustacean stretch receptor in this example) to in addition to those described above. These in
creased stimulus intensity. clude pain receptors (or nociceptors, from the
52 Sensory Reception
same root as the word noxious), which are of lights or colors. Experimentally, electrical
specialized to transmit information about se stimulation of auditory neurons can produce
vere mechanical changes (cutting, crushing, the illusion of sound. Electrical stimulation
and so on), or other stimuli that could cause of the median nerve may produce the illusion
damage. In the cochlea of the ear, specialized that the hand is being touched. In certain
mechanoreceptors are sensitive to the minute diseases that affect the peripheral nerve, pain
air-pressure oscillations caused by sound waves. ful or tingling feelings may occur in an arm or
In the vestibular apparatus, other mechanore leg without any outside stimulus. These pares
ceptors are sensitive to movements of the head thesias may be due to some abnormal stimula
and its position with respect to the pull of tion of the peripheral nerve involved or of the
gravity. CNS connections directly.
In addition to the mechanoreceptors, there
are receptors specialized to receive and trans
duce other forms of energy from the environ
ment. Thermoreceptors are most sensitive to
thermal energy (heat and cold). Chemoreceptors
are most sensitive to various forms of chemical
energy, for example, the solutions that cause
taste sensations in the tongue and the gases
that cause odor sensations in the nose. Photo-
receptors— the rods and cones in the retina—
are most sensitive to light energy.
Each receptor, then, is especially sensitive to
a particular form of energy, which is called its
adequate stimulus when the intensity is above
threshold. The receptor transduces this form of
energy into receptor potentials and then action
potentials. The action potentials activate a
sensory pathway in the central nervous system.
Since the action potentials from different re
ceptors are similar, it is believed that the type
(or modality) o f sensation associated with each
receptor is due to its central nervous system
connections— a modem restatement of one of
the earliest generalizations in physiology, the
law of specific nerve energies by Johannes
Muller.
The importance of central nervous system
connections allows us to understand why sensa
tions can sometimes be produced by unusual
stimuli. For example, rubbing the eyes (me
chanical stimulation that may activate sensory
neurons in the retina) may produce the illusion
53 Sensory Reception
Chapter 3
Sensory Reception
The pacinian corpuscle as an example of receptor function. The receptor potential (left), due to opening of mechanically-gated
channels, can initiate an action potential (right), due to opening of voltage-gated channels, if the depolarization is large enough to
cross threshold.
56 Sensory Reception
6. Cite gvamplps of rapidly and slowly adapting receptors, and describe their func
tions.
7. Describe how the timing of afferent nerve impulses can be related to vibratory
movement (for phasic receptors) and the intensity of a mechanical input (For
tonic receptors).
8. Speculate about the effect on sensory experience of a mythical drug that tem
porarily makes all receptors rapidly adapting.
57 Sensory Reception
9. State and explain the significance of the law of specific nerve energies.
10. ' Make a list of receptors classified in terms of the form of energy they transduce..
4. Synaptic Transmission
In the last chapter we saw how a nerve impulse Similar drugs are now used in a controlled
can travel the length of a nerve axon— even a way to aid muscle relaxation during surgery.
distance as long as one meter as in the case of The effect of such drugs illustrates how the
the motor-nerve fibers in the leg. What hap synaptic junctions between cells provide a site
pens when the nerve impulse reaches the end of of action for chemicals. The chemicals can be
the nerve fiber? Since nerve and muscle fibers poisons, as in the case of curare; drugs, as in the
are separated by a gap, how does activity reach case of the blocking agents used during surgery;
the muscle fibers to cause muscle contraction? or substances normally found in the body, such
This process, called neuromuscular transmission, as chemical transmitters.
is one form of synaptic transmission, the trans
mission of information from nerve cell to
muscle, nerve, or gland cell. Not only is
Nerve—Muscle Synapses
synaptic transmission a key to the complex
behavior of the nervous system, but its disorder Neuromuscular transmission is the transmission
gives rise to major diseases (such as Parkinson’s of action potentials from nerve fiber to muscle
disease), and its modification is the basis of the fiber by means of a chemical intermediary. A
action of many drugs (such as general anesthet single motor-nerve fiber will branch and con
ics and tranquilizers) that affect the nervous tact from 10 to 2,000 separate skeletal muscle
system. fibers; since the nerve fiber triggers an impulse
Long before the development of the neuro in all the muscle fibers it innervates, the nerve
sciences, a group of Indians living in the and muscle fibers together are called a motor
jungles of Ecuador modified neuromuscular unit (figure 4 -1 ). Fibers of different units are
transmission with a drug. Their survival de interspersed with each other to ensure smooth
pended on hunting tropical birds with a blow- contraction of the muscle as a whole.
gun— a thin hollow reed about 10 feet long. The neuromuscular junction (figure 4 -2 ) is
The arrow that fit inside was formed from a where the nerve ending meets the muscle fiber.
palm leaf with a point as sharp as a needle. The As it nears the muscle fiber, the motor axon
arrow tip was dipped in a poison called curare, loses its myelin covering and ends in a presyn-
which was prepared from the wild Wourali aptic terminal. In microscopic sections of the
vine. Even at distances of 200 feet, the hunters terminal one can see small synaptic vesicles. A
seldom missed the flying birds. Within three synaptic cleft of about 50 nm (nanometers)1
minutes of being hit by the arrow tip, the bird separates the nerve ending from the specialized
became motionless and fell dead. Curare is a postsynaptic membrane of the muscle fiber. In
neuromuscular blocking agent; it blocks this region the muscle membrane is thrown
neuromuscular transmission and thus paralyzes
the muscles. 'One nanometer equals 10~9 meter or 10 angstrom units.
60 Synaptic Transmission
MOTOR UNIT
/
Spinal
motoneuron
Myelinated
axon
Figure 4-3. Miniature end-plate potentials (MEPPs)
recorded from muscle fiber at neuromuscular junc
w tion.
Muscle
fibers ter, within the cytoplasm of the nerve ending.
The vesicles contain acetylcholine, a chemical
Figure 4-1. A motor unit consisting of a spinal transmitter. In brief, the role of the acetyl
motoneuron with its myelinated axon innervating choline is to leave the vesicles, cross the
several skeletal muscle fibers. synaptic cleft, contact the muscle membrane
on the other side, and there cause electrical
into a series of folds (the junctional folds), depolarization.
which increase its surface area. The neuromus Even when muscle fibers are at rest, without
cular junction is also called the motor end-plate. the influence of nerve impulses, the effects of
acetylcholine can be seen. The resting poten
tial for muscle fibers is about -8 0 mV (as in
Neuromuscular Transmission
nerve cells, inside negative to outside). Super
The synaptic vesicles resemble small mem imposed on this resting potential, however, are
brane-bounded bubbles, about 50 nm in diame- tiny depolarizations of about 0.4 mV— that is,
the potential may briefly become less negative,
going from -8 0 mV to -7 9 .6 mV and back
Figure 4-2. A neuromuscular junction (schematic
diagram). Note the synaptic vesicles within the nerve again (figure 4 -3 ). These small depolariza
ending (presynaptic terminal). tions, called miniature end-plate potentials
M y e lin '
A xon
M u s c le fiber/
N e u ro m u s c u la r
ju n ctio n
P o s ts y n a p tic
m em b ran e
61 Synaptic Transmission
Using the Nemst equation, we can calculate arises (and obsciires the end-plate potential)
that the equilibrium potential for sodium ion is after a smaller depolarization.
positive (E)s|a = +61 mV) while the equilib The location of the acetylcholine receptors
rium potential for potassium ion is negative can be shown by an experiment in which
(Ek = -9 0 mV). A compromise value between acetylcholine is released from a micropipette
the two ((ENa + Er )/2) is -1 5 mV. The end- onto the muscle membrane (figure 4 -6 ). When
plate potential is a depolarization that moves the micropipette is placed immediately over the
toward this value; however, an action potential neuromuscular junction, a normal-looking end-
plate potential results. In this procedure, the
Figure 4-6. Experiment demonstrating concentration micropipette is imitating the normal function
of acetylcholine receptors near motor nerve terminals. of the nerve ending. However, if the micropi
(1) Micropipette (E) releases acetylcholine at several
points near nerve terminal. (II) Greatest sensitivity pette is moved slightly to one side, the end-
is at points a and d, adjacent to nerve terminals, plate potential is much reduced. This illustrates
and least sensitivity is at points m between. (Ill) that the acetylcholine receptor sites are con
Endplate potentials elicited at points a through d. centrated at the muscle end-plate region.
Reprinted with permission from Eccles, J.C . The The same acetylcholine receptor sites can be
Understanding of the Brain. New York: McGraw-
blocked by the poison curare. This blockade
Hill Book Co., 1973. Adapted from Peper, K., and
McMahan, U.J. Distribution of acetylcholine recep causes the end-plate potentials elicited by
tors m the vicinity of nerve terminals on skeletal motor-nerve impulses to be much reduced in
muscles of the frog. Proc. Roy. Soc. Lond. (Series size, so that they fail to elicit muscle action
B) 181:431-440, 1973. potentials, and paralysis can ensue. A number
64 Synaptic Transmission
example (figure 4 -8 ). In this experimental synaptic terminal, the ensuing release of trans
setup, a microelectrode is inserted into the cell mitter from the vesicles is similar to that in
body, and an oscilloscope shows the membrane neuromuscular transmission (but the transmit
potential— the voltage inside the cell minus ter itself may be an amino acid or some other
the voltage outside it. At first, the membrane compound). Also similar is the synaptic delay
potential is at its resting level of - 7 0 mV. At of approximately 0.5 msec, the presence of
time zero, one or more presynaptic terminals receptor molecules on the postsynaptic mem
become depolarized by an action potential. brane, and the effect of the transmitter reacting
Although these inputs to the synapse may be with the receptor molecules— an opening of
either excitatory or inhibitory, we will begin by membrane channels to both sodium and potas
considering the events at excitatory synapses. sium ions, leading to a depolarization from the
After the action potential invades the pre- resting potential on the postsynaptic side.
Figure 4-8. Excitatory synaptic transmission, as Excitatory Postsynaptic Potential. The postsyn
shown by recording with microelectrode in spinal aptic depolarization is called an excitatory
motoneuron. Inset shows superimposed EPSP re postsynaptic potential (EPSP). The diagram (fig
sponses to activation of 5, 25, or 50 excitatory ure 4 -8 ) shows that five excitatory inputs cause
synaptic inputs at time indicated by arrow. Once
action potential begins at initial segment, it is con an EPSP that depolarizes the membrane by
ducted along axon. Reference electrode is outside the about 5 mV, from - 7 0 mV to -6 5 mV. In
celL itself, such a small EPSP is a transient and
65 Synaptic Transmission
localized event; it is subthreshold and does not One-Way Conduction. Synaptic transmission
trigger an action potential. Instead, it dies away proceeds from presynaptic nerve terminals to
after about 15 msec, in a process called temporal the postsynaptic cell, rather than in the oppo
decay. Also, even when the EPSP at the site direction. Although conduction of an
synapse is at its peak, it may be much reduced a action potential along an axon is also generally
few micrometers away from the synapse, a in one direction, it is possible for an action
process called spatial decay. Thus, the EPSP by potential to move in both directions away from
itself is a local process of limited extent, and, the site of stimulation if an axon is stimulated
therefore, is of no use in transmitting a signal somewhere between the cell body and the axon
for any distance along the axon. terminal. Thus, synaptic transmission differs
The amount of depolarization increases as from action potential conduction in this re
more presynaptic fibers fire at the same time. spect and may be compared with one-way
The diagram shows that increasing the number streets regulating the flow of automobile traffic
of inputs from 5 to 25 increases the amount of during rush hour.
depolarization to -6 0 mV. Thus, the EPSP is a
graded process that can increase in size as a Synaptic Delay. The time between depolariza
function of the amount of excitatory input. tion of the presynaptic terminal (generally
because of an invading action potential) and
Threshold. In the example shown in figure 4 -8 , the onset of depolarization in the postsynaptic
only when 50 inputs fire does the depolariza membrane is approximately 0.5 msec. This is
tion reach the threshold value of -5 5 mV. At the time required for the release of transmitter,
this point, an action potential starts in the its diffusion across the synaptic cleft, and its
initial segment, the most sensitive region of the activation of the postsynaptic membrane.
motoneuron. The action potential than sweeps
down the axon, jumping from node to node by Graded Response. We have seen that the EPSP
saltatory conduction. is graded in size, in that the amount of
The number of inputs and the threshold depolarization increases with the number of
voltage are mentioned only as typical values excitatory inputs active at the same time. The
found in some spinal motoneurons. The gen action potential, on the other hand, does not
eral principle is that, in most neurons studied, increase in size with increased input: it is an all-
more than one excitatory input is required to or-none event.
elicit an action potential output, and the In neuromuscular transmission, a graded
action potential originates at a particular low- end-plate potential can be seen when the
threshold area on the neuron, generally the muscle action potential is blocked with curare.
initial segment. Under normal conditions, however, a nerve
impulse in the motor axon results in an end-
plate potential large enough to elicit a muscle
Characteristics o f Synaptic Transmission
action potential, so that gradation of response
The excitatory synapse on the spinal moto is not a factor.
neuron exemplifies a number of general charac
teristics of synaptic transmission. These Summation. The ability of the EPSP to increase
characteristics are found in other chemical when more and more excitatory synapses are
synapses as well, including the neuromuscular simultaneously active is called spatial summa
junction and the inhibitory synapses, which tion, since it represents a summing of inputs
remain to be considered. distributed along the spatial dimension of the
66 Synaptic Transmission
postsynaptic neuron. In addition, when nerve mV, and that for chloride ion is - 7 0 mV, the
impulses invade an axon terminal one after the effect of the permeability increase for both ions
other at a high enough rate, the resulting EPSP is to drive the membrane toward the inter
will also increase, a feature called temporal mediate value of —80 mV. This represents a
summation. membrane potential that is more negative on
Under normal conditions, numerous sy the intracellular side (hyperpolarized) with
naptic inputs may arrive at different locations respect to the resting potential of -7 0 mV.
on the neuron, each at different frequencies When the membrane starts at its normal
and somewhat different times. Thus, both resting potential, the activation of an inhibi
spatial and temporal summation play a part in tory synaptic input elicits an IPSP that consists
determining whether or not an action potential of a hyperpolarization of a few millivolts or less,
is generated in the postsynaptic cell. which resembles an inverted EPSP (figure 4—
9). Like EPSPs, IPSPs have a synaptic delay,
are graded according to the number of inputs
activated, have no refractory period, show
Transmission at Inhibitory Synapses spatial and temporal summation, and decay
Some synaptic inputs have inhibitory instead of over time and distance.
excitatory effects. In postsynaptic inhibition, IPSPs have an effect antagonistic to that of
the response of the postsynaptic membrane EPSPs. While a large-enough EPSP will de
consists of an inhibitory postsynaptic potential polarize the membrane to the threshold level
(IPSP). In this case, the transmitter causes a and trigger an action potential, a concurrent
simultaneous increase in membrane permeabil IPSP will tend to counteract this effect and
ity to potassium and chloride ions. Since the keep the membrane potential below threshold
equilibrium potential for potassium ion is -9 0 so that no action potential results (figure 4 -
10).
An overall comparison of excitatory and
inhibitory synaptic transmission is shown in
Figure 4-9. A typical IPSP in a spinal motoneuron, figure 4-11- Both are initiated by action
beginning at a - 7 0 mV resting potential and moving
m a hyperpolarizmg direction. Inhibitory input occurs potentials (APs) arriving at presynaptic termi
at time indicated by arrow. nals. When the excitatory transmitter com-
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T h resh o ld
R e stin g Potential
67 Synaptic Transmission
Figure 4-10. Above-threshold EPSP (upper trace), potential) is indicated by a one, and its absence
and IPSP (lower trace), triggered at time indicated is indicated by a zero.
by arrow. Dashed line: when IPSP is triggered si
multaneously with EPSP, depolarization is below The columns under I illustrate the summa
threshold. tion of two excitatory inputs to produce an
action potential. An excitatory input signal in
A alone or B alone does not produce an output
bines with postsynaptic receptors, it produces a signal in C; only when both excitatory inputs A
localized depolarization (the EPSP) via an and B are active (as shown in the bottom row)
increased permeability to sodium and potassium does an output signal, equivalent to an action
ions. In the postsynaptic neuron, the EPSP, if potential, appear in C.
large enough to cross threshold, initiates an The columns under II illustrate the ability
action potential that is then conducted along of an inhibitory input to prevent an output
the axon as an output signal. A simultaneous signal. An excitatory input in A alone, equiva
IPSP, however, can prevent this by driving the lent to a large EPSP, and the absence of an
membrane potential away from the threshold inhibitory input in B, produce an output signal
level. in C (as shown in the second row). But an
inhibitory input in B alone (third row) or
together with an excitatory input in A (fourth
row) produces no output signal in C.
Electronic Logic Circuits
as Models for Synaptic Integration
Electronic logic circuits are important compo
Effect of Synaptic Location
nents of electronic computers and illustrate
some basic principles of synaptic integration The previous discussion has assumed nearly the
(like any other model, however, their resem same location for excitatory and inhibitory
blance to the real thing is quite incomplete). synapses on the postsynaptic neuron. In reality,
Consider simple logic circuits with two inputs, synapses are located at quite different sites
A and B, and one output, C (figure 4 -1 2 ). In along the cell body and dendritic tree, and
the columns on the right side of the figure, the these locations affect synaptic interaction. For
presence of a signal (equivalent to an action example, the model of a dendrite or dendritic
68 Synaptic Transmission
Figure 4 -1 1. Excitatory and inhibitory synaptic 1980. Copyright © 1980, Macmillan Publishing
transmission compared. (I) The action potential Co., Inc. Modified from Eccles, ]. The Under
(AP), consisting of a reversal of membrane polariza standing of the Brain. New York: McGraw-Hill,
tion, arrives at the presynaptic terminal and causes 1973; and from Katz, B., Nerve, Muscle, and
release of excitatory (upper half of diagram) or in Synapse. New York: McGraw-Hill, 1976.
hibitory (lower half) transmitter. E, = internal po
tential; dashed line = zero potential; Ex — Figure 4-12. Electronic logic circuits as models of
excitatory; In = inhibitory. (2) Combination of the synaptic integration in a simplified neuron. Inputs A
excitatory transmitter with postsynaptic receptors pro
and B may be compared with presynaptic terminals,
duces a localized depolarization, the EPSP, through
output C may be compared with axon. In tables on
an increased permeability to sodium and potassium
right, 1 indicates a signal and 0 indicates no signal
ions. The inhibitory transmitter produces a localized
hyperpolarization, the IPSP, through an increased
permeability to potassium and chloride ions. (3) The
EPSP, if large enough (summation of several excita
tory inputs is often necessary), initiates a conducted
Ap in the postsynaptic neuron. However, this can
be prevented by a concurrent IPSP. Reprinted with
permission from Gilman, A.G., Goodman, L.S.,
and Gilman, A. The Pharmacological Basis of
Therapeutics (6th ed.). New York: Macmillan,
69 Synaptic Transmission
i e
QQQQ® 6
Figwre 4 - 13. Dependence of synaptic integration on reduced by about 60% of its control value.
location of synaptic inputs along a dendrite, after Finally, the inhibitory input is most effective
Rail [29]. Above: Numbered circles represent points when located proximal to the excitatory input
along a dendrite, with I being at cell body and 5
being most distant. An excitatory synapse (e) is (C)— the recorded EPSP is now reduced to
located in the middle in all cases, while an inhibitory only 40% of its control value. The overall
synapse (i) is located more distandy (A), at the conclusion is that inhibitory synapses are more
same site (B), or closer (C) to cell body. Below: effective (with respect to control over the cell
Resulting synaptic potentials recorded at the cell body body) when they are closer to the cell body.
for these three cases. Dotted line represents response
to excitatory synapse alone. Source: Shepherd, G.M. The potential of the cell body, and the nearby
The Synaptic Organization of the Brain (2nd initial segment, is important to the initiation of
edL). New York: Oxford University Press, 1979. nerve impulses as output signals from the
neuron.
HCHpVCH-CHj-NHj
9 PHi
o ^ -o -C H r C H -N ^ a ^ H O -U o H
CH3
NOREPINEPHRINE
ACETYLCHOLINE
EPINEPHRINE
DOR^MINE
HO—
r^^Tj—jrCHj-CHj-NH, NH3-CHr CHr CH2-C-OH
GABA
SEROTONIN
nh2-c h 2-c -o h
GLYCINE
MET-ENKEPHALIN
A C ETY LC H O LIN E (A C hl A S A
T R A N S M IT T E R
A cetyl
CoA
r
+ C h olin e
S Y N T H E S IS
P R E S Y N A P T IC ACh
CELL
STO RA G E
e s ic le
A ction Potential
R E L E A SE
D IFF U S IO N f
A c e tic a . + C holine
^^ ^ Jcholinesterasel
S y n a p tic c le ft
BREAKDO W N
D IFF U S IO N
' R e c e p to r
P O S T S Y N A P T IC
CELL
Figure 4-16. Acetylcholine (ACh) as a transmitter. sympathetic system (for example, sym
Presynaptic action potential, m the presence of cal pathetic cholinergic fibers, which innervate
cium, triggers release of transmitter, which diffuses sweat glands)
across and combines with postsynaptic receptor sites.
Figure 4-17. Distribution of transmitters acetylcholine those arising in certain brainstem regions
(ACh), norepinephrine (NE), and epinephrine (E). (locus ceruleus and reticular formation)
D opa Decarboxylase
Drug Action at Synapses
' r Synaptic transmission can be modified with
D O P A M IN E appropriate compounds that mimic or affect the
action of native chemical transmitters. Such
D op am ine
compounds can be either drugs or poisons (as
3 - Hydroxylase we have seen for curare and related com
i r pounds, which block the acetylcholine recep
tor). There are several specific mechanisms for
NOREPINEPHRINE
such drug effects.
P henylethanolam ine
1. The compound can prevent release of the
N - M ethyltransferase
chemical transmitter from its presynaptic ter
' r
minal Botulinum toxin, which sometimes
EPINEPHRINE appears in improperly prepared canned
goods, blocks the release of acetylcholine,
Figure 4-18. Synthesis of norepinephrine and. causing neuromuscular paralysis.
epinephrine. 2. The compound can block the postsynaptic recep
tor site by binding to it and competing u/ith the
originating in particular regions of the brainstem transmitter. Curare, tubocurarine, and alpha-
(the midline or raphd nuclei). It has been tenta bungarotoxin block the acetylcholine recep
tively linked with drugs that affect mood and tors in skeletal muscle. Another drug,
cause hallucinations (e.g., LSD). atropine, blocks the acetylcholine receptors
in smooth muscle and glands innervated by
parasympathetic nerve fibers. This is the
GABA and Glycine
reason atropine eyedrops dilate the pupils:
Certain amino acids, in particular gamma- they block the pupillary constriction ordi
aminobutyric acid (GABA) and glycine, are narily caused by acetylcholine acting on the
found in high concentrations in the CNS and smooth muscle of the iris.
are effective in modifying neuronal discharge. 3. The compound can block enzymatic breakdown
While not yet conclusive, experimental evi o f the chemical transmitter. For example,
75 Synaptic Transmission
NOREPINEPHRINE (NE) AS A
TRANSMITTER
-S Y N T H E S IS
PRESYNAPTIC
CELL
Action potential
Inactive metabolite
II
POSTSYNAPTIC
CELL
Figure 4-19. Norepinephrine (NE) as a neurotrans- The effect of a drug depends on which
nutter. DA = dopamine.
chemical transmitter system it affects and on
the mechanism by which it works. There are,
for example, drugs that block the receptor site
neostigmine blocks acetylcholinesterase and for acetylcholine in striated muscle, others that
is, therefore, called an anticholinesterase drug. block the receptor site for acetylcholine in
Since the acetylcholine no longer breaks smooth muscle, others that block the receptor
down rapidly, it builds up to higher levels in site for norepinephrine in smooth muscle, and
the synapse. In the disease myasthenia gravis, others that block the receptor site for dopamine
weakness due to deficient cholinergic activ in the brain. Obviously, these drugs can have
ity may be counteracted by administering radically different effects. In addition, both
such anticholinesterase drugs. curare and neostigmine work on acetylcholine
4. The compound can block the re-uptake o f the transmission on skeletal muscle, but since they
transmitter. For example, the tricyclic anti- work by different mechanisms, they have differ
depressant drugs, such as imipramine and ent effects: curare blocks the action of the trans
amitriptyline, are thought to block the re mitter, while neostigmine can enhance it.
uptake of norepinephrine into presynaptic
terminals. This allows more norepinephrine
to accumulate in the synapse and coun
teracts depressive symptoms that, according
to one theory, are due to norepinephrine
deficiency.
76 Synaptic Transmission
The amino acid, glutamate, appears to be the main excitatory neurotransmitter in the brain,
and the excitatory neurotransmitter released from primary afferent neurons in the spinal
cord. There are several types of glutamate receptors. The major excitatory action of
glutamate on motoneurons is due to binding to two types of glutamate receptors, the
kainate and quisqualate A receptors, named for their agonists. Both bind to the glutamate
agonist AMPA (alpha-amino-3-hydroxy-5-methyl-4-isoxazole prioprionic acid). These
AMPA receptors seem to be responsible for the EPSP in motoneurons triggered by la
afferent fibers.
Another type of glutamate receptor is the NMDA receptor, selectively activated by N-
methyl-D-aspartate. This receptor controls a cation channel which is permeable to Ca as
well as to Na and K, and which is normally obstructed by extracellular Mg. Glutamate-
activated conductance increase can occur only with adequate depolarization to force Mg
out
At resting membrane potentials, EPSPs depend mostly on the non-NMDA receptors. But
with increased depolarization by such receptors, more contribution can come from the
NMDA receptors. The NMDA receptors contribute a delayed, late phase
to the EPSP.
GABA (gamma-aminobutyric acid) is a major inhibitory transmitter in the CNS,
producing IPSPs in spinal cord and brain. Benzodiazepine drugs such as Valium
(diazepam) are anti-anxiety agents which enhance the inhibitory action of GABA.
Benzodiazepines bind to a site on the GABA receptor.
Two types of synapses which differ in their ultrastructure also differ in their function.
Gray type I synapses, typically, have rounded vesicles, are excitatory, and secrete
glutamate. Type II vesicles, typically, have flattened vesicles, are inhibitory, and secrete
GABA. While type I synapses often are located on dendrites, type II synapses often are
located on the cell body, where they tend to have more effect on the amount of
depolarization at the initial segment and thus on the response of the neuron.
77 Synaptic Transmission
Chapter 4
Synaptic Transmission
Postsynaptic Potential Action Potential
AO K i©
Synapse
Dendrites
Axon
Excitatory postsynaptic potentials (left), due to opening of chemically-gated channels, can lead to an action potential (right), due to
opening of voltage-gated channels, if depolarization is large enough to cross threshold. IS =- initial segment
79 Synaptic Transmission
Review Questions
3. Define end-plate potential (EPP) and miniature end-plate potential (MEPP) and
describe their relationship.
4. Describe the changes in membrane potential and ionic conductance during neu
romuscular transmission.
5. State the relationship between the EPP and the muscle action potential. Define
threshold in this context.
7. Specify a value for synaptic delay, and describe the basis for it
8. State the relationship of EPSPs and IPSPs to the initiation of an action potential in
the postsynaptic neuron.
9. Describe the part of the motoneuron with the lowest threshold. State the
approximate value of die threshold.
10. State in what ways postsynaptic potentials differ from action potentials.
82 Synaptic Transmission
13. State the transmitters at the following synapses: (1) neuromuscular junction
(skeletal muscle), (2) autonomic ganglia, (3) sympathetic effectors, and (4)
parasympathetic effectors.
15. List four mechanisms by which drugs can act at synapses, and classify each as to
whether it increases (potentiates) of decreases the postsynaptic action of the trans
mitter involved.
msec
87 Somatic Sensation
Discriminative Somatic Sensation and 5. Position sense. Move the subject’s arm so that
the Dorsal Column System the angle at the shoulder joint is about 45
Discriminative somatic sensations are those degrees from the vertical. Ask the subject to
that allow you to discriminate a nickel from a duplicate this angle with the other arm.
quarter by touch alone, for example. These Change the angle of the first arm, and ask
sensations involve peripheral afferent fibers the subject to duplicate the new angle.
belonging to Group II and a CNS pathway
including the dorsal columns of the spinal cord. PeripheraL-Nerve Activity
Many of the skin and joint mechanoreceptors
Tests of Discriminative Somatic Sensation
with specialized endings, such as the pacinian
Several tests are described briefly below. All corpuscle, Meissner’s corpuscle, Merkel’s disc,
should be done with the subject’s eyes closed. hair-follicle receptor, and joint receptor, have
Similar tests are included in the neurological large-myelinated Group II fibers. The responses
examination. of such afferent fibers have been described in
chapter 3.
1. Touch. With a small, fine brush, lightly Stimulation of these peripheral nerve fibers
touch the skin over different areas of the by a mild electrical stimulus produces a sensa
body (fingers, palms, arms, back, and so tion that appears to originate in the peripheral
on). Ask the subject to say when he feels the distribution of the nerve. Thus, a mild electri
touch and where. cal shock to the median nerve at the wrist can
2. TwO'point tactile discrimination. This is the give a tingle, tap, or slaplike feeling to the area
ability to discriminate two closely spaced of the hand innervated by the nerve.
touch stimuli from a single one. Two points The large-myelinated Group II afferents can
(pencil tips, calipers with tips covered, or a be functionally separated from the small-
specially designed device called an es- myelinated and unmyelinated fibers in several
thesiometer) a few centimeters apart are ways.
lightly touched to the skin alternated with a
single point. The subject says “one” or 1. They can be stimulated by external elec
“two” as the points are felt. The distance trodes at a lower voltage.
between the two points is gradually reduced 2. They can be more rapidly blocked by a
until the subject can no longer report accu tourniquet, which reduces blood flow, caus
rately; this is the two-point tactile thresh ing ischemia. So discriminative touch may
old. Compare this value on the fingers and be blocked while pain sensation remains.
on the upper arm or back. Notice that the 3. They are generally more slowly blocked by
finger threshold is close to the separation of infiltration of local anesthetics, which can,
the dots in Braille, which is 2.5 mm. therefore, block pain sensation while touch
3. Pressure. Have the subject hold his palm sensation remains.
upward and put different weights on his
palm, one after the other. Have the subject
Central Nervous System Activity
say whether each weight is heavier or lighter
than the previous one. Group II fibers enter the spinal cord through
4. Vibratory sense. Strike a tuning fork so that it the dorsal roots, and then ascend on the same
vibrates, and place its base on different parts side of the cord in the dorsal columns, which
of the body. Ask the subject to indicate can be seen on the dorsal surface of the cord
when and where the vibration is felt. (figure 5 -4 ). The dorsal columns are composed
89 Somatic Sensation
CEREBRAL HEMISPHERES
AVUEG
Thalamus
(VPL nucleus)
Medial lemniscus
MIDBRAIN
MEDULLA
RECEPTORS:
■Touch
Dorsal root' ■Pressure
■Vibration
.•Joint position
SPINAL CORD
■Muscle and tendon receptors
Group I
fibers
Figure 5-4. The dorsal column-medial lemniscal Nerve fibers in the medial lemniscus synapse
(DC-ML) pathway for somatic sensation. D = in a thalamic relay nucleus, the ventral post
dorsal; V = ventraL
erolateral or VPL nucleus. Fibers from the VPL
nucleus then project to the cerebral cortex,
o f the cuneate fasciculus (from the arms) and
mainly to somatic sensory area I in the post
the gracile fasciculus (from the legs). They
central gyrus of the parietal lobe. Because of
appear white because they are composed of
the crossing over in the medulla, the left half of
myelinated nerve fibers. The fibers ascend and
the body is represented in the right postcentral
synapse onto neurons in the dorsal-column
gyrus, and the right half of the body is repre
nuclei (the cuneate and gracile nuclei) in the
sented on the left postcentral gyrus (a contralat
base of the medulla just above the spinal cord.
eral representation).
These neurons send fibers across the midline to
the opposite side of the medulla, where they
continue upward as the medial lemniscus. Thus, Somatotopic Mapping. On the postcentral gyrus,
this system is often called the dorsal column the contralateral half of the body is represented
system or the dorsal column-medial lemniscal by a somatotopic map, very much the way the
(DC-ML) system. world can be mapped on a sheet of paper. The
90 Somatic Sensation
body is mapped upside down, with the face respond. For example, a sensory nerve fiber
below and the legs above. The cortical area for may innervate a particular area of skin in the
each body part reflects density of innervation hand. An adequate stimulus within this area
and sensory function rather than skin area; will cause excitation (increase in firing rate) of
thus, the hands and mouth (which have a high the nerve fiber. This area is called the excitatory
degree of discriminative touch sensation as receptive field o f the nerve fiber (figure 5—5).
shown by the two-point tactile test) have large The size of the receptive field varies widely,
representations, while the back has a small from very small on the fingertips (a few square
one. This mapping has been demonstrated millimeters) to large on the back (several
physiologically by recording the electrical re square centimeters). This size corresponds log
sponses of the cortex to touch stimulation of ically with the two-point tactile threshold on
different parts of the body. The electrical different parts of the body. If the receptive field
response, or evoked potential, to stimulation of of a fiber innervating the fingertips were
the right hand is at a point about 7 cm to the large— 2 cm square— it would cover the whole
left of the midline. The evoked potential to fingertip, and two points touching the fingertip
stimulation of the legs is best recorded from the would both activate the same nerve fiber, so it
part of the postcentral gyrus located on the would be impossible to differentiate the two
medial surface of the hemisphere. points from a single point applied anywhere on
the fingertip. Since the two-point threshold for
the fingertips is actually 2 to 3 mm, the
Receptive Fields and Surround Inhibition. The
diameter of the receptive fields on the finger
receptive field of a neuron in the somatosensory
tips must generally be less, so each point can
system is a particular area of skin within which
an adequate stimulus causes the neuron to excite a different fiber.
Fibers in the median nerve ascend in the
cuneate fasciculus (part of the dorsal column)
Figure 5 -5 . The excitatory receptive field of an affer
ent fiber (a first-order neuron in the somatic sen and then synapse onto second-order fibers in
sory pathway). the cuneate nucleus. A microelectrode in the
STIMULUS
91 Somatic Sensation
cuneate nucleus could record activity from by inhibited neurons a' and c'. In reality, this
these second'Order fibers. They too have recep diagram could be extended above and below
tive fields in the arm and hand. Unlike the the plane of the paper, and neuron b' would be
first-order sensory fibers, however, their ex ringed on all sides with inhibited neurons. This
citatory receptive fields are surrounded by contrast between an activated neuron and its
inhibitory areas, or inhibitory surrounds, stimu inhibited neighbors is considered important for
lation of which decreases the firing rate of the precise localization of where the stimulus is and
nerve cells. the ability to differentiate between two closely
The mechanism of surround inhibition (also spaced stimuli, as in two-point tactile dis
called lateral inhibition) is diagramed in figure 5 - crimination.
6. A table edge is stimulating neuron b, a first- The diagram illustrates two synaptic mecha
order sensory fiber that ascends in the cuneate nisms for surround inhibition:
fasciculus, by pressing on its excitatory recep
tive field. Neuron b synapses directly with 1. Postsynaptic inhibition. A branch (axon collat
second-order neuron b' in the cuneate nucleus. eral) of axon b' synapses with an inhibitory
Neuron b' is also activated by pressure from the intemeuron. Two such inhibitory inter
table edge in its excitatory receptive field. Its neurons then synapse onto neighboring
neighbors, neurons a' and c', however, have neurons a' and c', and inhibit them via
the table edge pressing their inhibitory sur postsynaptic inhibition (generation of an
rounds, and respond with a decrease in firing IPSP and reduction of firing rate).
rate (inhibition) below their ongoing spontane 2. Presynaptic inhibition. A branch of first-order
ous rate. These firing rates are graphed on the fiber b synapses with an intemeuron, which
right of the diagram. Notice that the excited synapses with the axon terminal of neigh
neuron (neuron b') is surrounded on both sides boring fiber c. This axoaxonal synapse (from
one axon terminal to another) is able to
Figure 5-6. A schematic representation of surround depress the release of transmitter from nerve
inhibition in the somatic sensory system. fiber c, causing less activation of neuron c'.
SK IN
-W -U -
ISTIM. |_
R e ce p tiv e field ----------
T IM E
Peripheral n erv e fib e r
M edial lem niscus
since in most people the left hemisphere has If you stub your toe, you will probably feel
dominant importance in language. two phases of pain: an immediate sharp sensa
tion, followed a short interval later by a more-
generalized ache. These two phases are
Pain Sensation and the Spinothalamic attributed to two sets of peripheral-nerve fibers
System conducting impulses at different speeds.
Pain and temperature sensation are carried 1. Fast pain is the more-immediate, sharp, and
from peripheral receptors (nociceptors for pain, localized sensation. It is
thermoreceptors for temperature) by small- a. Carried by A-delta fibers, conducting at a
myelinated (Group III or Type A-delta) and velocity of about 25 m/sec and capable of
unmyelinated (Group IV or Type C) fibers in activating the brain in a fraction of a
the peripheral nerve. In the spinal cord, these second
fibers transmit activity to fibers that ascend in b. A sharp sensation, resembling a pinprick
the lateral spinothalamic tract to several areas c. Localized to the area stimulated
of the brain. For brevity we will refer here only d. Brief
to pain sensation, and we will call the whole 2. Slow pain is the delayed ache. It is
CNS pathway the spinothalamic system. a. Carried by C fibers, conducting at a
velocity of about 2 m/sec and capable of
Peripheral-Nerve Activity activating the brain after a delay of a
second or more
Some of the peripheral stimuli that cause pain b. An aching or burning sensation, more
are: unpleasant than fast pain
c. Diffuse, tending to spread from the im
1. Intense pressure on the skin, cutting, or mediate area stimulated
other tissue damage. Tissue damage can d. Sustained, often outlasting the stimulus
release a polypeptide, such as bradykinin, for some time
that stimulates nociceptors.
2. Sustained muscle contraction that causes
Central Nervous System Activity
ischemia (lack of blood supply) to the
muscle. The lateral spinothalamic tract and its origins and
3. Other causes of ischemia to a muscle, for projections are schematically diagramed in fig
example, pain arising in the heart (angina ure 5 -8 . A-delta and C fibers from nocicep
pectoris) when a coronary artery is blocked. tors may go up or down a few segments after
4. Stretch of a visceral organ, for example, they enter the spinal cord and terminate in the
stretch of the temporal artery associated dorsal horn of the cord. After one or more
with vasodilation in migraine headache or synapses, activity is relayed by dorsal-horn
distention of the ureter when it is blocked by neurons which send axons across the anterior
a kidney stone. commissure to ascend in the lateral spino
5. An abnormal stimulus along the course of a thalamic tract of the opposite side. These
peripheral nerve or its dorsal root causing spinothalamic fibers terminate in several areas
pain that seems to originate in the periph of the brain:
eral region innervated by the nerve. An
example is pain due to a damaged interver- 1. In the thalamus: the VPL nucleus, and a
tebral disc (a so-called slipped disc) which region posterior to it called the posterior
exerts pressure on the dorsal root. nucleus (or nucleus ventralis caudalis)
94 Somatic Sensation
CEREBRAL HEMISPHERES
s .g . = S u b s t a n t i a g e l a t i n o s a
Figure 5-8. The spinothalamic system for pain and fields, sometimes encompassing an entire arm
temperature sensation. The VPL nucleus projects or leg, and a response that well outlasts the
specifically to the somatic-sensory cortex, while the
stimulus (afterdischarge); these characteristics
nonspecific nucleus projects to diffuse areas of cortex,
as indicated by arrows. may be related to the diffuse and sustained
nature of slow pain. Lesions in parts of this
pathway (e.g., the lateral spinothalamic tract
2. Also in the thalamus: the nonspecific nuclei surgically sectioned in human patients as a
(intralaminal and medial thalamic nuclei). treatment for pain, and lesions in the reticular
Neurons in thalamic areas (1) and (2) formation in animals) have been followed by a
project to the cerebral cortex. lack of responsiveness to pain stimuli. How
3. In the brainstem: the reticular formation, a ever, such lesions by no means always abolish
column of neurons in the central core of the pain. Furthermore, there are many complex
brainstem. pain phenomena that cannot be predicted by
considering the anatomical pathway alone;
Except for the VPL nucleus, many of the these phenomena include referred pain and
neurons in these areas have large receptive gating.
95 Somatic Sensation
Referred pain is pain originating in one which are too mild to cause pain sensations
peripheral site that produces a sensation as if themselves but can activate Group II affe
coming from another site, presumably due to rent fibers.
the nature of the CNS projections. Pain signals
from the viscera are often felt at superficial sites This interaction could work by means of
inhibitory intemeurons, excited by the large-
that share the same or nearby spinal segments.
For example, cardiac pain is often referred to myelinated afferents in the dorsal horn of the
cord, which then inhibit synaptic transmission
the chest and the inside of the left arm, areas
supplied by the same or neighboring dorsal (closing the gate) from the A-delta and C fibers
roots of the thoracic cord (the heart is supplied to the dorsal-horn cells that give rise to the
by segments T1 through T5). This phenome lateral spinothalamic tract.
non is often attributed to convergence of There is also evidence that a similar inhibi
afferent fibers from the heart and from the skin tion in the dorsal hom could originate in either
onto the same neurons in the dorsal horn of the the somatic sensory cortex or the brainstem
cord, which then project upward via the lateral reticular formation, with descending activity
spinothalamic tract (figure 5 -9 ). carried by the reticulospinal and other tracts.
Gating is a mechanism by which pain signals
within the spinal cord can be suppressed by
Substance P as a transmitter in the
activity in the large-myelinated Group II (A- nociceptive pathway
beta) fibers due to touch, pressure, or electrical
stimuli. For example: In the dorsal hom of die spinal cord, both
A-delta and C fiber nociceptive afferents
1. Moving water and massage have been used contain small, clear vesicles that are
to suppress pathological pain, as in the thought to contain glutamate or other
neurological disorder causalgia. amino adds, which elidt fast EPSPs in dor
2. Pain arising from some peripheral site can sal hom neurons. In addition, C fibers, and
sometimes be suppressed either by mild possibly A-delta fibers, release neuropep
electrical stimulation of nearby skin or of a tides, induding Substance P. Substance P
peripheral nerve supplying the area, both of elitits slow EPSPs in dorsal hom neurons
that mimic the effects of high-intensity,
painful stimulation of nodceptors.
Figure 5-9. Convergence as a possible mechanism
A specific type of opiate receptor, the mu
for referred pain originating in the heart. Since the
dorsal horn neuron shown usually signals path from receptor, is located on the terminals of noci
the left arm, the same interpretation is given when it ceptive afferents in the dorsal hom There is
is activated by cardiac pain. ST .T r = (lateral) pharmacological evidence that opiates may
spinothalamic tract. inhibit the presynaptic release of Substance
P and glutamate, and this may be an impor
tant mechanism of opiate analgesia. Opiate
receptors in the dorsal hom may also act by
postsynaptic mechanisms. Major classes of
opiate receptors are the mu, delta, and kappa
receptors. Morphine is an agonist of the mu
receptor, and high concentrations of these
receptors are found in the dorsal hom and
file periaqueductal gray of the brainstem.
Both regions are involved in the modula
tion of pain responses.
96 Somatic Sensation
Chapter 5
Somatic Sensation: Dorsal Column System
L R
Stimulus
Peripheral Nerve
Group Type Velocity Diameter Function (Modality)
hapters
Schematic diagram of superior view of lateral spinothalamic pathway in spinal cord and brain, and sites at which analgesic
drugs act (top), and peripheral nerve fibers that enter this pathway (bottom). PN = peripheral nerve. Only representative
values o f velocity and diameter are shown in table.
101 Somatic Sensation
Review Questions
1. Describe methods for (1) clinical and (2) experimental measurement of peripher-
al-nerve-conduction velocity.
2. Classify the sensory fibers in peripheral nerves according to their group, alpha
betical type, conduction velocity, and functions.
3. List and briefly describe the functions of the DC-ML system for discriminative
somatic sensation.
4. Define the excitatory receptive field for a neuron in the somatic sensory system.
101 Somatic Sensation
Review Questions
1. Describe methods for (1) clinical and (2) experimental measurement of peripher-
al-nerve-conduction velocity.
2. Classify the sensory fibers in peripheral nerves according to their group, alpha
betical type, conduction velocity, and functions.
3. List and briefly describe the functions of the DC-ML system for discriminative
somatic sensation.
4. Define the excitatory receptive field for a neuron in the somatic sensory system.
102 Somatic Sensation
5. Define surround inhibition for such a neuron. Describe the synaptic mechanisms
involved.
8. List the structural and functional differences between the DC-ML and the lateral
102 Somatic Sensation
5. Define surround inhibition for such a neuron. Describe the synaptic mechanisms
involved.
8. List the structural and functional differences between the DC-ML and the lateral
103 Somatic Sensation
12. List and briefly describe pharmacological and psychophysiological means of pain
suppression.
6. Auditory and Vestibular Function
Sound Waves and the Ear culating the ratio of the pressure oscillations, P
(in dynes/cm2), to a reference value, Po (which
Sound Waves was chosen to be near hearing threshold for a
young person with good hearing). Then the
We all are familiar with wave motion in water:
if you move your hand back and forth underwa logarithm of that ratio is multipled by 20, to
ter, it generates waves that radiate outward yield the sound pressure level, L, in units called
from the point of origin. A loudspeaker cone in decibels (dB):
an audio system generates waves in air in a
similar way (figure 6 -1 ). The air molecules in P
L (dB) = 20 log —
front of the cone are compressed by each Po
forward motion of the cone and decompressed
(rarified) by each backward motion. If the The frequency, F, of a pure tone is the
speaker cone is moving back and forth in a number of cycles per second (expressed as c/s,
simple sine wave pattern, alternating regions of cps) or hem (Hz). Sound frequency corre
high and low air pressure will radiate outward, sponds to what we hear as musical pitch. A
and at any one instant, the air pressure will be a tone of frequency 256 Hz will be heard as
periodic function of distance as shown in figure middle C. Increased frequency corresponds to a
6 -1 . These pressure variations are called sound higher-pitched sound. Doubling the frequency
waves and are propagated at a velocity of 344 from 256 Hz to 512 Hz will generate a tone one
m/sec. octave above middle C. Another way of
A microphone placed at some fixed distance characterizing the sound is by measuring the
from the loudspeaker will pick up the variations time between recurring peaks— the period T
of sound pressure and convert them to elec (where T = 1/F).
tronic signals, which can be displayed as shown Musical instruments and the human voice
in figure 6 -2 . The simplest sound corresponds often produce complex periodic sounds, which
to a sine wave and is heard as a pure tone; are periodic (they repeat themselves in a
although rare in nature, this can be produced regular period of time) but more complex than
by a soft whistle, a tuning fork, or an electronic sine waves. An example is shown in figure 6 -3 ,
oscillator. A pure tone can be characterized by where a tone of frequency F is combined with
its amplitude and by its frequency. A greater another tone of frequency 2F to produce the
amplitude of movement by the loudspeaker complex periodic sound in line C. The pattern
produces a sound of greater amplitude or inten in line C repeats itself periodically at the lower
sity, which is heard as a louder tone. The frequency F. Such sounds can be produced by
intensity is commonly measured by first cal combinations of frequencies F, 2F, 3F, 4F, and
105
106 Auditory and Vestibular Function
Basilar membrane
Tympanum
Round window
Figure 6 -4 . Outer, middle, and inner ear, showing window, the momentary increase in pressure in
the effect of sound waves at a particular instant in the upper canal causes a downward movement
time. M = malleus (hammer), I = incus (anvil),
of part of the basilar membrane. The extra
S = stapes (stirrup). A compression wave momen
tarily causes an inward movement (shown by dashed pressure transmitted to the lower canal is then
lines) of the tympanic membrane, transmitted via the relieved by a bulging of the round window into
three middle-ear ossicles to oval window, basilar the middle ear (figure 6 -4 ). After its downward
membrane, and round window. Cochlea is viewed as displacement, the basilar membrane then re
if unrolled to form a tube 3.5 cm long, then sec
bounds upward; during a sustained tone, the
tioned longitudinally and simplified as described in
text basilar membrane moves up and down at the
same frequency as that of the tone.
Stroboscopic photographs show that the up-
Cochlear Mechanics
and-down movement of the basilar membrane
Structure o f the Cochlea. Although the cochlea is transmitted as a traveling wave toward the
resembles a spiraling snail’s shell, for simplicity end of the cochlear tube (figure 6 -6 ). (It may
it can be viewed unrolled to form a tube about help to visualize a long elastic band fastened
3.5 cm long (figure 6 -4 ). The tube is filled to the wall at its far end, while your hand
with fluid (perilymph and endolymph) and vibrates the near end up and down and generates
divided lengthwise into two canals by the waves that travel toward the far end). The up-
basilar membrane. The middle ear is separated and-down movement (vertical displacement)
from the upper canal by the oval window and reaches a maximum amplitude at a particular
from the lower canal by the round window distance along the basilar membrane and then
(both windows are membranes). At the far dies away. The distance depends on the fre
end, both canals are connected by a small quency of the stimulus tone:
opening called the helicotrema. The basilar
Frequency Distance at Maximum Amplitude
membrane supports the organ of Corti, a
sensory structure containing auditory receptors high short (near oval window)
(hair cells) and auditory nerve endings. moderate medium
low long (near helicotrema)
Traveling Waves in the Cochlea. When a tone A low tone involves a longer stretch of basilar
causes the stapes to push inward on the oval membrane than a high tone. The envelopes
109 Auditory and Vestibular Function
Figure 6-5. Schematic illustration of area difference brane that contacts the tips of the sensory hairs
between tympanum and footplate of stapes, and the participate in this motion. The basilar mem
accompanying pressure amplification between the two
brane and the tectorial membrane act as if
structures. The area ratio shown (14) illustrates
the principle of pressure amplification, but varying independently hinged at one wall of the coch-
estimates of this ratio have been given.
Figure 6-6. Traveling waves along the basilar mem
brane shown at (A) one instant of time, in a three-
(outer bounds) of traveling waves for some dimensional representation, and (B) at four
frequencies are shown in figure 6 -7 . This successive instants of time, from left to rigfit, with
mechanical behavior of the basilar membrane is basilar membrane represented only as a line corres
largely due to its structure— narrow and stiff ponding to the heavy midline in A. Dashed lines
show the limits of displacement, or the envelope, of
near the oval window, becoming wider and these waves all along the basilar membrane, and ar
more flexible toward the helicotrema. A tone of rows point to the maximal value of the envelope for
a particular frequency (pitch) will cause maximal the particular stimulus frequency used
displacement at a particular place along the basilar
membrane. This is one aspect of the place theory
of hearing. S ta p e s I Basilar m e m b ra n e I H elicotrem a
Neurophysiology of Audition
Reissner’s m em brane
Tectorial m em brane
Hair cells
ganglioi
Rods of Corti
Scala tynnpani
(perilymph)
A uditory nerve
.Tectorial membrane
Reticular lamina
Hair cells
Hinges'
Basilar membrane
Organ of Corti (schematic)
Figure 6-8. Crosssection of the cochlea. Inset shows Sound Localization. One can easily confirm that
a schematic diagram of the organ of Corti, and illus a person whose eyes are closed can localize the
trates a possible mechanism for stimulation of the origin of a finger-snap, key-ring jingle, or other
hair cells.
sound. The automatic tendency to attend to a
ponding to increased loudness, causes increased sound source has obvious survival value in a
spike frequency for sounds within the neuron’s world where either friends or enemies may be
tuning curve, as shown in figure 6 -1 0 (bottom, hidden in the forest, or where automobiles may
right). Such an increase also adds or recruits appear from either end of a street you are about
new neurons to those that are responding. to cross. There are two major mechanisms
Excitation of auditory nerve fibers is as involved; both are illustrated in figure 6-12 for
sociated with upward motion of the basilar a brief “tap” or “click” sound:
membrane. For a steady tone of low frequency
1. The sound arrives first at the ear nearest the
(below about 2,000 Hz), nerve impulses tend to
source. The time delay between the two ears
occur at a particular phase of the stimulus sine
is estimated by dividing the difference in
wave (figure 6 -1 1 ). Such behavior is called
path length (in meters) by the speed of
phase locking. Although each fiber may not fire
sound (in m/sec).
on each cycle, a group of neighboring fibers will
2. Because the farther ear is partially blocked
tend to reproduce the periodicity of the
by the head, there is an intensity difference
stimulus in the timing of their impulses. There
between the two ears, the sound being less
is evidence that such timing contributes to
intense at the farther ear.
pitch perception for lower frequency sounds.
The phase-locked response of groups of nerve For the neural basis of localization, we focus
fibers and its contribution to pitch perception is on the superior olivary nucleus in the medulla,
described in the volley theory of hearing. the first place where nerve fibers from left and
112 Auditory and Vestibular Function
Figure 6-9 . The auditory pathway (schematic). Only ences also are preferred by different neurons.
representative connections are shown. Thus sound localization can be coded in terms
of which group of neurons in the superior
right cochlear nuclei converge on the same olivary nucleus becomes most active. A similar
cells. These cells receive a binaural input (from coding occurs in cells of the inferior colliculus.
both ears), and respond precisely to particular For low-frequency tones, the time delay
combinations of time or intensity difference. effect involves phase-locked nerve impulses
One neuron, for example, might respond best from left and right cochlear nuclei, similar to
to time delays of 0.4 msec (left ear preceding those shown in figure 6-11 for the auditory
right), corresponding to a sound source a few nerve. Phase-locked nerve impulses from the
degrees of angle to the left of the midline. ear nearest the sound source slightly precede
Another neuron might respond best to a sound those from the other ear, and these time delays
source 90 degrees to the right of the midline, result in the firing of a particular group of cells
and so on. Various left-right intensity differ which receive a binaural input.
113 Auditory and. Vestibular Function
EN V ELO PE O F V IB R A T IO N FO R T O N E S O F :
3 2 0 0 Hz
A U D IT O R Y N E R V E — FIBE R R E S P O N S E S TU N IN G C U R V E
FO R FIBE R b dB SPL
dB SPL
80- u m -
5©
-J
©
3
CO 40-
in
©
0.
■O
c
E
CO 0-
Tone
8 0 0 Hz 1 6 0 0 Hz 3 2 0 0 Hz Freq u e n cy
Figure 6-10. Auditory-nerve-fiber responses (spikes) Auditory Cortex. At the upper end of the
to tones of various frequencies and intensities. Left: auditory pathway, nerve impulses are relayed
Fiber b responds to a very low-mtensity tone (near from the medial geniculate nucleus (in the
0 dB SPL) of 1,600 Hz with a small increase in
spike race (a threshold response). An 800 Hz tone thalamus) to the primary auditory cortex (on
requires much greater intensity (80 dB) to elicit such top of the temporal lobes). The primary audi
a response. Right: Frequency-intensity combinations tory cortex is also called Heschl’s gyrus and
giving threshold responses for fiber b and fiber a. Brodmann’s area 41. It is connected to sur
Each fiber has a unique best frequency, fiber b at rounding auditory association areas (Brod
1600 Hz (arrow). Top: Traveling wave envelopes
mann’s areas 42 and 22).
for 1,600 and 3,200 Hz tones, showing that audi
tory nerve fibers a and b innervate different areas of Neuronal response characteristics are prob
basilar membrane. Microelectrode can be used experi ably more complex in the auditory cortex than
mentally to record spikes. at lower levels of the pathway. Some cortical
114 Auditory and Vestibular Function
Figure 6-11. Phase-locked response of two represen Hearing Sensitivity and Hearing Loss
tative auditory nerve fibers, a and b, to a 200 Hz
tone. In this example, each fiber tends to fire at the Normal Hearing Sensitivity. When the fre
same phase of the sinusoidal stimulus—the peak, as
shown by the dashed lines. A summation of the quency range of hearing is measured with pure
activity of fibers a and b would provide a signal (c) tone stimuli, it ranges from a low of about 20
having the same period—5 msec—as the stimulus. Hz to an upper frequency of 20,000 Hz (figure
6 -1 3 ). This range is for a child with normal
hearing; with increased age, the upper-
neurons do not give a sustained response to ffequency limit is reduced (to about 10,000 Hz
a steady, pure tone [11]. Instead, they respond at age 50). The threshold o f hearing (auditory
more reliably to changes in sound amplitude, threshold) is the lowest sound pressure at which
such as the onset or offset of a sound, or to a tone is just audible; it varies with the
changes in sound frequency (a frequency- frequency of the tone, reaching a minimum
modulated sound). Such changing stimuli are (hearing is most sensitive) between 1,000 and
more similar to the complex sounds that make 4,000 Hz. At 1,000 Hz, the ideal threshold of
up animal communication and human speech. hearing is a sound pressure (measured between
Because of extensive crossing fibers in the upper and lower peaks of the sound wave) of
auditory pathway, Brodmann’s area 41 receives 0.0002 dyne/cm2, which has been established
binaural input (from both ears), although there as the reference pressure (Po) for measurements
is some predominance of input from the con of sound intensity, L, in decibels (dB). For
tralateral ear. Thus destruction of this area on other sound pressures, P, the intensity can then
one side does not cause complete hearing loss be expressed as:
but only partial hearing loss in the contralateral P
ear. Deafness would be caused by bilateral L (dB) = 20 log —, where Po = 0.0002
destruction of Brodmann’s area 41, but this 0 dyne/cm2
condition is rare.
115 Auditory and Vestibular Function
20 20
Brainstem Auditory Evoked Response. Electrical
20 - 'J I 30
.-3 i— - 4 E— t-- responses can be recorded from the brainstem
40
"O auditory pathway with electrodes on the scalp.
50
The stimulus is generally a click presented to
u 60 60
UJ left or right ear via earphones. The electrodes
70
O are metal disks, like those used in recording the
Z BQ 80
£ EEG, attached with conducting paste on the
90
z skin at the top of the head (vertex) and the ear.
100 100
The voltage between the two electrodes is
110
greatly amplified, and the response to 1,000 or
250 500 1000 2000 4000 800C
more clicks is averaged by computer electron
> Bon* eond. ics. With this technique, we can record a
X Air cond.
complex electrical potential originating in the
brainstem auditory pathway (figure 6-15). This
FAR-FIELD RECORDING OF AUDITORY BRAIN STEM RESPONSES
LATENCIES MEASURED IN H U M A N SUBJECTS
P R O P O S E D F U N C T IO N A L -A N A T O M IC A L C O R R E L A T IO N S
CHART 5
EEG SERIES
3 « S 6 7 8
1 [T ; 1 l| 1 } III T J I 1 j
I m GZ 3zn
LATERAL INFE RIO R MEDIAL a u d it o r y
ACO USTIC CO CHLEAR SUPER IO R
NERVE :8ihi NUCLEI OLIVARY LEM NISCUS CO LLIC U LU S G ENICULATE RADIATIONS
(P O N S ) C O R TIC A L)
SUPERIOR
OLIVARY
CO M PLE X
O R G A N OF
CORTI
N U CLEU S OF
LA T E R A L
LEM N ISC US
Figure 6-15. A normal brainstem auditory evoked thousand presentations of the click. Although precise
response to clicks of 60 dB above normal hearing neuronal origin of each wave is uncertain, lesions at
threshold presented at the onset of the trace. Total different levels of auditory pathway diagrammed be
length of trace is 10 msec; normal latencies of num low tend to produce response abnormalities beginning
bered peaks are shown at dotted lines. An upward with indicated waves. Courtesy of E. Grass and J.J.
movement of the trace represents positive potential at Stockord, and Grass Medical Instruments, Quincy,
the vertex electrode, as averaged over several Mass. Reprinted with permission.
117
118 Auditory and Vestibular Function
Figure 6—16. The orientation of the right semicircular lar apparatus, is located next to the cochlea in
canals as seen from above, and the response of the the temporal bone, and its nerve, the vestibular
horizontal canal to rotations. As head rotates to right
(clockwise), there is relative movement of fluid (en- nerve, runs adjacent to the auditory nerve
dolymph) to left (counterclockwise). (together they make up cranial nerve VIII).
A ffe re n t to brainstem
Efferent from brainstem
Posterior brainstem
Testing position
(horizontal canals h o riz o n ta l
Figure 6-17. Physiologic nystagmus produced by in one direction throws them backward in the
rotating the head to the right with the head bent
forward 30 degrees so that the horizontal canals opposite direction.
(HC) are parallel to the floor. Dorsal view of post The vestibular nerve fibers, which innervate
erior brainstem, semicircular canals, and eyes. Rotat the hair cells, have a steady resting discharge
ing head to the right causes endolymph due to inertia rate. Bending the hair cells in one direction
to move to the left, stimulating vestibular nerve fi increases the rate, while bending them in the
bers in the ampulla of the right horizontal canaL
Vestibular nerve fibers enter the brainstem to synapse opposite direction decreases it. Rotating the
in the vestibular nucleus (VN), which projects to head to the right (figure 6-17) excites the right
the nuclei of cranial nerves III, TV, and VI. Effe vestibular nerve fibers. This causes reflex eye
rent output via the cranial nerves to eye muscles movements called physiological nystagmus.
(left lateral rectus, LLR, and right medial rectus, First, the eyes move slowly in a direction
RMR) causes slow movement of eye to the left; a
fast movement (saccade) to the right follows. The opposite to the rotation, as if to compenstate
pontine gaze center (PGC) influences these eye for the head movement. This slow phase o f
movements, and impulses between nuclei are carried nystagmus is due to connections from the
by a fiber bundle, the medial longitudinal fasciculus vestibular nuclei in the medulla to the eye-
(MLF). Source: Gay, A.J., Newman, N.M., Kelt- muscle nuclei (cranial nerve nuclei III, IV, and
ner, J.L , and Stroud, M.H. Eye Movement Dis
orders. St. Louis: The C.V. Mosby Co., 1974.
VI) higher in the brainstem. Second, the eyes
Reprinted with permission. flick back quickly in the same direction as the
120 Auditory and Vestibular Function
rotation. This fast phase of nystagmus appears to utricle are fluid-filled structures containing hair
be due to inputs to the eye-muscle nuclei from celb, and the hairs are imbedded in a gelati
sources in the cerebral cortex or reticular nous structure. If the head b tilted, gravity acts
formation or both. With loss of consciousness, on this structure and causes the hairs to bend,
which lowers cortical and reticular activity, the and a specific pattern of activity b transmitted
fast component of nystagmus is reduced. through vestibular nerve fibers to the brain.
The general stimulus to these sensory organs b
Clmcal Testing linear acceleration, of which gravity b one
form. Other forms would occur during accelera
Clinical testing of the function of the semicir tion or braking of a car, rising and falling in an
cular canals and the vestibular pathways can be
elevator, or any sudden movement of the body
tested in two ways: by rotation, and by thermal forward, backward, up, down, right, or left.
stimulation (the caloric test). Automatic, reflex responses of the neck, trunk,
Rotating the patient in a chair stimulates the
arms, and legs allow a person to maintain
canals on the right and left side of the head at equilibrium during these movements. An ex
the same time. If the head is tilted slightly
ample b the leaning forward of a runner as he
forward, as shown in figure 6-17, the horizon
accelerates during a race. The vestibulospinal
tal canals are parallel to the ground and are
tract b involved in these reflexes.
maximally stimulated. The normal response is
nystagmus and the sensation of spinning. Dur
ing a rotation to the patient’s right, his eyes
move slowly to the left and then quickly to the
right; this back and forth movement is repeated
during the rotation. After the rotation is
stopped, for a short time there should be
nystagmus in the opposite direction. This post
rotatory nystagmus is due to inertia: just as
passengers in a ring rotated clockwise would be
flung clockwise after the rotation stops, so the
sensory hairs in the semicircular canals react in
a similar way.
Thermal stimulation allows one to test the
semicircular canals on the right and left sides
separately. With the head tilted backward so
that the horizontal canab are in a vertical
plane, warm water is squirted into the external
auditory meatus. This warms the endolymph at
the outer edge of the horizontal canal on that
side, and the warmed endolymph rises, flows
around the canal, and bends the sensory hairs.
If the vestibular system is functioning normally,
this unusual stimulus gives rise to nystagmus.
Review Questions
1. Using a diagram of sound waves, define sine waves, amplitude, and frequency.
Note how they correspond to pure tones, loudness, and pitch. Also define har
monics and noise.
4. Prepare a schematic diagram of the ear, indicating the tympanic membrane, ossi
cles, oval window, round window, cochlea, basilar membrane, organ of Cord, and
helicotrema.
6. On die diagram of the ear, indicate where and how transduction occurs.
7. List the major structures of the auditory pathway in order, and group diem
accordingly to major divisions of the brain (brainstem, thalamus, cortex).
124 Auditory and Vestibular Function
4. Prepare a schematic diagram of the ear, indicating the tympanic membrane, ossi
cles, oval window, round window, cochlea, basilar membrane, organ of Cord, and
helicotrema.
6. On die diagram of the ear, indicate where and how transduction occurs.
7. List the major structures of the auditory pathway in order, and group diem
accordingly to major divisions of the brain (brainstem, thalamus, cortex).
125 Auditory and Vestibular Function
8. Describe auditory nerve-fiber responses, using the terms bestfrequency and tuning
curve.
11. Explain why a unilateral lesion in the auditory cortex does not lead to complete
deafness in the contralateral ear.
126 Auditory and Vestibular Function
17. Describe the normal stimulus to the semicircular canals. Relate it to the physio
logical nystagmus.
18. Give two methods for clinical testing of semicircular canal functions.
20. Describe a possible common origin for the following combination of signs and
symptoms. A patient complains of a hearing difficulty in the right ear and a spin
ning sensation and shows a slight nystagmus. A caloric test shows normal
vestibular responses on the left side but reduced responses on the right side. An
audiometric examination shows sensorineural deafness in the right ear. A neuro
logical examination reveals no other abnormalities.
7. Vision
How the eye works and how the visual system converts light
into sensed visual images
We are given tiny distorted upside-down images in convert light into electrochemical potentials.
the eyes, and we see separate solid objects in Therefore, no information concerning the cir
surrounding space. From the patterns o f stimula cle can be sent to the visual area of the brain,
tion on the retinas we perceive the world o f objects, and the circle seems to disappear. This disap
and this is nothing short of a miracle [12]. pearance, like other aspects of vision, was not
due to the visual stimulus alone, but to its
interaction with the eye and brain.
Vision requires a combination of light in the In the first stage of this process, the eye
outside world with the human eye and brain. If focuses light from its visual field onto its back
you would like to try an experiment, look with surface. In doing so the eye resembles a camera
your right eye only (by covering the left with or any other optical system.
your hand) at the X in figure 7 -1 . This diagram
now represents part of the outside world as it
impinges upon your right eye, or the visual field
The Eye as an Optical System
of your right eye. The X, on which you are
fixing your eye (or staring at), is your fixation When the eye is viewed from the front, its most
point. The circle is on the right side of your prominent feature is the doughnut-shaped iris,
visual field; although you are aware of it, it will pigmented gray, blue, green, or brown (figure
be less distinct than the X. Now if you move 7-3 ). The hole in the doughnut is the dark-
the book so that it is about 20 cm from your colored pupil. The pupil is really an adjustable
eye, the circle should seem to disappear. At opening through which light passes into the
that point, it is located in the blind spot of your eye. In front of the iris and pupil is the
right eye. transparent cornea, surrounded by the white
The disappearance of the circle can be sclera with overlying conjunctiva.
explained by considering the perspective of an When the eye is viewed in horizontal section
outside observor looking down on you from (figure 7 -4 ), we can see the path traveled by
above your head and able to see into your eye rays of light from an object in the visual field.
(figure 7 -2 ). The observor would see that the The light rays pass through the transparent
fixation point (the X) is focused on a tiny area cornea, the anterior chamber filled with a clear
called the fovea in back of the eye. The circle, fluid called aqueous humor, the lens, and the
which is to the right of the X in the visual field, transparent, gelatinous vitreous humor, and are
is focused to the left of the X in the eye. When finally focused onto the retina at the posterior
the book is at the correct distance, the circle surface of the eye. Befor entering the lens, the
falls upon a region of the eye at the exit of the rays must pass through the pupillary opening,
optic nerve. This region has no receptor cells to the size of which is controlled by the iris.
129
130 Vision
with a lens that is more curved horizontally accommodation, and requires the lens to change
than vertically. shape— it becomes rounder, or more convex
(figure 7 -8 ). The more convex the lens, the
greater its refractive power. Some intricate
mechanical adjustments are involved in this
Accommodation
process. Left to itself, the lens would be almost
When a photographer wants to take a close-up round when viewed in horizontal section.
picture, he focuses the lens by increasing the However, in the eye, the lens is being flattened
distance between the lens and the film surface. by the pull of suspensory ligaments (zonular
The mammalian eye works on a different fibers) on its outer border (these ligaments form
principle. When an object is moved closer to a ring around the lens as viewed from front or
the eye, the distance between cornea and rear). Contraction of the ciliary muscle releases
retina does not change. Instead, the refractive this tension and allows the lens to assume a
power of the eye’s optical system is increased to more spherical shape due to its own elastic
keep the image in focus. This process is called properties.
The ciliary muscle is controlled by parasym
pathetic fibers from cranial nerve III (oculo
motor), which secrete acetylcholine at the
Figure 7-8. Accommodation of the lens for near vi
sion (right), compared with the unaccommodated eye neuromuscular junction. The accommodation
(left). Arrows indicate contraction of ciliary muscle response involves areas of cerebral cortex and
during accommodation. the superior colliculus. As in other parasym-
D ISTA N T O B JE C T
H ORIZON TAL S E C T IO N O F L E N S
134 Vision
pathetic responses, accommodation can be a well-lit room, you should see both pupils
blocked by the drug atropine. constrict in response to the increased illumi
The capacity of the lens to become more nation.
convex when tension is released is also essential This constriction in response to light is called
for accommodation to occur. With increasing the pupillary light reflex. It can be demonstrated
age, the lens tends to become more rigid, and by asking another person to look ahead in a
accommodation is impaired. This condition is moderately lit room and then shining a pen-
called presbyopia. It may be partly compensated light into the eye. The pupillary constriction
for with bifocal lenses, in which the lower part that results is an automatic reflex response that
of the spectacle lens is used for near vision. protects the retina against too-intense light,
just as a small iris-diaphragm protects film
against overexposure. The effector muscles are
Adjustment o f Pupil Size
the sphincter (circular) muscles of the iris,
The amount of light entering the lens is which receive a parasympathetic, cholinergic
determined by the size of the pupil, which in innervation from cranial nerve III (oculo
turn is controlled by the surrounding iris. The motor). The pupillary light reflex pathway
iris-diaphragm in a camera performs a similar involves the following:
function: in bright sunshine it may be adjusted
light —» optic nerve tract —* pretectal
to provide an opening as small as a pinhead
nucleus —►E-W nucleus —* cranial nerve III —*•
(signified as f-stop f-22), while in dim indoor
sphincter muscle of iris —* constriction
light it may be opened wide (f-2.8) to allow
enough light to enter the camera to provide where E-W stands for the Edinger-Westphal or
adequate exposure for the film. If you look at parasympathetic nucleus of cranial nerve III,
people’s eyes in a wide range of lighting located, along with the pretectal nucleus, in
conditions, you may find pupillary diameters the midbrain. The time required to traverse
ranging from about 2 to 8 mm (figure 7-9). this pathway, from light onset to pupillary
Although other influences are also involved, constriction, is approximately 200 msec. In the
brighter light causes smaller pupils. You can try pupillary light reflex, both pupils constrict even
a simple test on yourself to demonstrate the though only one may be stimulated. This is due
pupil’s response to a change in illumination. If to fibers that cross over in the posterior com
you close your eyes for about 25 seconds and missure, from pretectal nucleus on one side to
then open them while looking into a mirror in E-W nucleus on the other. Pupillary constric
tion is also called miosis.
Figure 7-9. The approximate range of pupil diameter. Active dilation of the pupil is caused by
135 Vision
contraction of the dilator (radial) muscles of buildup of acetylcholine, which also activates
the iris, which receive innervation from sym the sphincter muscles and causes pupillary
pathetic fibers that secrete norepinephrine. Via constriction.
this pathway strong emotions like fear or You may have noticed that cholinergic fibers
excitement can cause pupillary dilation. from cranial nerve III cause two effects on the
The balance between the opposing cholin eye: accommodation for near vision and pupil
ergic and adrenergic effects on the iris explains lary constriction. During an eye examination,
how many drugs can affect pupil size (figure 7 - the atropine that blocks pupillary constriction
10). Atropine or similar substances given in (and leads to a dilated pupil) will block accom
eyedrops block acetylcholine receptors in the modation as well. The patient will have diffi
sphincter muscles of the iris; the unopposed culty seeing clearly, especially nearby objects,
dilator muscles then cause pupillary dilation, until the effects have worn away. Another
an effect used by ophthalmologists when they occasion in which both effects are evident is
examine the eye. Amphetamine also causes the near response: when a person looks from a
dilation, although by a different mechanism; it distant object to a nearby one, accommodation
enhances the response of the dilator muscles. and pupillary constriction occur automatically.
Acetylcholine-like substances, such as pilocar The pupillary constriction sharpens the image,
pine, activate the sphincter muscles of the iris because light rays are forced to pass only
and cause pupillary constriction. Cholinester- through the center of the lens, reducing the
ase inhibitors (eserine, prostigmine) allow a distorting effects (spherical and chromatic
aberrations) of light passing through the edges.
These aberrations would otherwise tend to
Figure 7-10. Opposing effects of sympathetic (ad
worsen as the lens becomes rounder.
renergic) and parasympathetic (cholinergic) innerva
tion of the iris, and the effect of drugs. From
Westhehner, G. The eye, including central nervous
system control of eye movements. In Mountcastle, The Neurophysiology of Vision
V.B. (Ed.). Medical Physiology (12th ed.). VoL
2. St. Louis: The C.V. Mosby Co., 1968. Re The ability to see light and dark differences,
printed with permission. forms, and details (pattern vision) may, for the
I Noradrenaline I
Sympathetic Parasympathetic
innervation innervation
Dilator Sphincter
135 Vision
contraction of the dilator (radial) muscles of buildup of acetylcholine, which also activates
the iris, which receive innervation from sym the sphincter muscles and causes pupillary
pathetic fibers that secrete norepinephrine. Via constriction.
this pathway strong emotions like fear or You may have noticed that cholinergic fibers
excitement can cause pupillary dilation. from cranial nerve III cause two effects on the
The balance between the opposing cholin eye: accommodation for near vision and pupil
ergic and adrenergic effects on the iris explains lary constriction. During an eye examination,
how many drugs can affect pupil size (figure 7 - the atropine that blocks pupillary constriction
10). Atropine or similar substances given in (and leads to a dilated pupil) will block accom
eyedrops block acetylcholine receptors in the modation as well. The patient will have diffi
sphincter muscles of the iris; the unopposed culty seeing clearly, especially nearby objects,
dilator muscles then cause pupillary dilation, until the effects have worn away. Another
an effect used by ophthalmologists when they occasion in which both effects are evident is
examine the eye. Amphetamine also causes the near response: when a person looks from a
dilation, although by a different mechanism; it distant object to a nearby one, accommodation
enhances the response of the dilator muscles. and pupillary constriction occur automatically.
Acetylcholine-like substances, such as pilocar The pupillary constriction sharpens the image,
pine, activate the sphincter muscles of the iris because light rays are forced to pass only
and cause pupillary constriction. Cholinester- through the center of the lens, reducing the
ase inhibitors (eserine, prostigmine) allow a distorting effects (spherical and chromatic
aberrations) of light passing through the edges.
These aberrations would otherwise tend to
Figure 7-10. Opposing effects of sympathetic (ad
worsen as the lens becomes rounder.
renergic) and parasympathetic (cholinergic) innerva
tion of the iris, and the effect of drugs. From
Westhehner, G. The eye, including central nervous
system control of eye movements. In Mountcastle, The Neurophysiology of Vision
V.B. (Ed.). Medical Physiology (12th ed.). VoL
2. St. Louis: The C.V. Mosby Co., 1968. Re The ability to see light and dark differences,
printed with permission. forms, and details (pattern vision) may, for the
I Noradrenaline I
Sympathetic Parasympathetic
innervation innervation
Dilator Sphincter
136 Vision
Distribution (in retina) Distance from fovea as High concentration in Absent in fovea; predo
measured in degrees of fovea; concentration minant in periphery of
visual angle declines toward periph retina
ery of retina
Light sensitivity Minimum intensity of Low sensitivity High sensitivity (can see
light stimulus causing in dim light after dark
response (sensitivity is adaptation)
the inverse of this
value)
Color sensitivity Wavelength Color vision Black-white vision; also,
insensitive to pure red
light
Visual acuity Minimum visual angle be High acuity (can see fine Low acuity
tween objects (acuity is details)
the inverse)
in the immediate vicinity of the fixation point, half represent the right half of the retina (right
which is focused on the all-cone fovea. As hemiretina). The two optic nerves meet at the
figure 7-15 shows, visual acuity falls off with optic chiasm, where a regrouping of nerve fibers
distance from the fovea in the same manner as occurs. This regrouping is most easily under
does cone concentration. Rod vision is capable stood in terms of the visual field, which can be
of much less acuity than is cone vision. Thus, divided into a left and right half at the fixation
we are much more capable of distinguishing point. For each eye, the left half of the visual
details, such as fine print, in bright light when field is focused onto the right half of the retina.
fixating directly on the object, than in dim The corresponding optic nerve fibers combine
light or in the periphery of the visual field, at the optic chiasm to form the right optic tract.
where rod vision predominates. Thus, at the optic chiasm, uncrossed fibers
In summary, as shown in table 7 -1 , rod and from the right hemiretina of the right eye are
cone vision differ in several characteristics. joined by crossed fibers from the right
Rod vision is capable of high sensitivity to lighthemiretina of the left eye. These fibers carry
(it responds to low light intensity) but cannot information representing the left half of the
distinguish colors (wavelength) and has poor visual field to the visual pathway of the right
visual acuity (it distinguishes between points side of the brain. Similarly, for each eye, the
only if they are separated by a large visual right half of the visual field is focused onto the
angle). Cone vision has only low sensitivity but left half of the retina. The corresponding nerve
is capable of color vision and high acuity in fibers form the left optic tract, which consists of
bright light. uncrossed fibers from the left eye and crossed
fibers from the right eye. These fibers carry
information to the visual pathway of the left
The Visual Pathway in the Central Nervous
side of the brain.
System
The main visual pathway on each side con
In each eye, the axons of the approximately sists of the optic tract, the lateral geniculate nu
one million retinal ganglion cells come to cleus, the optic radiation, and the primary visual
gether to form the optic nerve (figure 7—17). cortex (Brodmann’s area 17). A large lesion in
About half of these cells represent the left half this pathway can cause blindness in the opposite
of the retina (left hemiretina) and the other half of the visual field of both eyes, a condition
142 Vision
termed hemianopia. Nerve fibers in the optic located in layer IV of the primary visual cortex
tract synapse onto cells in the lateral geniculate, near the incoming nerve endings from the
which is a relay nucleus in the thalamus. The lateral geniculate. Other cortical cells, how
fibers from these cells form the optic radiation, ever, have a new type of receptive-field organi
which projects to cells in the primary visual zation. These cells, studied extensively by
cortex, in the occipital lobe. The retinal surface Hubei and Wiesel [17,18], require not only
is mapped onto the visual cortex, just as the that the stimulus be located in a specific area of
body surface is mapped onto the somatic sensory the visual field (as do retinal ganglion cells),
cortex. The heavy concentration of incoming but also that, within that area, excitatory and
nerve fibers from the optic radiation in layer IV inhibitory areas are arranged along a line at a
of the six-layered visual cortex forms a horizon particular angle from the vertical (a particular
tal striation, which gives rise to another name orientation). As shown in figure 7-18, the
for this region, the striate cortex. Outgoing optimal stimulus is a bar of light at an angle
fibers leave the striate cortex for the adjacent that covers the excitatory but not the inhibit
visual association cortex (areas 18 and 19). ory receptive field. Rotation of the bar reduces
the neural response, since both excitatory and
inhibitory areas are activated; a bar that is
Neuron Activity in the Visual Cortex oriented perpendicular to the correct angle
evokes little or no response (figure 7-18D).
The circular organization (center-surround re
These cells may be called orientation-specific
ceptive field) of retinal ganglion cells is pre
edge detectors; Hubei and Wiesel called them
served, with slight change, in the lateral
simple cells to contrast with other, more-
geniculate nucleus and in some of the cells
complex cells in the visual area.
What is the basis for the response specificity
Figure 7-17. Schematic diagram of the visual pathway
as viewed from above the head. T = temporal, N = of the simple cortical cells? One possibility,
nasal half of visual field; L = left, R = right. suggested by Hubei and Wiesel, is that these
143 Vision
a
B Bar at optimal orientation
a
D Bar at 90*angle to
optimal orientation
a LIGHT
Figure 7-18. Receptive field of a neuron in visual Wiesel presented a model of how the receptive
cortex. This receptive field responds maximally to a fields of cortical cells are organized [17,18]. A
vertical bar. + = excitatory area, — = inhibitory
simplified version of this model appears in
area After Hubei and Wiesel [16],
figure 7-20. A microelectrode lowered perpen-
cells receive the convergent input of several
neurons in the lateral geniculate nucleus or in
Figure 7-19. Schematic model explaining receptive
layer IV of the striate cortex, each with circular field of a simple visual cortex cell. Several cells in
receptive fields adjacent to each other along a the lateral geniculate nucleus (LGN), with receptive
single line in the visual field (figure 7-19). The fields lined up vertically in the retina, synapse onto a
compound receptive field composed of the single cortical cell as shown (e = excitatory syn
apse). After Hubei and Wiesel [16]. Source:
adjacent circular receptive fields would have
Shepard, G.M., The Synaptic Organization of the
the orientation specificity of the simple cortical Brain (2nd ed ). New York: Oxford University
cells. Press, 1979. Reprinted with permission.
The more-complex cortical cells are of sev
eral types. Some have orientation specificity
CORTEX
but less location specificity than simple cells— (SIMPLE CELL)
they will respond when the bar or edge is
e
shifted in position over a moderate range. RETINA
Others respond best to a comer at a particular
angle, moving in a particular direction.
We have noted that the visual cortex (like
other primary sensory-cortical areas) may be
divided into six cell layers. After studying the
responses of cells encountered by microelec
trodes passing through all six layers at different
angles from the cortical surface, Hubei and
144 Vision
Hypercolumn
600 « — 180*
Figure 7-20. A model of receptive field organization about 33 p. in monkey cortex. Neurons in the
in visual cortex. 1 to VI represent the layers of adjacent column, labeled column 2, have re
cerebral cortex (I is the surface layer). Short lines
ceptive fields with preferred orientations about
and circles represent receptive fields of neurons in the
cortex as described in text Numerical measurements 10 degrees clockwise from the vertical, neurons
given are representative only. Adapted from D.M. in column 3 have fields with orientations about
Hubei and T.N. WieseL Brain mechanisms of vi 20 degrees clockwise, and so on. With each
sion. Sci. Am. 241:150-162, 1979. column covering about 10 degrees, all possible
angles (180 degrees) could be covered by 18
dicular to the cortical surface might encounter orientation columns, occupying a width of
a number of cells whose receptive field covers about 600 p, in the cortex. The 600 p.-wide
approximately the same area of the visual field. hypercolumn would be able to respond to all
Some cells in layer IV, as we have mentioned, line orientations within a particular receptive
would have circular receptive fields, as indi field, covering a certain number of minutes or
cated by the small circles in figure 7-20. degrees of visual angle. The next hypercolumn,
However, other neurons encountered by the starting with column 19, would represent an
microelectrode would have vertically oriented immediately adjacent area in the visual field.
receptive fields as symbolized by the short Figure 7-21 shows typical relationships be
vertical lines in figure 7-20. These neurons tween two hypercolumns, labeled a and b, and
could be visualized as a slab, or orientation their corresponding receptive fields. Note that
column, of cells with similar response proper the adjacent hypercolumns correspond to adja
ties, and are labeled column 1 in the figure. cent and partially overlapping receptive fields.
The width of the column was found to average The upshot of this model is that within a block
145 Vision
Figure 7-21. Projection of receptive fields (below) ocular vision and produce a shift of the visual
onto hypercolumns of visual cortex (above) in the image especially for nearby objects. With both
model suggested by Hubei and Wiesel [17,18], eyes fixed on the same object, the resulting
binocular vision involves a fusion effect, in
o f cortex approximately 600 p. (one hyper- which the object is perceived as being in a
column) by 2,000 p. (the depth of the cortex) is single location. This binocular fusion results
the neural machinery for analyzing all possible when the image of the object folk on corres
light-dark borders or lines within a given area ponding areas of both left and right retinas.
of the visual field. An adjacent block of cortex Neurons carrying information about these
analyzes an adjacent area of the visual field. two corresponding areas remain essentially sep
Within each block is a similar series of orienta arate all along the visual pathway until the
tion columns. striate cortex. Even in the striate cortex, a
number of neurons are monocular— they have
receptive fields in one eye only. The model in
Binocular Vision
figure 7-21 shows how right or left eye domi
By alternately closing the left and right eyes, nance of neural responses appears to alternate
you can alternate between right and left mon at right angles to the sequence of orientation
146 Vision
Two classes of retinal ganglion cells are the M cells, which project to the magnocellular
(large celled) visual pathway, and the P cells, which project to the parvocellular (small
celled) visual pathway. The M cells have widely branching dendrites and correspondingly
large receptive fields. They are sensitive to large, transient stimuli, and can respond even
at low contrast. The P cells have smaller dendritic fields, respond to fine details, and are
color sensitive. These two cell types form parallel connections with photoreceptors, and
within each class there are both on-center and off-center cells.
These two classes of neurons project to different layers of the lateral geniculate nucleus.
The M cells project to magnocellular layers 1 and 2, and the P cells project to
parvocellular layers 3-6. The M pathway is associated with detection of movement, while
the P pathway is associated with the detection of detail and color.
In addition to the retinotopic map on the primary visual (striate) cortex, there are several
retinotopic maps in areas 18 and 19, and also in the posterior temporal lobe and nearby
parietal lobe. These areas are responsible for parallel processing of different aspects of the
visual image. For example, in area 19 a region called V4 analyzes color and shape, while
in the temporal lobe a region called V5 analyzes movement
Connections between the P (parvocellular) and M (magnocellular) pathways and the
several visual maps in the cortex have been described by researchers. For example, the
magnocellular pathway projects to various layers of area 17, region VI, and then to area
18, regions V2 and V3, and to the temporal lobe, region V5. This pathway specializes in
the movement and location of visual images, but is insensitive to color differences.
In tests with human subjects, the perception of movement appeared to disappear when
stimuli varied in color only and not in brightness. The results suggested that movement is
detected by the magnocellular system. Clinically, bilateral lesions including temporal lobe
region V5 have led to the inability to detect movement, while other visual sensations were
retained. Other types of impairment in visual perception, or visual agnosias, for form,
color, depth, and other attributes, have been associated with lesions of other specific
cortical regions. In addition, dyslexia, or developmental reading disorder, according to
some researchers, may be associated with less response to rapidly changing, low-contrast
stimuli, and a possible impairment in magnocellular system function.
Sources:
Kandel, E.R., Schwartz, J.H., and Jessdl, T.M. Principles o f neural science, 3rd ed. Elsevier, New York, 1991, pp. 445-
449.
Livingstone, M., Rosen, G., Drisland, F., and Galaburda, A. Physiological and anatomical evidence for a magnocellular
defect in developmental dyslexia. Proc. National Academy o f Sciences, USA 88:7943-7947,1991
154 Vision
11. Diagram the visual pathway, identifying optic nerve, optic chiasm, optic tract, lat
eral geniculate nucleus, and cortical area 17. On the diagram, show the projection
of temporal and nasal halves of the visual field onto the retina and the visual cor
tex. Illustrate effects of lesions in an optic nerve, tract, or chiasm on visual fields.
12. Use the term simple cell in describing neuronal activity in the visual cortex.
Describe and diagram the receptive field of such a cell
13. State the current view of the mechanism for color vision.
155 Vision
14. State die current view of the mechanism for binocular vision.
15. Early astronomers discovered that a dim star was best seen if they looked slight*
ly to one side of it rather titan directly at i t How can this be explained on the basis
of (1) the distribution and (2) the light sensitivity of the rods and cones in the reti
na?
16. After receiving atropine eye drops in the left eye only, a patient is asked to focus
on a near object Describe and explain the different responses in the two eyes.
17. A patient is found to have no vision in the left half of both visual fields. Name this
field defect and suggest the site of a possible lesion or dysfunction.
158 Chemical Senses
Pore
Figure 8-1. (A) A taste bud, showing several sen others. Since a single fiber can respond to more
sory receptor cells, Supporting cells are associated than one stimulus quality, the unambiguous
with receptor cells but are not shown. (B) The
quality of, for example, saltiness probably re
neural pathway for taste. Afferent fibers travel m
cranial nerve VII (facial nerve, chorda tympani quires the central nervous system to correlate
branch) and nerve IX (glossopharyngeal nerve, ling- the activity of a number of different fibers.
ual branch). ST = solitary tract, NST = nucleus There is a parallel in the auditory system: since
of the solitary tract, Th. = thalamus (arcuate nu each auditory nerve fiber responds to a range of
cleus), Cx. = cortex (taste area in postcentral gy
frequencies, determining the pitch of a tone
rus). Associated pathways shown with dashed lines:
HN = hypoglossal nucleus, Amyg. = amygdala. presumably requires the correlation of the
activity of a number of fibers to determine
which is responding the most.
bitter substances and is innervated by the
In addition to quality, another characteristic
glossopharyngeal nerve. Thus, a lesion to the
of taste sensation is intensity. Judgments of
glossopharyngeal nerve would reduce sensitiv
stimulus intensity could be based on informa
ity to bitter taste with little effect on the other
tion provided by afferent fibers in the form of
taste qualities.
spike frequency, which is a function of the
The receptor potential of single receptor
concentration of the stimulating chemical.
cells and the action potentials of single afferent
fibers in response to chemical stimuli can be
recorded. In general, each cell or fiber has its Behavioral Responses to Taste Stimuli
own pattern of response, responding maximally Human fetuses can swallow amniotic fluid as
to one or more stimulus qualities and less to early as 12 to 16 weeks gestation. This response
159 Chemical Senses
is evidently influenced by the taste of the fluid, Conversely, sour or bitter solutions lead to a
since with the addition of saccharin as part of a rejection reflex, in which the tongue moves the
medical procedure, swallowing has been found solution out of the mouth.
to increase. Newborn infants demonstrate In animals, behavior tests have been designed
strong taste preferences, which have been to measure the amount of preference or aversion
studied by placing drops of distilled water or a elicited by various solutions [27]. Sucrose and
weak solution of sugar, salt, or other solution sodium chloride elicited preference (drinking)
on the tongue [4]. Sugar solution placed on the in moderate concentration, but aversion in
sweet-sensitive tip of the tongue causes an higher concentrations. The preferred concen
intake reflex: the tip of the tongue curls trations could be altered by deprivation: salt-
upward, and the tongue retracts in such a deprived animals preferred a more concentrated
manner that liquid is moved toward the open solution, an adaptive response in that it would
ing of the pharynx. The tongue movement is help them maintain salt homeostasis.
controlled by the hypoglossal nerve (figure 8 - Such experiments, and others in human
1). In addition, the infant may demonstrate subjects, have led to speculation that, under
associated facial expressions, such as a smile, natural conditions, humans might naturally
that communicate a feeling of enjoyment. prefer the foods they need for proper nutrition.
Once liquid reaches the opening of the If so, such adaptive eating behavior could be
pharynx, it can activate a swallowing trigger threatened by processed and artificial foods. For
zone and thereby elicit a swallowing reflex. example, a breast-fed infont will be encouraged
to drink its mother’s milk by the sweet taste of
the lactose in the milk; at the same time the
Figure 8-2. Distribution on the tongue of sensitivity infant obtains the protein, fot, and other
to the four qualities of taste.
nutritional components of the milk. Later, the
sweet taste of fructose will encourage the child
to eat nutritious, ripe fruit. But if soft drinks
containing sugar are taken with equal zest, no
such nutritional benefits accompany them.
Figure 8-3. The olfactory epithelium, bulb, and region. The axons of the mitral cells form the
tract Left: Location with respect to the nasal cavity.
olfactory tract, a nerve bundle on the underside
Right: Neural connections.
of the frontal lobe, which sends projections to
the olfactory bulb on the opposite side and to
several regions of the brain. These regions
appetizing foods may be approached and include an olfactory area of the cortex, located
odoriferous poisons avoided while there is time on the ventral and medial surfaces of the
to do so. frontal and temporal lobes, and the amygdala.
Review Questions
2. List the four basic qualities of taste, and give examples of each.
3. Describe the behavioral responses to taste stimuli. Explain why they are gener
ally adaptive.
Review Questions
2. List the four basic qualities of taste, and give examples of each.
3. Describe the behavioral responses to taste stimuli. Explain why they are gener
ally adaptive.
Review Questions
2. List the four basic qualities of taste, and give examples of each.
3. Describe the behavioral responses to taste stimuli. Explain why they are gener
ally adaptive.
5. Describe the physiological responses that have been recorded from tire olfactory
system.
6. A patient has lost tire sense of taste on tire anterior part of the tongue; however,
the ability to taste quinine and other bitter substances on the posterior third of
the tongue remains. In what location is a lesion suspected?
164 Chemical Senses
5. Describe the physiological responses that have been recorded from tire olfactory
system.
6. A patient has lost tire sense of taste on tire anterior part of the tongue; however,
the ability to taste quinine and other bitter substances on the posterior third of
the tongue remains. In what location is a lesion suspected?
9. Posture and Movement
A tennis player waiting for the ball typically general, all normal members of the same
assumes a modified standing posture, erect but species respond in the same way to the same
with knees slightly flexed and trunk bent stimulus. Spinal reflexes are those responses
forward at the waist. When the ball ap whose essential neuronal pathways are con
proaches, this posture is dissolved; the player tained within the peripheral nerves and spinal
runs to get into position and swings the cord, although, as we shall see, they may be
racquet. Two elements of motor behavior are subject to descending influence from the brain.
shown in this example: posture (the upright The simplest spinal reflex is the muscle stretch
stance while waiting for the ball) and move reflex; others are the inverse muscle stretch
ment (the run and swing). Motor behavior can reflex and the flexor withdrawal reflex.
also be divided into reflex and supraspinal
components. In the standing posture, for exam
ple, the muscle that extends the lower leg (the Muscle Stretch Reflex
quadriceps femoris muscle, in the front part of
An example of the muscle stretch reflex is the
the thigh) must contract to resist the pull of
Achilles tendon or ankle jerk reflex, as shown
gravity. It can be shown that this contraction is
in figure 9 -1 . Starting with the subject’s leg
largely due to a spinal reflex, an automatic
relaxed, the stimulus (1) is a tap of the Achilles
response to gravity built into the neuronal
tendon provided by a reflex hammer. This
circuitry of the spinal cord. But the strength of
causes an almost immediate stretch (2) of the
this reflex, and the changes that occur when
gastrocnemius muscle in the calf. After a short
the player moves toward the ball and then
delay, the reflex response is (3) a contraction of
swings the racquet, are controlled by motor
the gastrocnemius muscle, causing a plantar
centers in the brain; they are subject to supras
flexion of the foot.
pinal motor control. Such movements are of
The electrical activity associated with the
course not built into every human being (or
muscle contraction can be viewed by placing
else everyone would be a good tennis player);
recording electrodes (metal disks) on the skin
but are matters of will, purpose, and learning.
overlying the muscle and amplifying the elec
They are called voluntary, or purposeful, move
trical potential difference between them. The
ments to distinguish them from purely auto
electrical response recorded in this manner is
matic reflex responses.
called an electromyogram (EMG); it represents a
compound potential due to the action poten
tials of numerous muscle fibers beneath the
Spinal Reflexes
electrodes.
A reflex is stereotyped motor response to a The EMG provides a measurement of the
sensory stimulus. It is stereotyped in that, in time interval, or latent period, between the
165
166 Posture and Movement
Figure 9-1. The ankle jerk reflex, a (phasic) muscle Afferent limb: Distance 1.2 m
stretch reflex. Amp. = amplifier, S = stimulus, R = Velocity 100 m/sec
response, EMG = electromyogram. Conduction
time 1.2 m/100 m/sec)
12 msec
Efferent limb: Distance 1.2 m
stimulus and the response. In the example Velocity 80 m/sec
shown (in figure 9 -1 ), the latent period is 30 Conduction
msec. Most of the latent period can be attrib time 1.2 m/(80 m/sec)
uted to conduction time along the afferent limb 15 msec
of the reflex (the nerve fibers from the muscle- Total conduction time = 27 msec
stretch receptor to the spinal cord) and the
efferent limb (the nerve fibers from the spinal The remaining 3 msec of the latent period is
cord to the muscle). The distance between the composed of additional delays due to excitation
center of the muscle and the cord is approxi of muscle receptors, synaptic delay, and excita
mately 1.2 m. The afferent nerve fibers are tion of muscle fibers.
members of Group la, with velocities of ap The basic phenomenon in the muscle stretch
proximately 100 m/sec. The efferent nerve reflex is that a stretched muscle responds by
fibers are A-alpha motoneurons, with velocities actively contracting. Although in the ankle
of approximately 80 m/sec. The resulting con jerk reflex, the stretch is induced by tapping
duction times can be calculated as follows: the tendon, more common stimuli are gravity
167 Posture and Movement
or other forces during everyday life. Three the upper arm. The resulting reflex contrac
examples are listed below. tion of the same muscle helps to keep the
arm in its original position.
1. During standing, the body tips forward
under the influence of gravity, causing dor A common element in this reflex is that the
sal flexion of the foot relative to the lower muscle that is stretched is itself contracted. In
leg. This stretches the gastrocnemius mus addition, antagonist muscles at the same joint
cle. The resulting reflex contraction of the are automatically relaxed, so that the opposing
same muscle causes plantar flexion of the muscle groups do not fight each other. This
foot, assisting in the maintenance of an relaxation of antagonists is called reciprocal
upright posture. inhibition.
2. The influence of gravity during standing also
Neuronal Mechanism. In the muscle stretch re
causes the knees to bend, stretching the
flex, the basic reflex arc is shown in figure 9 -2 ,
quadriceps femoris muscle in the anterior
consisting of an afferent limb, A, and an
thigh. The resulting reflex contraction of
efferent limb, B (ignore part C for the mo
the quadriceps femoris muscle causes the
ment). On the afferent side, the stretch recep
lower leg to straighten at the knee, assisting
tor consists of thin muscle fibers (intrafusal
further in the maintenance of an upright
fibers) contained within a spindle-shaped cap
posture.
sule (the muscle spindle). The annulospiral
3. When the hands are outstretched to catch a
(primary) sensory-nerve endings spiral around
falling weight, the weight causes a brief
the thin muscle fibers in the expanded, fluid-
extension of the arm at the elbow and,
filled region at the center of the capsule, and
therefore, a stretch of the biceps muscle in
send impulses to the spinal cord via laxge-
diameter, Group la sensory-nerve fibers, which
Figure 9-2. The muscle stretch reflex, showing the
neuronal connections. A = efferent limb of the re conduct impulses at a high velocity (about 100
flex, B = efferent limb of the reflex, C = gamma m/sec). The intrafusal muscle fibers lie parallel
efferent control, e = excitatory synapse. to the ordinary, extrafusal muscle fibers outside
EXTRAFUSAL
MUSCLE FIBER '______________ J . A L P H A , MOTONEURON
168 Posture and Movement
the spindle that make up the bulk of the muscle the muscle is stretched, the spiral of nerve
and cause it to contract. ending is pulled apart, causing an increased
The group la afferent fibers enter the cord spike frequency as a response (figure 9-3B).
via the dorsal root and pass through the gray The increased spike frequency lasts throughout
matter to the alpha motoneurons in the ventral the stimulus (the stretch receptor is a slowly
horn. There they end in excitatory synapses adapting receptor). The afferent nerve impulses
(labeled e in figure 9 -2 ). Because only a single enter the cord and travel to excitatory synapses
synapse in the CNS is involved, the muscle in the ventral horn. Alpha motoneurons are
stretch reflex is monosynaptic and is the sim then activated and, in turn, activate the
plest reflex in the nervous system. extrafusal muscle fibers, causing the reflex
The efferent limb of the reflex labeled B in response— the muscle contracts and shortens
figure 9 -2 consists of the alpha motoneurons, (figure 9-3C ). If there were no other inputs to
their axons, which exit the cord via the ventral the muscle spindle, this shortening would lead
roots and travel via the peripheral nerve to the to slackness in the intrafusal fiber and an
muscle, and their neuromuscular junctions on absence of response, or silent period, as shown
the extrafusal muscle fibers. The result of nerve in figure 9-3C .
impulses traveling along this pathway is the Note that the reflex response is a contraction
contraction of the muscle. of the same muscle that was stretched. This in
The basic dynamics of the reflex are shown effect counteracts the stretch. At the same
in figure 9 -3 . When the muscle is at its normal time, other muscles acting at the same joint,
resting position, there is typically a fairly steady both synergists (which act in the same way as
resting discharge from the stretch receptor and the stretched muscle) and antagonists (which
its sensory nerve fiber (figure 9 -3 A). When act in the opposite way) are influenced by their
motoneurons in a manner that supplements the
reflex response.
Figure 9-3. Sequence of events m the musde stretch
reflex. (A) Muscle at rest, showing resting discharge We can examine some of these events at a
in Group la afferent fibers from stretch receptors. synaptic level. In brief, the alpha motoneuron
(B) Muscle is stretched (increased in length), show going to the stretched muscle is activated— it
ing increased afferent response. (C) Muscle contracts has an EPSP followed by a spike. At the
(the reflex response), showing the silent period in the neuromuscular junction, release of acetyl
afferent response that would occur without gamma
efferent activity; normally, this silent pause would choline is followed by an end-plate potential,
not occur because the gamma efferents would reduce which leads to a muscle action potential and
the slack in the intrafusal muscle fibers. then muscle contraction. The alpha moto
169 Posture and Movement
neurons going to the synergist muscle are up and down by fibers from the brain that
facilitated— they have subthreshold EPSPs not innervate the gamma motoneurons.
followed by a spike but that make it easier for
other imputs to cause spikes. The alpha
motoneurons to antagonist muscles are in Supraspinal Control
hibited— they have IPSPs, which block spike Both alpha and gamma motoneurons receive
generation and leads to relaxation of antagonist synaptic input from descending tracts originat
muscles. ing in the brain (in supraspinal centers). Major
descending tracts are the corticospinal tract
from the cerebral cortex and the rubrospinal
Gamma Motoneurons
tract, vestibulospinal tract, and reticulospinal
Alongside the cell bodies of alpha motoneurons tract from the brain stem motor areas.
in the cord are smaller gamma motoneurons, In brief, the supraspinal inputs act to:
which send gamma efferent fibers to innervate
the thin, intrafusal muscle fibers within the 1. Control the muscle stretch reflex via a normal
muscle spindle (figure 9 -2 ). The gamma effe balance o f excitatory and inhibitory synaptic inputs
rent fibers are smaller in diameter and lower in onto the motoneurons. Lesions of supraspinal
conduction velocity than the alpha efferent motor structures often may result in an imba
fibers, as shown in table 9 -1 . lance that favors excitation. The muscle
Gamma efferents end in neuromuscular junc stretch reflexes then show a release of function
tions that are located only on the end (polar) and become hyperactive.
regions of the intrafusal fibers. Their activation 2. Initiate voluntary activity. The same spinal
results in contraction of these end regions, motoneurons involved in reflex activity can
resulting in a stretch of the central region with also participate in a voluntary movement when
its associated sensory nerve endings. One effect properly activated from brain centers. Sudden
of this gamma innervation is to take up the activation of either alpha or gamma mo
slack on the intrafusal fibers that would other toneurons may result in muscle contraction. A
wise occur in a contracting muscle, as shown in direct pathway involves activation of alpha
figure 9-3C , and thus prevent the silent period motoneurons, which then activate extrafusal
in firing of the stretch receptor. Gamma activ muscle fibers. A possible indirect pathway in
ity, by filling in the silent period, eliminates volves activation of gamma motoneurons,
this possible gap and helps maintain a continu which then activate the ends of the intrafusal
ous feedback to the CNS of information on muscle fibers, which in turn cause an afferent
muscle length, which is helpful in regulating response in the stretch receptors and their
muscle length. Another effect of gamma activ afferent fibers, which in turn excite the alpha
ity is to make the muscle stretch receptors more motoneurons and lead to a reflex contraction of
sensitive. This sensitivity control can be turned the muscle as a whole. Clearly the indirect
170 Posture and Movement
pathway is more delayed. Actually, recent together in front of the chest and pulled apart;
research shows that some human voluntary this appears to increase motoneuron excitabil
movements involve both direct and indirect ity in a general manner. On the other hand, in
pathways activated at about the same time, a some patients the tendon jerk reflexes may be
phenomenon called coactivation. generally brisk, presumably due to an increased
motoneuron excitability, for example in tense
Clinical Examination o f Stretch Reflexes people.
The tonic stretch reflexes can also be ex
Reflexes are examined by giving the patient a amined clinically. Absent or weak stretch
stimulus (input), and observing the response reflexes can result in reduced muscle tone, or
(output). Different time-courses of stimulus flaccidity. Hyperactive stretch reflexes can re
and response results in either phasic or tonic sult in increased muscle tone.
stretch reflexes. Phasic stretch reflexes result The absence of a muscle stretch reflex may
from a brief stimulus providing a sudden and indicate some lesion of either afferent or effe
transient stretch o f the muscle. Tapping a rent limbs, or the cord at the appropriate
tendon provides such a stimulus. Thus, the segmental level. Such a lesion affecting spinal
tendon jerk or deep tendon reflexes are phasic. motoneurons may be called a lower moto
The brief input results in a reflex contraction neuron lesion. Conversely, hyperactive stretch
that is also brief. Tonic stretch reflexes resultreflexes may suggest a lesion in the descending
from a long-lasting stimulus, such as that fibers from supraspinal motor centers, which
provided by gravity. The long-lasting stimulus may be called an upper motoneuron lesion. For
results in a long-lasting muscle contraction. cerebral lesions such effects tend to be crossed;
During normal posture, this continuous muscle that is, a lesion affecting descending fibers from
contraction is known as muscle tone. Tone is the right hemisphere will lead to hyperactive
also reflected in a normal resistance to passive stretch reflexes on the left side of the body. A
movement: when an examiner tries to dorsiflex hyperactive stretch reflex may give rise to
a foot, there should be some resistance, pro spasticity, a heightened resistance to passive
vided by a tonic stretch reflex in the gastroc movement, and in some cases to clonus, in
nemius muscle. which reflex contraction dies away and returns
Tendon jerk reflexes may be elicited at a rhythmically, leading to a series of muscle
number of sites involving spinal roots at various contractions at a rate of about six per second.
segments of the cord. For example, the biceps
jerk involves spinal segments C5 and C6, the
Inverse Stretch Reflex (Golgi Tendon Organ
triceps jerk involves C6 and C7, the knee jerk
Reflex)
involves L2 to L4, and the ankle jerk involves
S I. The inverse stretch reflex was so named be
It is important to compare the same reflexes cause it causes stretch of a muscle, at least past
on the right and left sides to see if they are a certain point, to result in reflex relaxation of
symmetrical. Tendon jerk reflexes may be small the same muscle, which is opposite to the
or absent throughout the body in some cases, action of the stretch reflex. Clinically, this is
presumably when the excitability of alpha and particularly evident in some spastic limbs: for
gamma motoneurons is low. Reflexes can then example, passive flexion of a joint initially
be reinforced by asking the patient to voluntar leads to great resistance, due to a hyperactive
ily contract some other muscles, as in Jendras- stretch reflex, but after a point, the resistance
sik’s maneuver where the hands are clasped collapses, due to the inverse stretch reflex. In
171 Posture and Movement
such cases, the inverse stretch reflex is thought vents overall change in muscle length. The
to protect the tendon from being tom by GTO impulses travel to the cord along the
excessive tension. In ordinary life, this reflex is large Group lb afferent fibers, which are only
thought to regulate the tension of a muscle, as slightly smaller than the Group la afferents
opposed to the stretch reflex which regulates from the annulospiral stretch receptors.
the length of a muscle. In many activities, such In the cord, the synaptic effect of these
as, paddling a canoe, one might wish to impulses from the GTOs is to exert inhibition
maintain a fairly constant muscle tension even (labeled i in figure 9 - 4 ) on the alpha
while joint angles and muscle lengths are motoneurons leading to the same muscle.
continuously changing, and here the inverse There is probably a short, inhibitory inter
stretch reflex could play a role. neuron in the reflex arc between the afferent
and efferent neurons; thus, the reflex is thought
Neuronal Mechanism. The receptors are the to be disynaptic. The effect of inhibiting the
Golgi tendon organs (GTOs) located in the impulses in the alpha motoneuron is to cause
muscle tendon (figure 9 -4 ), in series with the relaxation of the muscle, which is the reflex
muscle fibers. These receptors respond to in response.
creased tension, whether due to an externally
applied stretch of the muscle (passive stretch),
or to contraction of the muscle itself, particu Flexor Withdrawal Reflex
larly against an external resistance, as in The flexor withdrawal reflex is typically the
isometric contraction when the resistance pre response of an arm or leg to a painful or hot
stimulus, which results in the withdrawal of the
figure 9-4. The inverse stretch reflex; e = excita limb from the stimulus, associated with flexion
tory synapse, i = inhibitory synapse. at several joints in the limb. Common exam-
pies are when you pull your hand away from a also excite a chain of intemeurons in the gray
hot frying pan or lift your foot away from a matter of the cord that eventually excite a
thumbtack. In these examples, the reflex re number of alpha motoneurons in the ventral
sponse generally begins before you are con horn of the cord. The motoneurons are diffuse
sciously aware of the painful stimulus— because in that they may be located in several segments
the spinal reflex arc can function before the of the cord, and their effect is to cause the
afferent impulses have affected the brain. In contraction of flexor muscle fibers in the
the second example, the opposite foot must several joints of the affected limb. The spread
support the body alone; it is aided in this task of activity from a limited input to a widespread
by an associated reflex called crossed extension, output is called divergence.
which activates motoneurons to the extensor The reflex is termed polysynaptic because a
muscles of the opposite leg to withstand the number of CNS synapses are involved. The
pull of gravity. greater latent period, as compared with that of
the muscle stretch reflex, is attributed in part to
Neuronal Mechanism. The receptors are pain the large number of synaptic delays in the reflex
and temperature receptors in the skin, giving arc. The reflex response may continue for
rise to Group III (or A-delta) and Group IV (or several seconds after a strong stimulus is
C) fibers which enter the cord via the dorsal stopped; this afterdischarge may be due to the
roots. In chapter 5, we saw how these fibers intemeuronal circuits, some of which are circu
synapse in the dorsal gray horn of the cord and lar and lead to repeated excitation of the same
relay pain and temperature information to pathway.
neurons whose axons cross over to ascend via
the lateral spinothalamic tract to the brain. In
Other Reflexes and Responses
addition (figure 9 -5 ), these dorsal root fibers
Figure 9-5. The flexor withdrawal reflex. S.G. = In addition to the elementary spinal reflexes, a
substantia gelatmosa, e = excitatory synapse. number of other reflexes and automatic pos-
173 Posture and Movement
tural responses have been described. Several of motor areas in the brain). These centers are of
these, such as the tonic neck reflex, can be major importance physiologically and clinic
seen in infancy, disappearing with the matura ally. Unfortunately for the student, the struc
tion of the supraspinal motor mechanisms. ture and activity of these centers are complex
Only three examples will be listed here. Al and not completely understood. The approach
though largely spinal, they are subject to of this book is to simplify this subject; more
supraspinal influence. advanced textbooks may then be consulted for
further detail.
Tonic Neck Reflexes. When the head is turned The four major centers to be described here
to the right, the right arm and leg are are the motor cortex, the brain stem motor
extended, and the left limbs are flexed; when centers, the basal ganglia, and the cerebellum.
the head is turned to the left, the left limbs These areas interact with each other, and with
are extended, and the right limbs are flexed. spinal reflex mechanisms including afferent
This resembles, and might have led to, the fibers and motoneurons.
pose of the boxer or archer. The sensory
receptors are in the joints of the cervical
vertebrae. Motor Cortex
Positive Supporting Reaction. When the sole of
the foot touches the ground, both the exten Structure. The motor cortex is located in the
sors and flexors of the leg contract, making frontal lobe of the cerebrum; it includes the
the leg a rigid pillar, as in the standing precentral gyrus (just anterior to the central
posture. sulcus), and is also known as Brodmann’s area
Plantar Reflexes. The stimulus is applied to the 4.
outer border of the sole of the foot by The motor cortex is a layered structure
scratching it with a key or other blunt featuring [31] large cells in layer V that because
object. The normal response, after the first of their shape are called giant pyramidal cells and
year of life, is a plantar flexion (downward send axons to the spinal cord; the length of
bending) of the toes, usually combined with some of these axons exceeds one meter. The
upward flexion of the foot at the ankle. axons of these and other pyramidal cells form
During the first year of life, before the the corticospinal tract, so named because it is a
supraspinal motor systems (in particular the direct pathway from the cortex to the spinal
pyramidal or corticospinal tract) have com cord (figure 9 -6 ). In the cerebrum, as this tract
pletely developed, the response is an upward passes between the thalamus and the basal
movement of the big toe. In adults, a similar ganglia, it is part of the internal capsule, a
upgoing toe response can be found during structure of great clinical importance as a
sleep, and also with lesions of the pyramidal frequent site of damage in cerebrovascular
or corticospinal tract. Another term for the accidents (or strokes); in the medulla, it forms
upgoing toe response is Babinski's sign. two prominent ridges along the ventral surface
known as the medullary pyramids. In the
medulla, near its border with the spinal cord,
these fibers cross over to the opposite side and
Supraspinal Motor Control
then descend the cord as the lateral corticos
The spinal reflex mechanisms discussed in the pinal tract. At various segments of the cord,
previous section are normally under the control fibers turn off from this tract to synapse on
of supraspinal motor centers (higher centers, motoneurons in the ventral hom, either di
174 Posture and Movement
Figure 9-6. The lateral corticospinal tract, from axons of these cells control. These movements
neurons m the motor cortex to spinal motoneurons. are mostly on the contralateral (opposite) side
of the body, by virtue of the largely crossed
rectly or indirectly via intemeurons. A smaller nature of the descending tracts. When the
number of fibers (about 10 %) do not cross and induced movements are recorded on a diagram
form the medial corticospinal tract. Other fibers
of the motor cortex, the result is a somatotopic
from motor cortex project to brainstem motor map— a map of the body on the motor cortex. 1
areas, basal ganglia, and cerebellum. On this map the legs are located on the medial
surface of the cerebral hemisphere, and the
Somatotopic Mapping. Artificial stimulation of hands and face are located ventrolaterally,
area 4 with low-voltage electrical pulses can closer to the temporal lobe. The map is
cause movement of various parts of the body. distorted in that a disproportionately large area
The pulses activate pyramidal cells in a particu
lar part of the motor cortex, and the resulting 'Another type of somatotopic map is found on the somatic
movements show which parts of the body the sensory area of the cortex, as described in chapter 5.
175 Posture and Movement
of the cortex is devoted to control of the hands corticospinal fibers. Other changes, such as
and face, areas capable of the complicated and enhanced stretch reflexes and spasticity, are
precise movements. due to cortical lesions that extend beyond
Most neurons that originate in the motor area 4 to area 6 and involve a relay in
cortex and project via the corticospinal tracts brainstem motor areas. Spinal-reflex
affect spinal motoneurons indirectly via one or changes are also on the side opposite to the
more spinal intemeurons. However, some brain lesion.
neurons project directly to spinal motoneurons,
particularly those that govern discrete move Brainstem Motor Areas
ments of finger muscles. Both cases are illus
trated in figure 9 -6 . In addition to the direct pathway from motor
cortex via internal capsule, pyramidal tract,
Function. The motor cortex functions to initi and corticospinal tract to spinal motoneurons,
ate voluntary movement and to control spinal there are important indirect pathways in which
reflexes. Simple voluntary hand movements, the motor cortex relays information via several
such as pressing a button with the thumb, have brainstem motor areas and their descending
been studied in humans and animals [8,33] tracts to spinal motoneurons. These brainstem
enough to present a tentative outline of the areas also receive inputs from noncortical
neural events involved. About 100 msec before sources. The brainstem motor areas and their
the movement, some motor cortex neurons descending tracts contribute to the normal
begin firing; others, which were tonically ac control of posture and movement and also
tive before, now stop firing. After a time account for many signs of motor disorder after
interval necessary for conduction down the brain lesions.
corticospinal tract and synaptic delay, spinal
motoneurons in the cervical cord are activated. Structure. Three brainstem motor areas are de
Both alpha and gamma motoneurons appear to scribed here (figure 9 -7 ).
be activated at about the same time, a process
known as coactivation. The alpha moto 1. The red nucleus (nucleus ruber) is located in
neurons cause muscle contraction directly, the midbrain, the highest division of the
while the gamma motoneurons work indirectly brainstem. It receives inputs from the motor
via the muscle stretch reflex arc. cortex, cerebellum and possibly the basal
Destructive lesions of motor cortex (area 4) ganglia. It gives rise to the rubrospinal tract,
or of its descending projections in the internal which descends in the lateral column of
capsule or other levels of the brain have the spinal white matter, adjacent to the lateral
following effects: corticospinal tract.
2. The reticular formation is a column of
1. Voluntary movements are diminished neurons at the core of the brainstem ventral
(paresis) or lost (paralysis) in the corres to the fourth ventricle. The many cells with
ponding parts of the body, on the side short, branching axons and their numerous
opposite to the lesion. synapses running the length of the brainstem
. 2. Certain spinal reflexes are altered. In par have an appearance that is netlike or re
ticular, a positive Babinski’s sign appears, in ticulated, giving rise to the name. (See more
which stroking of the sole of the foot results detailed texts for discussion of lateral, me
in upward extension of the big toe. After dial, and other subdivisions.) The reticular
infancy this reflex is normally suppressed by formation receives inputs from the motor
Can Brain
Connectivity Improve
with Practice?
Recent research suggests that
corticospinal tract axons might
develop more connectivity (or
connectedness) to spinal
motoneurons with practice. Brain
imaging in volunteers used the
direction of water molecules to
analyze the microstructure of specific
nerve pathways by means of
Diffusion Tensor Imaging (DTI).
References:
2. Cranial nerves
I Smell
n Vision: acuity, fields, fundus
m Pupils, accommodation
in, iv, vi Ocular motility
V Masseter
V Facial touch and pain
VII, IX, X Taste
vn Facial movement
vm Hearing
IX, X Gag
X Palate
XI Stemomastoid and trapezius
XII Tongue
179' Posture and Movement
3. Motor
General
Gait
Heel-toe walking
Tandem gait
4. Reflexes
Deep Tendon Reflexes (DTRs):
Biceps C5,6
Brachioradialis C5,6
Triceps C7
Knee jerks (quads) L3,4
Ankle jerks (Achilles) SI
Plantar reflex: Babinski sign
Superficial reflexes:
Abdominal
Cremasteric
5. Sensory
Cutaneous (small fiber): light touch, pain, temperature
Discriminative (large fiber): vibration, two-point tactile, position sense, others
6. Cerebellar (coordination)
Finger to nose
Heel to knee
Alternating movements
7. Gait
Apraxia, ataxia, spasticity, other
179' Posture and Movement
3. Motor
General
Gait
Heel-toe walking
Tandem gait
4. Reflexes
Deep Tendon Reflexes (DTRs):
Biceps C5,6
Brachioradialis C5,6
Triceps C7
Knee jerks (quads) L3,4
Ankle jerks (Achilles) SI
Plantar reflex: Babinski sign
Superficial reflexes:
Abdominal
Cremasteric
5. Sensory
Cutaneous (small fiber): light touch, pain, temperature
Discriminative (large fiber): vibration, two-point tactile, position sense, others
6. Cerebellar (coordination)
Finger to nose
Heel to knee
Alternating movements
7. Gait
Apraxia, ataxia, spasticity, other
181 Posture and Movement
In addition to posture, the reticulospinal and (together known as the striatum) receive affe
vestibulospinal tracts help control the moto rent input from the association and motor
neurons innervating muscles of the central and cortex, and the third sends efferent output back
proximal parts of the body, including the neck, up to the motor cortex (via a relay in the
back, shoulder, and hip joints, which are thalamus) and down to the brainstem motor
important in large, integrated movements of areas. These nuclei connect to each other and
the body and limbs, such as turning, bending, also to other nearby nuclei that are often
or walking. The rubrospinal tract, in contrast, included in the basal ganglia: the substantia
may assist the adjacent lateral corticospinal nigra and the subthalamic nucleus.
tract in controlling independent movements of Compared with other brain regions, the
the more distal muscles, such as those of the basal ganglia have extremely high concentra
fingers [28]. tions of two neurotransmitters, acetylcholine and
dopamine. Neurons originating in the substantia
Basal Ga.ngfia nigra secrete dopamine at their synaptic ter
minals in the striatum, and acetylcholine is
Structure. The basal ganglia are a group of secreted at synapses between striatal neurons,
nuclei located well beneath the surface of the according to present evidence [31]. These
cerebral hemispheres, separated from the transmitters are of practical importance in
thalamus by the fibers of the internal capsule. diseases affecting the basal ganglia.
The major nuclei are the caudate, putamen, and
globus pallidus (figure 9 -8 ). The first two Normal and Disordered Function. Most of the
interest in basal ganglia function derives from
Figure 9-8. Basal ganglia, indicating major neuronal the movement disorders associated with lesions
connections between nuclei in the basal ganglia and and dysfunction in the human basal ganglia.
relationships with other structures. C = caudate nu Parkinsonism, the most common movement
cleus, P = putamen, Sm = striatum, GP = disorder, has both negative and positive symp
globus pallidus, SN = substantia nigra, STN =
subthalamic nucleus. Thalamic relay nucleus between toms. The negative symptom (reflecting insuf
basal ganglia and motor cortex is not shown. ficient activity) is both a difficulty in starting
and a decrease in voluntary movements (hy-
pokinesis). The positive symptom (reflecting
M O TOR C O R T E X
excessive activity) is an increased tone of both
extensor and flexor muscles, leading to rigidity;
the rigidity switches on and off at 5 to 10 c/s
during passive movement of a limb (cogwheel
rigidity) or at rest (resting tremor).
In most cases, examination of the brain by
B R A IN ST E M B A S A L GANGLIA
to O T O R A R E A S pathologists shows degeneration in the substan
C *P
GP
Sm . tia nigra, associated with a reduction in the
amount of dopamine in the substantia nigra
s and striatum. Normally, a group of nerve cells
in the substantia nigra sends axons to the
RF striatum, where they secrete dopamine; the
axons are called nigrostriate dopaminergic fi
bers. A loss of these neurons is thought to be
n the underlying lesion in parkinsonism (figure
9 -9 A and 9-9B ).
181 Posture and Movement
In addition to posture, the reticulospinal and (together known as the striatum) receive affe
vestibulospinal tracts help control the moto rent input from the association and motor
neurons innervating muscles of the central and cortex, and the third sends efferent output back
proximal parts of the body, including the neck, up to the motor cortex (via a relay in the
back, shoulder, and hip joints, which are thalamus) and down to the brainstem motor
important in large, integrated movements of areas. These nuclei connect to each other and
the body and limbs, such as turning, bending, also to other nearby nuclei that are often
or walking. The rubrospinal tract, in contrast, included in the basal ganglia: the substantia
may assist the adjacent lateral corticospinal nigra and the subthalamic nucleus.
tract in controlling independent movements of Compared with other brain regions, the
the more distal muscles, such as those of the basal ganglia have extremely high concentra
fingers [28]. tions of two neurotransmitters, acetylcholine and
dopamine. Neurons originating in the substantia
Basal Ga.ngfia nigra secrete dopamine at their synaptic ter
minals in the striatum, and acetylcholine is
Structure. The basal ganglia are a group of secreted at synapses between striatal neurons,
nuclei located well beneath the surface of the according to present evidence [31]. These
cerebral hemispheres, separated from the transmitters are of practical importance in
thalamus by the fibers of the internal capsule. diseases affecting the basal ganglia.
The major nuclei are the caudate, putamen, and
globus pallidus (figure 9 -8 ). The first two Normal and Disordered Function. Most of the
interest in basal ganglia function derives from
Figure 9-8. Basal ganglia, indicating major neuronal the movement disorders associated with lesions
connections between nuclei in the basal ganglia and and dysfunction in the human basal ganglia.
relationships with other structures. C = caudate nu Parkinsonism, the most common movement
cleus, P = putamen, Sm = striatum, GP = disorder, has both negative and positive symp
globus pallidus, SN = substantia nigra, STN =
subthalamic nucleus. Thalamic relay nucleus between toms. The negative symptom (reflecting insuf
basal ganglia and motor cortex is not shown. ficient activity) is both a difficulty in starting
and a decrease in voluntary movements (hy-
pokinesis). The positive symptom (reflecting
M O TOR C O R T E X
excessive activity) is an increased tone of both
extensor and flexor muscles, leading to rigidity;
the rigidity switches on and off at 5 to 10 c/s
during passive movement of a limb (cogwheel
rigidity) or at rest (resting tremor).
In most cases, examination of the brain by
B R A IN ST E M B A S A L GANGLIA
to O T O R A R E A S pathologists shows degeneration in the substan
C *P
GP
Sm . tia nigra, associated with a reduction in the
amount of dopamine in the substantia nigra
s and striatum. Normally, a group of nerve cells
in the substantia nigra sends axons to the
RF striatum, where they secrete dopamine; the
axons are called nigrostriate dopaminergic fi
bers. A loss of these neurons is thought to be
n the underlying lesion in parkinsonism (figure
9 -9 A and 9-9B ).
182 Posture and Movement
A B C
Normal Parkinsonism Phenothiazines
Striatum:
CK - 0 * 0 -O Q .
S.N.:
(J!> -■ o -
Figure 9 -9 . Proposed neuronal mechanism in parkin als at synapses in the striatum (figure 9-9B ), or
sonism. S.N. = substantia nigra, DA = (2 ) blockade of postsynaptic dopamine receptors
dopamine. Dashed lines show sites of degeneration or at the same synapses (figure 9-9C ). (The
chemical blockade. (A) Normal mechanism. (B) In
parkinsonism, cells in substantia nigra and their favorable effect of phenothiazines on psychiat
dopamine-secreting terminals degenerate. (C) ric symptoms may be due to their action at
Phenothiazine drugs can block dopamine receptors, other synapses.)
leading to side effects that resemble parkinsonism. In addition to L-dopa, anticholinergic drugs
such as atropine may also be given to treat
The presumed dopamine deficiency cannot parkinsonism. Their effect seems to be on
be treated by administering dopamine itself, cholinergic synapses in the striatum, whose
because it cannot cross the barrier from the action opposes that of the dopaminergic sy
circulation into the brain (the blood-brain napses. Drugs that mimic or increase the effects
barrier). The deficiency can be relieved by of acetylcholine tend to worsen parkinsonian
administering L-dopa (L-dihydroxyphenyla- symptoms.
lanine), which can cross the barrier and then O th er movement disorders— chorea,
be converted into dopamine. L-dopa provides athetosis, and ballismus— have mainly positive
symptomatic relief in many cases and represents symptoms: involuntary movements, that is,
a major advance in neurological treatment, in movements over which the patient seems to
that it represents a partial replacement of a have no control. In chorea, the movements are
transmitter known from other evidence to be jerky or birdlike. In athetosis, the movements
deficient. are writhing or snakelike. Both chorea and
A syndrome resembling parkinsonism can athetosis are associated with lesions in the
also be induced as a side effect of the striatum. In ballismus, the movements resemble
phenothiazine drugs, such as chlorpromazine those involved in flinging a ball. Ballismus is
(Thorazine) and trifluoperazine (Stelazine), associated with lesions in the subthalamic
that are commonly prescribed in psychiatric nucleus, which is closely connected with the
practice. A possible basis for this side effect is globus pallidus.
that the phenothiazines block dopamine recep A possible interpretation for these move
tor sites in the postsynaptic membrane of ment disorders is that a controlling influence,
neurons (figure 9-9C ). Thus, it seems that possibly inhibitory, normally exerted by basal
similar symptoms can result from ( 1) degenera ganglia neurons on the motor cortex, is dis
tion of dopamine-secreting presynaptic termin turbed by such lesions, leading to excessive
183 Posture and Movement
activity in the motor cortex. This interpreta cerebrum, it has a cellular cortex surrounding a
tion is supported by some animal experiments white region of nerve fibers that contains
on basal ganglia function. When electrical several subcortical nuclei. All cerebellar output
stimulation of the motor cortex was used to signals are carried by axons of the large Purkinje
initiate a movement, the movement could cells, the principal cells of the cerebellar cortex.
often be stopped by stimulation of the striatum. These cells have many-branched dendritic
In other experiments, spike activity recorded trees, containing thousands of synapses from
from motor cortex neurons was reduced in input fibers and the numerous intemeurons
frequency by electrical stimulation of the (basket cells, granule cells, and stellate cells)
striatum. Both effects could result from striatal that modify the activity of the Purkinje cell.
inhibition of motor-cortex neurons. A corol The three divisions of the cerebellum are
lary to this interpretation is that lesions in the shown in figure 9 -1 0 . The archicerebellum
substantia nigra increase the inhibitory activity (Arch, in the figure), the most primitive part,
of the same striatal neurons, leading to the has input and output connections with the
negative symptoms of parkinsonism (reduced vestibular nuclei, one of the brainstem motor
and slowed movement). areas described earlier in the chapter. The
Other experiments showed that striatal paleocerebellum (Pal.) receives afferent input
neurons, recorded with microelectrodes, in from muscle spindles, Golgi tendon organs,
creased their firing rate at the beginning and joint receptors, and other peripheral receptors,
end of voluntary movements, particularly when via the rapidly conducting spinocerebellar
the movements were slow. This has led to tracts (and others), and sends output to the
speculation that the basal ganglia are involved brainstem motor areas. The neocerebellum
in the programming (initiation, guidance) of (Neo.), the major part in humans, receives
slow voluntary movements. input from several areas of cerebral cortex
The two different effects that striatal neurons including the motor cortex, relayed through
may exert on motor cortex— inhibition and the pons; it sends output fibers back to the
initiation of activity— are not necessarily in motor cortex, relayed through the thalamus.
conflict. Initiating a new movement often
requires an inhibition of the preexisting move Function. The archicerebellum developed in
ment or posture (e.g., sitting down requires a connection with the vestibular nuclei and
suppression of the muscle contractions in functions with it in maintaining balance. Le
volved in the standing posture). Both effects sions of this area lead to nystagmus, head
could therefore be part of the same overall rotation, and/or difficulty in maintaining
basal ganglia-motor cortex system. balance while walking.
The paleocerebellum and neocerebellum
Cerebellum function in the control of posture and move
ment. One effect of cerebellar lesions, related
Like the basal ganglia, the cerebellum forms a to posture, is muscular hypotonus due to under
closed loop of neural connections with the active muscle stretch reflexes. Other effects,
cerebral cortex and modifies cerebral output to related to voluntary movement, include the
spinal motoneurons. following:
M O TO R C O R T E X
CEREBELLU M B R A IN ST E M
M O TO R A REA S
N eo.
Pal.
V E S T IB U L A R A rch.
ORGAN
E 2
RF
JO IN T
M U SC L E
Figure 9-10. Cerebellum, indicating major divisions closed and arms stretched out to both sides,
and relationships with other structures. Arch. = who then attempts to bring together the middle
archicerebeUum, Pal. = paleocerebellum, Neo. =
fingers of both hands. Without visual informa
neocerebellum. Afferent information from joint and
muscle receptors ascends in the spinocerebellar and tion this action is difficult but still possible.
other tracts. Observe how errors (deviations from the target)
are corrected, and how the movement slows
over a glass when reaching for it; failure to down when the target is nearly reached. In
terminate a motion properly. everyday life, similar corrections in skilled
2. A diadochokinesis— difficulty in making movements, such as reaching for and grasping a
rapidly alternating movements, for example, cup, are done automatically and unconsciously,
inability to rotate the palm up and down in large part by the cerebellum and its interac
quickly. tions with other structures. According to one
3. Scanning speech— difficulty in making the model, even before the motor cortex initiates
rapid movements of the vocal apparatus the movement, the cerebellum may be in
necessary for smooth speech. volved in planning (programming, sequencing)
4. Intention tremor— a tremor that appears dur the movement. For this stage, the cerebellum
ing a voluntary movement rather than dur receives input information from association
ing rest. cortex, processes it, and sends output informa
tion to motor cortex. At the initiation of the
These effects on movement have been related movement, the motor cortex sends signals not
to the role of the cerebellum as a system that only to the spinal motoneurons but also to the
uses feedback signals to correct errors. cerebellum, informing it of the intended move
To understand the concept of error correc ment (“reach for the cup”). The signals sent to
tion, you might watch a subject, with eyes the cerebellum are processed within its com
185 Posture and Movement
plex neuronal circuitry and combined with The tone of the antigravity muscles during
feedback signals from muscle, joints, and other the upright posture is maintained by the stretch
peripheral receptors, which provide informa reflex, utilizing afferent path 1 as shown in the
tion on the actual moment-to-moment position diagram, with the strength of that reflex deter
and speed of the hand as it approaches the cup. mined largely by tract 3. Voluntary, purposive
Output signals then are sent from the cerebel movement is associated with control signals in
lum back to the motor cortex and lead to descending tracts 2 and 3; the precise move
correction of errors (for example, “more to the ments of the hands and fingers are particularly
right,” “slow down,” “stop”) during the dependent on tract 2 .
movement. Information descending through tracts 2 and
On the basis of this model, cerebellar lesions 3 is produced by neural activity in four inter
could lead to deficiencies in planning voluntary related supraspinal motor systems: motor cor
movements, in correcting errors during the tex, brainstem motor areas, basal ganglia, and
movements, or both. If, for example, the hand cerebellum. The loop from cortex to basal
strays to one side of the target and the error is ganglia and back suggests that motor cortex
overcompensated by moving it too far to the activity may be slowed down, inhibited, or
other side, a back-and-forth movement (inten otherwise modified by the basal ganglia. The
tion tremor) can result. If the hand moves too loop from cortex to cerebellum and back
rapidly toward the target, and this error is not suggests that the cerebellum could play a
corrected in time, the hand will overreach its similar role. In addition, afferent input to the
target (dysmetria). cerebellum from muscle, joint, and other sen
sory receptors, and the outcome of cerebellar
lesions, suggests that the cerebellum could
function as the central component of a ser
An Overview of Motor Mechanisms
vomechanism. That is, it could combine infor
A schematic diagram of the various motor mation from the motor cortex on the intended
systems combined is shown in figure 9-11. action (e.g., “reach for the cup”) with informa
Note the central position of the alpha tion from sensory receptors on the current
motoneuron, receiving and integrating infor situation (“the hand is moving too quickly”) to
mation from the following sources: produce an error-correction signal (“slow
down”). The error-correction signal could go
1 muscle stretch receptors (and other afferent back up to the motor cortex and/or be transmit
input not shown). ted downward via the brainstem motor areas.
2 the motor cortex, via corticospinal (py What precedes activity of the motor cortex
ramidal) tract fibers. in the execution of a voluntary movement?
3 the brainstem motor areas, via the descend Neural activity in the following areas has been
ing rubrospinal, vestibulospinal, and re suggested: ( 1 ) other areas of cerebral cortex,
ticulospinal tracts (extrapyramidal tracts). especially the premotor cortex, Brodmann’s
area 6 , and the prefrontal lobes, anterior to
The result of all this information is muscle area 6 ; (2) basal ganglia; and (3) cerebellum.
contraction determined by the frequency of These possibilities are under investigation. For
action potentials coming from the alpha one important type of voluntary activity,
motoneuron. The motoneuron has therefore speech, the influence of a frontal lobe region
been called the final common path of the motor (known as Broca’s area) anterior to the part of
systems. the motor cortex that controls the muscles of
186 Posture and Movement
M O TO R C O R T E X
CEREBELLUM B R A IN ST E M B A SA L GANGLIA
M O TO R A REA S
C *P
N eo.
EJpx^Hl GP
Sm .
Pal. —
1s t n | I S N |
V E S T IB U L A R
Arch. N
ORG AN
RF
r r
JO IN T
M U SC L E
(D
—V--
PN S CNS
Review Questions
1. Diagram and describe the muscle stretch reflex, including die role of muscle spin
dle receptors, alpha motoneurons, and gamma motoneurons.
2. Give two examples of how the muscle stretch reflex helps to maintain an upright
posture.
190 Posture and Movement
5. Compare the activation of alpha motoneurons either directly or indirectly via the
gamma motoneurons. Define coactivation.
6. Define phasic and tonic stretch reflexes. Describe how each can be tested clinical
ly, giving examples.
191 Posture and Movement
7. Define muscle tone, flacddity, spasticity, and clonus. Relate each term to spinal
reflexes.
10. For the three major spinal reflexes described (muscle stretch, inverse stretch, and
flexor withdrawal), list the following information in a table: effective stimulus,
receptor, afferent fiber type (I, n, etc), number of synapses in the CNS, and effect
on motoneurons (showing which are excited and which are inhibited).
193 Posture and Movement
12. Describe the tonic neck reflexes, positive supporting reaction, and plantar reflex.
Define Babinski's sign and describe its significance.
13. List in outline form the major supraspinal motor-control centers and their com
ponents and output tracts.
14. Describe tire somatotopic map on the motor cortex and the effects of motor-cor-
tex stimulation.
194 Posture and Movement
16. Describe the functions of the brainstem motor areas, according to current infor
mation.
17. Describe the symptoms of parkinsonism and their relationship to dopamine defi
ciency, L-dopa medication, and undesirable side effects of die phenothiazine
drugs.
19. Diagram and describe the function of the cerebellum as a servomechanism utiliz
ing feedback to control voluntary movement
20. List the possible manifestations of lesions in the following areas: motor cortex,
internal capsule, striatum, substantia nigra, archiceiebellum, paleocerebellum,
and neocerebellum.
21. Describe how alpha motoneurons integrate information from several sources.
10. Autonomic Function
A novice skier about to start down a mountain While the posture and movement of the
slope is excited and a little anxious. His heart body in the outside environment are controlled
beats rapidly and with more force than usual, by the somatic motor nerve fibers, the function
and his digestive processes seem to be shut of the heart, blood pressure and flow, body
down. The circulation to his skin is reduced, temperature, and other variables of the internal
making it pale and cool. Sugar is released into environment are controlled in large part by the
his blood from storage sites within liver and visceral motor nerve fibers of the autonomic
muscle. His pupils are dilated. nervous system. While the somatic motor
All these various responses are due to activa nerve fibers control the skeletal muscles, the
tion of the sympathetic nervous system, one of visceral motor nerve fibers control the smooth
the two divisions of the autonomic nervous muscles, cardiac muscle, and exocrine glands.
system. The responses are also related in that In both cases, motor output is influenced both
they all can contribute to physical performance by sensory input and by control centers in the
under challenging conditions, especially in the brain.
presence of danger or cold. The increased
cardiac output, measured as liters of blood
pumped per minute, is largely available to
Functional Anatomy
supply the skeletal muscles, since the require
ments of the digestive organs and the skin are
Sympathetic Nervous System
reduced. The muscles, about to produce extra
work, will be nourished by the additional sugar The sympathetic efferent nerves originate in a
and oxygen supplied by the blood. The dilated group of nerve cells running along the lateral
pupils allow obstacles to be seen even in dim horn of the spinal gray matter in the thoracic
light. and upper-lumbar segments (from T1 to L3) of
In contrast, hours later, the same person is the spinal cord (figure 10-1). These cells send
relaxed and drowsy after a good dinner and a axons through the ventral roots to the chain of
glass of beer. His heart rate is slow; digestion is sympathetic ganglia alongside the cord or to
active; skin is flushed and warm, supplied by an more peripheral ganglia and are known as the
increased blood flow; and pupils are con preganglionic sympathetic neurons. Within the
stricted. These responses are all due to activity ganglia, preganglionic nerve fibers synapse
in the parasympathetic nervous system, the onto postganglionic sympathetic neurons. Some
other division of the autonomic nervous sys preganglionic fibers travel several segments up
tem. In many but not all instances, parasym or down the chain of ganglia before synapsing,
pathetic effects are associated with relaxation and some synapse only on more peripheral
and rebuilding of stores of metabolic energy. ganglia. The postganglionic sympathetic axons
197
198 Autonomic Function
submandibular
and sublingual
glands
parotid gland
bronchial tree
stomach
adrenal
medulla
large
intestine
urinary
D
bladder
Figure 10-1. The sympathetic nervous system, show are slowly conducting, unmyelinated C fibers.
ing efferent fibers; chains of sympathetic ganglia on They travel outward via the peripheral nerves
both sides of the spinal cord; peripheral ganglia
(e.g. = celiac ganglion, s.m.g. = superior mesen to various effector organs, where they supply
teric ganglion, and i.m.g. = inferior mesenteric smooth muscle, cardiac muscle, and exocrine
ganglion); and effector organs. Source: Ban, M .L glands.
The Human Nervous System: An Anatomic The secretory cells of the adrenal medulla
Viewpoint (3rd edL). Hagerstown, MdL: Harper & have an embryonic origin similar to that of the
Row Publishers, Inc., 1979. Reprinted with
permission. postganglionic sympathetic neurons. Like those
199 Autonomic Function
Figure 10-2. The parasympathetic nervous system, neurons, they are innervated by preganglionic
showing efferent fibers in cranial nerves 111, VII, sympathetic fibers. The secretions of the
IX, and X, and in pelvic nerves; ganglia (e.g. = medullary cells, however, are released into the
ciliary ganglion, s.g. = submandibular ganglion, and
o.g. = otic ganglion); and effector organs. Source: general circulation.
Barr, M .L The Human Nervous System: An
Anatomic Viewpoint (3rd ed.). Hagerstown, M i:
Harper & Row Publishers, Inc., 1979. Reprinted Parasympathetic Nervous System
with permission. Like the sympathetic system, the parasym
200 Autonomic Function
pathetic nervous system has preganglionic and Supraspinal Control o f the Autonomic
postganglionic neurons (figure 10-2). The pre Nervous System
ganglionic parasympathetic neurons originate in
the brainstem, sending axons outward via the Like the muscle stretch reflex and other spinal
oculomotor, vagus, and other cranial nerves, reflexes that control posture and movement,
and in sacral segments of the spinal cord, the autonomic reflexes of the spinal cord also
sending axons outward via the pelvic nerve are normally under the control of the supra
(also called nervi erigentes). The preganglionic spinal centers. Thus, in the adult, emptying of
nerve fibers are long, and synapse in parasym the bladder and bowel are under voluntary
pathetic ganglia, onto postganglionic parasym control, involving neurons from the brain that
pathetic neurons. Since the parasympathetic descend to the sacral spinal cord. Temperature
ganglia are located near or within their effector regulation is coordinated by neural centers in
organs, the postganglionic axons are shorter the hypothalamus. In patients suffering spinal-
than those of the sympathetic system. cord injury, however, this supraspinal control
can be lost, and reflex responses may be
released from higher control.
Autonomic Reflexes
Similarly, the baroreceptor reflex mecha
The activity of autonomic neurons is controlled nisms within the brainstem are normally subor
in part by reflex activity at a brainstem or spinal dinated to activity within higher centers,
level. Some examples of autonomic reflexes are including areas of the hypothalamus and cere
given below. bral cortex. Thus, for example, emotional
states, such as anger, can cause a rise of blood
1. Within the carotid and other large arteries, pressure that is not corrected by the barorecep
stretch receptors called baroreceptors are ac tor reflex.
tivated by increased arterial blood pressure,
and transmit information to centers in the
brainstem. Reflex output to the heart and
the blood vessels generally causes the blood
Chemical Transmitters
pressure to return toward a normal level. As discussed in chapter 4 (see figure 4-17),
This is known as the baroreceptor reflex. synaptic transmission within the autonomic
2. When stretch receptors within the walls of nervous system involves two types of chemical
the bladder are activated by a full bladder, transmitters: ( 1 ) acetylcholine, secreted by
they transmit information to the sacral cholinergic fibers, and ( 2 ) norepinephrine, also
spinal cord. Within the cord, signals are known as noradrenaline, secreted by adrenergic
passed via intemeurons, leading to reflex fibers. In addition, epinephrine, or adrenaline,
output to the smooth muscles of the bladder. is secreted by the adrenal medulla.
The bladder contracts and empties. A simi
lar spinal reflex leads to emptying of the
Chemical Transmission in the Sympathetic
bowel.
Nervous System
3. Warming the skin can lead to increased
blood flow and reddening of the affected The sympathetic preganglionic fibers secrete
area, while cooling the skin can have an acetylcholine at their synapses with postgan
opposite effect. These effects involve a glionic neurons. The postganglionic fibers in
spinal reflex originating in cutaneous ther general secrete norepinephrine at their syn
moreceptors, with reflex output to cutane apses with effectors and are known as sym
ous blood vessels. pathetic adrenergic fibers (the sympathetic
201 Autonomic Function
Chapter 10
Effectors
Schematic diagram of autonomic nervous system and its neurotransmitters. T = thoracic, L = lumbar, S = sacral segments of spinal
cord. ACh = acetylcholine, E = epinephrine, NE = norepinephrine, ad. med. = adrenal medulla. Dotted line surrounds sympathetic
branch.
203 Autonomic Function
cholinergic fibers that innervate sweat glands Norepinephrine and epinephrine also act
are among the few exceptions to this rule). The upon two different types of receptor sites, both
sympathetic preganglionic fibers to the adrenal within effector organs activated by sympathetic
medulla also secrete acetylcholine at their postganglionic nerve fibers. Alpha receptors
synapses with secretory cells in the adrenal cause constriction of blood vessels, relaxation
medulla. These cells then secrete a mixture of of the intestinal wall, and other effects when
mostly epinephrine with some norepinephrine activated. They can be stimulated by certain
into the bloodstream, which carries it to effec drugs, such as phenylephrine. Beta receptors
tor organs throughout the body. cause dilation of blood vessels, constriction of
the intestinal wall, and other effects when
activated. They can be stimulated by certain
Chemical Transmission in the Parasympathetic drugs, such as isoproterenol.
Nervous System
As in the sympathetic nervous system, the
parasympathetic preganglionic fibers secrete
Responses of Effectors
acetylcholine. Unlike the sympathetic system,
the parasympathetic postganglionic fibers also The autonomic nervous system affects many
secrete acetylcholine at their synapses with physiological functions. A few important ex
effectors, and are known as parasympathetic amples are discussed below.
cholinergic fibers.
Cardiovascular System
Receptor Sites
The heart is constantly affected by both sym
Chemical transmitters react with receptor sites pathetic and parasympathetic neural activity.
in cell membranes of neurons and of effectors Sympathetic stimulation causes an increased
(smooth muscle, cardiac muscle, and exocrine heart rate and an increased force of contrac
glands). A change in membrane potential and tion, which together lead to an increase in the
in neural or effector activity follows. The amount of blood pumped by the heart per
receptor sites are of great practical importance minute or an increased cardiac output.
in that numerous drugs and poisons act upon Parasympathetic stimulation, on the other
them, and such pharmacological substances are hand, causes a decreased heart rate and (in
often classified in terms of the appropriate part) a decreased contractile force, leading to a
receptor site. decreased cardiac output.
Acetylcholine acts upon two different types Sympathetic stimulation causes constriction
of receptor sites. Nicotinic receptors are found of the blood vessels that supply the abdominal
in the postganglionic cells within all autonomic viscera and the skin. Thus, during the excite
ganglia. They are also the receptor type in ment before an athletic event, activation of the
skeletal muscle membrane. The term nicotinic is sympathetic nervous system can cause not only
used because nicotine activates all of these an increased cardiac output but also can cause
receptors. Muscarinic receptors are found in an increased fraction of that output to be
the effector cells activated by the parasym diverted away from the gut and the skin, to be
pathetic nervous system and also by the few available to supply the muscles as they prepare
sympathetic cholinergic fibers. The term mus for extreme exertion. A dilation of blood
carinic is used because muscarine activates all of vessels that supply skeletal muscle (via stimula
these receptors. tion of adrenergic beta receptors), preparatory
204 Autonomic Function
Other Functions
Numerous other functions, described in human
and mammalian physiology, are affected by
autonomic activity. For example, sexual func
tion is influenced by a combination of parasym
pathetic and sympathetic activity. Sympathetic
stimulation can dilate the bronchi in the lungs,
and drugs with a similar effect are often given
to relieve asthma attacks. Sympathetic stimula
tion increases the release of glucose from the
liver and the breakdown of muscle glycogen to
form glucose and thus tends to increase blood
glucose concentration; it also can promote the
breakdown of stored fat, releasing free fatty
acids into the blood.
205 Autonomic Function
Review Questions
2. Give two examples of autonomic reflexes. Describe the influence of sensory input
and of supraspinal control.
205 Autonomic Function
Review Questions
2. Give two examples of autonomic reflexes. Describe the influence of sensory input
and of supraspinal control.
206 Autonomic Function
In the cockpit o f a commercial airliner leaving wakefulness and sleep. Abnormal EEG patterns
Baltimore for London, a pilot communicates in occur in epilepsy, some other brain disorders,
a brisk dialogue with the control tower to gain and some types of metabolic dysfunction.
permission to take off. He must be prepared for
up to 10 hours of alert wakefulness while paying
Method
attention to the information gained through
the cockpit window, from the bank of over 100 An EEG recording electrode is usually a gold or
instruments facing him, and through radio silver disk or cup about one centimeter in
communications. Crucial decisions based on diameter with a wire attached. One electrode,
this information, such as whether to take off the active electrode, is pasted onto the scalp at
once maximum ground speed is attained, must some predetermined site over a particular area
often be made within milliseconds. In order to of cerebral cortex, and another, the reference
decide properly, important information must be electrode, is placed on a relatively inactive site,
remembered from periods of seconds to hours such as an earlobe. The voltage difference
and evaluated on the basis of criteria learned between the two electrodes is them amplified
over thousands of hours of flying time. and displayed on a chart recorder as a graph
Wakefulness and alertness, language, and against time (figure 11-1). The resulting fluc
memory are some of the complex physiological tuations range from 10 to 100 microvolts
and psychological processes that govern how a (millionths of a volt, or 10 “ 6 volt) before
pilot interprets sensory input and produces the amplification, and from 0.5 to 50 Hz in
motor output that guides an airplane. frequency. They change with electrode site,
These higher processes are associated par age, sensory input and levels of sleep and
ticularly with cerebral activity. In some cases, wakefulness.
such cerebral activity is reflected in electrical In neurological diagnosis, electrodes are usu
changes, such as the electroencephalogram; in ally placed on a number of standardized record
others, it is inferred from the deficits following ing sites. In the widely used 10-20 system of
lesions and functional disorders. electrode placement, O, P, T, and F stand for
occipital, parietal, temporal, and frontal lobe
sites, respectively; C stands for the central
sulcus region; odd number subscripts stand for
The Electroencephalogram in Wakefulness, left; even for right; and z for midsaggital
Sleep, and Epilepsy locations (figure 11-2). Thus, O j is a left
The electroencephalogram, or EEG, is a re occipital site and O 2 is the corresponding right
cording of the electrical activity of the cerebral occipital site. P3 and P4 are corresponding sites
cortex, normally associated with levels of over left and right parietal lobes, and so on.
207
208 Cerebral Physiology and Higher Functions
Figure 11-1. Recording the EEG on a single chan electrode is believed to originate in a large
nel AMP = amplifier, O = occipital, P = number (estimates go up to one million) of
parietal, T = temporal, and F = frontal. neurons in the cerebral cortex underlying the
The voltage difference between each electrode electrode site. The small amplitude of the scalp
and the reference electrode is displayed on one EEG compared with potential fluctuations near
of a number of parallel graphs. Alternatively, the neuron is largely due to the distance and
the voltage difference between two active barriers (meninges, blood vessels, bone, skin)
electrodes, such as Oi and P3, can be displayed. between the cortex and the electrode.
Major sources of EEG waves are the postsy
Origin naptic potentials (EPSPs and IPSPs) of cortical
neurons, particularly from the apical dendrites
The EEG obtained from a particular active of cortical pyramidal cells (figure 11-3). When
these occur simultaneously in a number of
Figure 11-2. The International 10-20 electrode sys neurons, they summate at the recording site
tem. Source: Jasper, H.H. The ten-twenty electrode
system of the International Federation. EEG Clin. and produce large, regular waves in the EEG,
Neurophysiol. 10:371-375, 1958. Reprinted with called synchronized EEG waves. Otherwise,
permission. the EEG tends to be small in amplitude and
209 Cerebral Physiology and Higher Functions
A | B C
I
X “
A / V _ ---- F F G
------ EPSPs
!■{ 4 4! 58^ -
Figure 11-3. Schematic diagram of relationship of waves are high-amplitude and low-fre
EEG with postsynaptic potentials of cortical cells. quency, including frequency ranges called
(A) EEG electrode on scalp can record activity theta (4 to 7 Hz) and delta (0.5 to 4). Bursts
originating in underlying brain area, shown by of waves at about 14 Hz (sleep spindles) may
dashed lines, containing cortical neurons of which a,
be interspersed during lighter sleep, while
b, and c, are examples. (B) Synchronized EEG.
(C) Desynchronized EEG. slow delta waves predominate during deeper
sleep. Because these stages lack rapid eye
irregular, or desynchronized. The former can be movements or REMs (see below), they are
compared to the fluctuations of a sound meter called non-REM or NREM sleep.
in an auditorium when people are clapping in 4. Another stage of sleep is characterized by
unison; the latter to the smaller fluctuations rapid eye movements (the eyes typically
when people clap randomly. move left and right, back and forth) and is
called REM sleep. Compared with NREM
Normal EEG Patterns
Figure 11-4. Normal EEG waveforms. (1) Low-am
EEG patterns typically change with levels of plitude beta waves during alertness with eyes open.
wakefulness and sleep in the following manner (2) Moderate-amplitude alpha waves, here at a fre
(figure 11-4): quency of 10 Hz, during relaxed wakefulness with
eyes closed. (3) High-amplitude delta waves during
deep, NREM sleep.
1. During alertness, with eyes open, waves are
low-amplitude, high-frequency, and irregu
lar; they are desynchronized. The frequency 1
range (above 13 Hz) is called beta.
2. During relaxation, with eyes closed, waves
are of moderately high amplitude, moderate
frequency, and regular; they are syn 50 /«-V
chronized. The frequency range (8 to 13 Hz)
is called alplia. Such waves are not always
found in all normal subjects and are most
prominent in occipital sites. Opening the 3 ]
eyes makes the waves disappear and shift to
the beta pattern, a phenomenon called alpha I------------- 1
blocking. 1 sec
3. During most of the night’s sleep, the EEG
210 Cerebral Physiology and Higher Functions
sleep, the EEG is lower in amplitude and A proposed model for the origin of the alpha
higher in frequency (including frequencies rhythm [1] may be summarized as follows: The
in the alpha and beta range). REM sleep has alpha rhythm at about 10 Hz is triggered by a
excited a great deal of interest because vivid similar rhythm of action potentials in non
dreams are often reported by people who are specific thalamic neurons. These neurons can
awakened from REM sleep. Several physio be called principal cells since they send axons
logical changes, such as loss of postural out of the thalamus to the cerebral cortex [31].
muscle tone, increased heart and respiratory They fire at intervals of about 100 msec (0.1
rate, and activity of the sex organs also occur sec) separated by 100-msec-long IPSPs
during this period. In adults, each REM generated by inhibitory intemeurons con
period lasts about 20 minutes and alternates nected to the principal neurons by a feedback
with longer periods of NREM sleep in cycles circuit within the thalamus (figure 11-5B).
of about 90 minutes, with three to five of Since the interneurons have numerous
these cycles occurring during the night. branches, they can synchronize the activity of
Infants spend a larger fraction of their sleep numerous principal cells in the thalamus,
time in the REM stage. which in turn can synchronize the activity of
numerous cortical cells. The nerve impulses
conducted along principal cell axons to cortical
Subcortical EEG Control Centers
synapses result in depolarizing EPSPs on the
What coordinates the millions of neurons cortical surface. The summation of these EPSPs
throughout the cerebral cortex that give rise to results in a synchronized EEG.
the EEG and associated changes in the level of E lectrical stim ulation of nonspecific
alertness and sleep? Two centers, the non thalamic nuclei in animal experiments at about
specific thalamic nuclei and the brainstem 10 pulses per second has been found to trigger
reticular formation, play major roles. Several synchronized, depolarizing potentials at the
other centers are thought to be involved in the cortical surface, consistent with the role pro
onset and regulation of sleep. posed for these nuclei in control of the EEG.
Sleep can also result, suggesting a relationship
Nonspecific Thalamic Nuclei. The appearance of between the synchronized EEG and relaxation
synchronized waves, such as the alpha rhythm, or sleep.
may be related to the activity of neurons in the
nonspecific thalamic nuclei. These nuclei in Reticular Form ation. Desynchronized EEG
clude the intralaminar and midline nuclei and waves (beta waves) and alert behavior are
project to other thalamic nuclei and to diffuse associated with activity in the brainstem reticu
areas of cerebral cortex, in contrast to specific lar formation, a column of neurons ascending
thalamic nuclei, such as the ventroposterola- in the central core of the brainstem and
teral and lateral geniculate, which project to containing numerous synapses (figure 11-5A).
specific, limited areas of cortex (figure 11-5A ). An early clue to this association was that after
The nonspecific projections are via nerve fibers epidemics of encephalitis lethargica, or sleep
that synapse on superficial cortical layers, par ing sickness, a disease often characterized by
ticularly onto the apical dendrites of pyramidal excessive sleeping and reduced wakefulness,
cells, while the specific projections synapse patients were found to have lesions in part of
more densely on the middle layers. The non the reticular formation surrounding the cere
specific thalamic nuclei and their cortical pro bral aqueduct in the midbrain (the eye-muscle
jections are called the nonspecific, generalized, or paralysis often associated with the disease was
diffuse thalamocortical system. probably due to invasion of the nearby
211 Cerebral Physiology and Higher Functions
SENSORY
INPUT
■EEG
-LIT \ psp
Figure 11-5. (A) Subcorneal areas that coordinate and other sensory pathways. (B) A model for the
the EEG: the nonspecific thalamic nuclei (NSp.), origin of the alpha rhythm. Intracellular recordings
and the reticular formation m the brainstem. from principal cells of thalamus (those which project
Neurons in die nonspecific thalamic nuclei project to up to cortex) are shown at lower right; e = excita
diffuse areas of cortex, in contrast to a specific tion, i = inhibition. Sources: G.M. Shepherd. The
thalamic relay nucleus (Sp.), such as the medial Synaptic Organization of the Brain (2nd e<L).
geniculate, which projects to a limited sensory area New York: Oxford University Press, 1979; and P.
of cortex, such as the primary auditory area. (Syn Andersen and S. A. Andersscm. Physiological Basis
apses of auditory pathway in inferior colliculus not of the Alpha Rhythm. New York: Appleton-Cen-
shown.) In the reticular formation, neurons are ac tury-Crofts, 1968.
tivated by axon collaterals from the lateral lemniscus
212 Cerebral Physiology and Higher Functions
oculomotor nuclei). Since then it has been controls consciousness and EEG activity by
found that damage to the brainstem reticular means of ascending fibers is known as the
formation through cerebrovascular accidents, ascending reticular activating system (ARAS).
tumors, or increased pressure on the brainstem
from other causes can cause loss of con Sleep-Inducing Centers. We have already men
sciousness. In comparison, large areas of cere tioned that low-frequency stimulation of non
bral cortex can be removed without causing loss specific thalamic nuclei can result in sleep. In
of consciousness. addition, several other areas may be important
Reticular neurons can be activated by audi in inducing sleep. A basal forebrain area,
tory inputs carried up the lateral lemniscus, including the anterior part of the hypothalamus
somatic sensory inputs carried up the medial and an adjacent area of cerebral cortex, has
lemniscus, and by other sensory inputs, all of been found to trigger slow-wave sleep under
which converge on reticular neurons (figure certain conditions. Animal lesion experiments
11—5A). Reticular activation experimentally show a brainstem nucleus called the raphe
leads to a low-amplitude, high-frequency nucleus, a major source of the presumed
(desynchronized) EEG, along with alert, wake neurotransmitter serotonin, to be necessary for
ful behavior. At least in part, these effects are slow-wave sleep. The role of serotonin is
transmitted by projections from the reticular suggested by experiments in which its synthesis
formation to the nonspecific thalamic nuclei, is blocked by drugs, resulting in insomnia.
which then affect cortical activity. Lesions at Another brainstem nucleus, the locus ceruleus
the upper end of the reticular formation block in the pons, which contains a great deal of
these effects, resulting in somnolence and a norepinephrine, seems to be necessary for REM
synchronized EEG. sleep.
With increasing dosage of general anesthet
ics [2 ], both the reticular formation and its Abnormal EEG Patterns in Epilepsy
projections to the thalamus respond less to
The diagnosis of epilepsy is a major clinical
sensory stimulation, as shown by neuronal
application of the EEG. Epilepsy has been
recordings. EEG desynchronization and behav
defined as “an occasional, an excessive and a
ioral alerting are also diminished; these and
disorderly discharge of nervous tissue” [19].
several other effects of general anesthetics have
This abnormal electrical discharge occurs in
been attributed to actions on the reticular
the cerebral cortex or thalamus and (possibly)
formation that modify sensory input to the
brainstem reticular formation. The physical or
cortex. Direct effects of anesthetics on cortical
mental manifestations of the abnormal neural
neurons may also occur.
activity are known as seizures. Epilepsy may be
In addition to its effects on consciousness
generalized, involving wide areas of the brain at
and the EEG, the brainstem reticular formation
the same time, or partial, localized to small
has several other functions: it receives projec
areas. A text on clinical neurology should be
tions from the spinothalamic pain pathway
consulted for a more complete treatment of this
(chapter 5), it is a brainstem motor area
subject; only some forms of epilepsy are de
controlling posture and movement via descend
scribed here, and seizures can occur without
ing pathways to the spinal cord (chapter 9 ),
epilepsy.
and it contains control centers for respiration,
heart rate, vasomotor activity and various Generalized Epilepsy:
functions of the autonomic nervous system. Grand Mai. This is probably the most dra
That portion of the reticular formation that matic form of epilepsy. The patient loses
213 Cerebral Physiology and Higher Functions
originate in the synaptic activation of neurons sociated with language disorders. Obviously,
in the middle layers of the postcentral gyrus, oral paralysis interferes with speaking and
where most of the fibers ascending from the deafness interferes with hearing words, but
thalamic relay nucleus arrive. Later waves there are other types of language disorders, or
(positive peaks at about 50, 100, and 200 msec, aphasias, that are not due to simple motor or
and a negative wave at about 140 msec) are sensory deficits.
thought to be due to less direct pathways to the
cortex or to the activation of cortical associa The Language-Dominant Hemisphere. Over 90%
tion areas that surround the primary receiving of the population is right-handed and manipu
area in the postcentral gyrus. lates pens, forks, and so on under control of the
In addition to this somatosensory evoked left motor cortex. Almost all of these right
potential (SEP) it is also possible to record an handers also have left hemispheres dominant
auditory evoked potential (AEP) and a visual for language, as indicated by the location of
evoked potential (VEP). All these responses lesions producing language disorders. Of left-
provide information about the physiological handed people, some have left-hemisphere
function of human sensory pathways by means dominance like right-handers, others have
of a noninvasive procedure. They have been cerebral dominance simply reversed so that
applied to neurological diagnosis; one example their right hemisphere is dominant, and a third
is that reduced conduction velocity in the optic group have mixed dominance, in which control
nerve may be associated with increased latency of language is shared by both sides.
of particular peaks in the visual evoked poten
tial. Other applications are to research in Expressive (Broca’s) Aphasia. Paul Broca, in
sensory physiology, psychology, and psychiatry. 1861, found that damage to a region of the left
For example, the amplitudes of the late waves frontal lobe gives rise to a disorder in speech
are either augmented (increased) or reduced production, commonly called expressive
with greater stimulus intensity, and these two aphasia. Broca's area (figure 11-8), also called
response types have shown some correlation the frontal speech area and including Brod-
with diagnostic categories in psychiatry [5]. mann’s area 44, is adjacent and sends projec
tions to the facial and pharyngeal areas of
motor cortex. But with lesions limited to
Broca’s area, these areas of motor cortex and
Language and Hemispheric Specialization the corresponding muscles can still function in
Language is one of the most uniquely human eating and singing tunes. Speaking may be
abilities. It depends on cerebral structures and either absent or limited, labored, and non
processes that violate the rule of bilateral fluent, and is often characterized by word-
symmetry— they are generally concentrated in finding difficulties.
one cerebral hemisphere rather than the other.
The other hemisphere may specialize to some R eceptive (W ernicke’s) Aphasia. In 1874,
extent in certain higher functions that do not shortly after Broca’s discovery, Carl Wernicke
involve language. reported that a different lesion in the left
hemisphere produced a different language dis
Language Abilities and Disorders order. Wernicke’s area (figure 11-8), also called
the temporal speech area and including part of
Brain mechanisms underlying language were Brodmann’s area 22, adjoins the auditory cor
discovered on the basis of brain lesions as tex on top of the temporal lobe and receives
216 Cerebral Physiology and Higher Functions
Figure 11-8. Broca’s area, Wernicke’s area, and the information from the neighboring visual associ
angular gyrus on the left hemisphere. F, P, T , and ation cortex and to send information on to
O indicate frontal, parietal, temporal, and occipital
Wernicke’s area for further processing. Thus,
lobes, respectively. Source: Geschwind, N. Speciali
zations of the human brain. Sci Am. 241:180-201, reading a sentence and then repeating it out
1979. loud probably involves transmission of neural
information from visual cortex, to visual associ
ation cortex, to the angular gyrus, to Wer
information about auditory input from it via
nicke’s area, to Broca’s area, to the oral area of
short nerve fibers. The main language disorder
the motor cortex. The pathway from Wer
is a difficulty in understanding speech that
nicke’s area forward to Broca’s area is a fiber
extends to written language as well. Patients
bundle known as the arcuate fasciculus [9 ].
often produce speech fluently but it contains
inappropriate words and is relatively incom Left-Right Hemisphere Specialization. Do lawyers
prehensible; for example: “I came here but the and artists exercise different brain functions in
restrictions from the roof falling in on the bam their work? It could be argued that lawyers,
made the army difficult.” Because of the pa using verbal and logical skills involving step-
tient’s difficulty in understanding the meaning by-step analysis of information, are demonstrat
of language, he is generally not aware of his ing largely left-hemisphere function, while
own errors.
artists, using other abilities, such as perception
of visual patterns as a whole, are demonstrating
Alexia. The loss of the ability to comprehend a greater amount of right-hemisphere function.
written language due to a brain lesion is called This sort of speculation, although overly sim
alexia (although inability to read without plistic, is one product of studies on left versus
known brain lesions may be given the same right hemisphere specialization. Some methods
name). When spoken language and calculation of study are the following:
ability remain relatively intact, the lesion is
generally in the angular gyrus in the posterior 1. Patients with unilateral brain lesions have
parietal lobe on the left side (figure 11-8). The been compared with normal subjects and
angular gyrus is thought to receive visual those with similar lesions on the opposite
Reading and the Brain
.
Revised
Functional MRI. Advances in brain
scanning have resulted in a revised view of
how the brain works during reading.
Functional Magnetic Resonance Imaging, or
fMRI, is a brain scanning method that allows
us to detect the function of specific brain
regions in people while they do various
tasks. The method resulted from observing
that when the neurons in a brain region
become active, they use more oxygen for
metabolic activity. After a short delay, this
leads to a local increase in oxygenated
blood flow to that region. When the head is
within a special magnetic field, the localized
increase in oxygenated blood (Blood
Oxygen Level Dependent or BOLD
response) produces an image that can
change second by second, from which we
can infer localized changes in neural
activity. Needless to say, there are many
theoretical and technical steps in this
process.
side. Lesions in the language areas are ponding to the words can be picked out with
examples already discussed. the left hand. An explanation is that left-
2. Several patients have had the corpus cal hemisphere input can be processed by centers
losum cut surgically for therapeutic reasons. for language comprehension and production in
Subsequent study of the abilities of these the same hemisphere; right-hemisphere input
“split-brain” patients has led to conclusions can be processed to a limited extent for com
about the capacities of left and right hemis prehension in the same hemisphere but cannot
pheres when not directly connected to each cross over to the more capable centers for
other. comprehension and expression in the left
3. In normal as well as brain-damaged subjects, hemisphere. Similar tests have shown the right
visual input can be limited to the left or hemisphere to be superior to the left in some
right half-field, sending input initially to the nonlinguistic tasks, such as copying a drawing.
opposite hemisphere. Similarly, auditory in
put can be provided to the left or right ear, Right-Hemisphere C om p ared with L e ft -
sending input primarily to the opposite Hemisphere Function. The different methods
hemisphere because of the somewhat greater support the concept that one hemisphere,
effectiveness of crossed as opposed to un generally the left, is dominant for language,
crossed fibers in the auditory pathway. although they also show a limited amount of
4. Cortical electrical activity, both EEG and language comprehension in the right hemi
evoked responses, can be recorded from left sphere. Arithmetic ability seems to be repre
and right scalp electrodes. A new method, sented on the same side as language. The
the positron-emission tomography (PET) results also suggest (although there are some
scan, provides images of metabolically active differences of opinion) that the right hemi
areas within the functioning human brain. sphere plays a major role in the perception,
recognition, and interpretation of visual pat
The split-brain studies have attracted par terns or objects in space, the person’s own
ticular interest (figure 11-9). They utilize the body, music, and the meaning of situations.
knowledge that the left hemisphere receives Lesions in the association cortex of the right
input from the right visual half-fields and posterior parietal lobe (as described in chapter
controls the right arm, while the right hemi 5) may result in defective perception and
sphere receives input from the left visual half neglect of visual objects and of the patient’s
fields and controls the left arm. In right-handed own body, particularly on the left side. Several
patients, when visual input is restricted to the neuropsychological tests have shown that prob
right visual fields and thus the left hemisphere, lems in visual recognition are more severe
objects can be named and words can be read following right-sided than left-sided lesions. In
aloud or written down, confirming the lan- normal subjects also, recognition of visual
guage-dominance of the left hemisphere. patterns is often superior when they are pro
When visual input is switched to the left visual jected to the right hemisphere rather than to
fields and thus the right hemisphere, objects the left hemisphere (from the opposite visual
cannot be named and words cannot be read half-fields).
aloud or written down. However, the right Recent neurophysiological experiments [30]
hemisphere shows recognition of objects and may be related to the right-parietal or neglect
understanding of spoken and written words to a syndrome. Single neurons in posterior parietal
limited extent as long as linguistic responses are cortex of monkeys were found to discharge only
not demanded; for example, objects corres when a visual stimulus appeared in part of the
218 Cerebral Physiology and Higher Functions
Figure 11-9. (A) In experiment with spkt-brain pa right hemisphere in right-handed subject Source:
tient, names of objects flashed in left half-field acti Sperry, R. Lateral specialization in the surgically sep
vate right visual cortex and can be read, understood, arated hemispheres. In Schmitt P.O., and Worden,
and picked up by left hand, but cannot be spoken. F.G. (Eds.). The Neurosciences: Third Study
(B) Schema showing visual input, auditory input, Program. Cambridge, Mass.: MIT Press, 1974.
hand control, and higher specializations of left and Reprinted with permission.
219 Cerebral Physiology and Higher Functions
contralateral visual field and when the monkey different patients and under different environ
attended to the stimulus, a combination often mental conditions. Two simultaneous behav
followed by eye movement to fixate the iors, such as speaking and tapping, or tapping
stimulus. Such neurons may be partly responsi different rhythms with each hand, may or may
ble for visual perception associated with atten not interfere with each other, possibly as a
tion, and their loss may be associated with result of the spread of activity between the
visual neglect. cortical areas involved. The thalamus and
Certain aspects of musical perception have other subcortical regions also play a role in
also been related to right-hemisphere function. cerebral systems that govern complex behavior.
Patients with right temporal lobe lesions were Such examples suggest that cerebral activity in
inferior to patients with left temporal lobe language and other higher processes is complex
lesions in perception of melody and other and our understanding of it incomplete.
nonlinguistic auditory stimuli. In addition,
experiments in which separate stimuli entered
left and right ears simultaneously (dichotic
listening) showed a left-ear superiority for Memory
recognizing pitch and chords, implying a right-
Suppose you have to look up a seven-digit
hemisphere superiority. Similar results have
telephone number. If there are no distractions,
been observed during a test sometimes em
you may retain the image of the number for an
ployed prior to neurosurgery: when the right
instant after you read it and remember it for a
hemisphere is anesthetized, the patient may be
few more seconds while dialing it or repeating it
able to speak adequately but unable to hum a
to the operator. If you had to, you could
tune.
probably memorize the number for a longer
Finally, a right hemisphere role in recogni
period of time, by repeating it over and over
tion of the meaning of situations is suggested by
again or possibly associating the number se
several lines of evidence. Patients with right-
quence with some idea. At any stage, you may
hemisphere lesions have shown greater diffi
well forget the number, especially if you try to
culty interpreting cartoon sequences and
memorize several others.
complex pictures than patients with left-
These everyday experiences of memory can
hemisphere lesions, although they can identify
be associated with memory processes postulated
individual parts of the pictures. They may also
on the basis of objective psychological experi
deny or minimize their own illness, and display
ments on human memory. As diagrammed in
little emotional reaction in general, a tendency
figure 11-10 [3,22], visual or auditory input
called flat affect.
can be thought of as first entering a sensory
store, where it is stored with little change for up
Localization and Interrelation o f Cerebral Activity. to one second— corresponding in our example
Although a number of higher functions have to the image of the telephone number. With
been localized to particular areas of the cere out attention this information is lost, but with
brum, the interrelation of brain areas and attention it can be transferred to short-term
activities makes the picture more complex [13]. memory (STM). Without either effort or dis
Recovery of language functions after left- tractions, information can remain in STM for
hemisphere lesions may reflect compensatory 15 to 30 seconds, corresponding in our example
activity of a different part of the brain, for to the time it takes to dial or repeat the
example, the right hemisphere. The effects of number. Finally, by repeating (rehearsing) the
apparently similar brain lesions may vary in information several times, or associating it with
720 Cerebral Physiology and Higher Functions
J-Rehearsal Retrieval
Sensory Short-term Long-term
Sensory Encoding
Storage Memorv Memory
Input Consolidation
Attention
II INI
-Decay (msec) Decay(sec) -Decay (?)
Displacement r Interference
Figure 11-10. A multistage model of human mem- called sensory store (sensory storage, iconic
ory. Downward arrows show loss of information. store for vision, echoic store for audition).
Adapted from Atkinson and Shifrin [3] and Loftus
and Loftus [22].
Short-Term Memory
some idea, it can be stored in long-term memory Although most of the information deluging our
(LTM) for a period of minutes to years— like sensory store from various sensory inputs is lost,
your own telephone number and a few others focusing attention on it can transfer some of it
that are important to you. to STM. STM can be tested by giving someone
The stages of sensory store, STM, and LTM a list of digits and asking him to repeat back as
were based on experiments on human memory many as possible immediately afterward. This
and do not have known anatomical or physio procedure, known as a digit-span test, is part of
logical correlates, although it is possible to mental-status examinations (given when some
make some educated guesses. impairment of consciousness or thought process
is suspected), and of IQ tests, such as the
Wechsler Adult Intelligence Scale. Adults can
Sensory Store typically repeat back about seven digits cor
A slide displayed for only 50 msec contains rectly (corresponding to the number of digits in
rows of letters, for example, a telephone number), with a range of five to
nine, or 7 ± 2. The same limit to the capacity
T D R of STM applies to letters, words, or other items
L O N of information. STM has been compared to a
K S T section of computer memory, with five to nine
locations containing slots to be filled with
Shortly after the slide goes off, a tone sounds, items of information.
signalling the viewer to repeat the letters in one Although rehearsing (repeating) a short list
of the rows, corresponding to thepitch of the of items can hold it in STM indefinitely, if
tone— high, medium, or low.If the delay rehearsal is prevented (for example, by giving
between the slide and the tone is no more than some other mental task immediately after the
one second, all the letters can generally be list is presented) the list will be totally forgot
repeated. The explanation has been that an ten in 20 to 30 seconds. Thus STM seems to
image of the visual stimulus is maintained, have a built-in limit on both capacity (7 ± 2
probably somewhere in the visual system, for items) and duration (20 to 30 seconds).
this brief period. A similar sound image is These limits govern the way information is
thought to be left by an auditory stimulus. The lost (forgotten) from STM. Because of the
system postulated for this process has been limited capacity, new items can enter and
221 Cerebral Physiology and Higher Functions
displace the old. Because of the limited dura duration can last from minutes to a lifetime. A
tion, information can be lost by simple decay if number of experiments suggest that the overall
it is not rehearsed or processed further. meaning of a written passage rather than the
literal words tend to be stored in LTM. Forget
ting from LTM is thought to be largely due to
Long-Term Memory
interference; for example, memory of one list of
How is information transferred from STM to words is interfered with by other lists presented
LTM, or consolidated? Two thought processes before or after the lists in question.
known to be effective are rehearsal and organi
zation. As you might expect, the chance of
remembering a single word tends to go up with Disorders o f Memory
the number of times the word is rehearsed [22 ].
For more-complex information, such as a scien Amnesia is the general term for loss of memory.
tific article, memory is aided by organizing the When amnesia is associated with some clinical
material, with or without rehearsal. condition or episode, it may be retrograde, in
Methods of organizing material may be di which memory is lost for events preceding the
vided into two categories. The first is by using episode by a period ranging from minutes to
some preexisting framework. An ancient exam years; anterograde, in which memory is lost for
ple, cited by the Roman writer Cicero (in De events following the episode; or global, which
oratore), was the “method of loci.” A public includes memory loss for events both preceding
speaker would imagine a familiar place, such as and following the episode.
his house, and mentally place the parts of his
speech in different places— the introduction at Disorders o f Memory Associated with Specific
the front door, the first anecdote in the living Brain Lesions. The clearest association of mem
room, and so on. When he gave the speech he ory impairment with specific brain lesions has
would then mentally walk through the house, probably been found in patients with lesions of
finding (retrieving) each part of his speech in the hippocampus and several other structures to
the corresponding part of the house. Although which it is anatomically connected [13,23].
a preexisting framework is not always available, The hippocampus is a part of the limbic system
most people recognize that organizing informa within die temporal lobe and adjacent to the
tion themselves, whether by outline, diagram, lateral ventricle. Several patients sustaining
images, or developing overall ideas about the lesions of the hippocampus, often as part of
subject, helps to remember it. Psychology surgical treatment for temporal-lobe epilepsy,
experiments have often used simple lists of have been left with a specific impairment of
unrelated words to study these effects. When memory function. Their level of consciousness,
asked to memorize ten-word lists, those stu concentration, and attention did not seem
dents who made up a story around the words impaired, their memory for long-past events
(for example, “on the chair in the kitchen was a was good, and they could perform the digit-
cup, from which a canary drank water” used to span test well, even retaining items in memory
remember the list chair, cup, canary, water) for several minutes by constant rehearsal. But
retained most of the words on the list, while new information could not be remembered for
others remembered few of the words. Similarly, any longer period if rehearsal was blocked by
mental images of parts of such a story helped to distractions. In terms of the proposed model,
remember the words. their STM appeared to function adequately,
Unlike STM, the capacity of LTM is rela their LTM was largely intact, but consolidation
tively unlimited, or at least unknown, and its or transfer of information from STM to LTM
222 Cerebral Physiology and Higher Functions
seemed to be blocked. 1 The result was an sensory store, short-term memory, and at least
inability to retain memory for experiences a large part of long-term memory appear to
occurring after the surgery (anterograde am function adequately.
nesia), and sometimes for a period of months to
a few years before (a variable amount of Disorders o f Memory Associated with More Gen
retrograde amnesia). These lesions were bilat eralized Brain Dysfunctions. A number of mem
eral (on both sides). ory disorders are not associated with specific
Unilateral lesions of the temporal lobe, brain lesions, at least according to current
including the hippocampus, have been re knowledge. Amnesia can be induced by a blow
ported to be associated with memory loss for to the head (trauma), electroshock, epileptic
certain information only. These defects corres seizures, cerebral anoxia, or anesthesia. This
pond to the different functions of the two amnesia is often transient and pertains to
cerebral hemispheres, in that left-temporal events during, shortly before, and sometimes
lesions impair verbal memory, while right- shortly after the episode.
temporal lesions impair nonverbal memory, A blow to the head may cause concussion,
such as recognition of visual designs seen which typically involves a transient loss of
earlier. Although many of these patients had consciousness and of memory for events shortly
entire temporal lobes removed, the effects of preceding the injury. Observations of several
more-limited lesions suggest the critical role of athletes immediately after concussion in a
hippocampal lesions in causing the defect. football game showed that they recalled such
In patients with Korsakoff’s syndrome, a events when questioned immediately after the
memory disorder accompanies a wide range of injury (indicating STM function) but were
conditions including alcoholism, nutritional unable to retain the information when ques
deficiency, poisoning, and cerebrovascular le tioned 3 to 20 minutes later.
sions. Patients have an impaired ability to
consolidate new memories (anterograde am Physiological Mechanisms o f Memory
nesia) as in hippocampal damage, together
with a variable loss of earlier memories (retro In models of memory based on both human and
grade amnesia), although very early memories animal research, short-term memory is thought
tend to be preserved. Lesions have been found to involve temporary, easily disrupted changes
in certain thalamic nuclei (the dorsomedial or in neural activity or excitability. One possible
anterior), a portion of the hypothalamus (the change is the activation of closed circuits of
mamillary bodies), and in some cases the neurons, in which, for example, neuron A
hippocampus. These structures are part of a activates neuron B which then reactivates
neural circuit projecting from hippocampus via neuron A, and so on. Such reverberating circuits
the fornix to the mamillary bodies, from there are similar to those thought to occur among
to thalamic nuclei, and then to the cingulate spinal intemeurons during afterdischarge of the
gyrus of the cerebral cortex. flexor withdrawal reflex; they could be main
These cases suggest that the hippocampus, tained for several seconds but also could be
along with some connected structures, plays a disrupted by another stimulus. Another possi
role in memory consolidation. This process ble short-term change is found in spinal
may be impaired even when consciousness, the motoneurons involved in the stretch reflex. If
they are activated repetitively at high fre
'There is recent evidence, however, that some skills can be
learned, and presentation of cues can improve recall of
quency for a few seconds, a procedure called
verbal material 124]. tetanic stimulation, their postsynaptic response
223 Cerebral Physiology and Higher Functions
The Mental Status Examination includes an assessment of the higher functions discussed
in this chapter. These topics can be organized in the following outline. Sample tasks are
included.
Level of consciousness
alert, lethargic, stuporous, comatose
Attention
ability to recall a series of digits, subtract serial sevens from 100, spell WORLD
backwards, concentrate on interview
Language
comprehension, expression, repetition, reading, writing, naming
Visual-motor function
ability to copy diagrams, construct block designs
Memory
immediate: repeat series of digits; repeat three unrelated words
recent: repeat words after several minutes delay
remote: name most recent war in which the US was involved; name past schools
Observations
appearance, ability to relate, eye contact, psychomotor activity
Fund of information
names of recent presidents, location of states (depends on learning and memory)
Orientation
tine, place, person, situation (depends on learning and memory to update information)
Calculations
subtract serial sevens from 100, perform simple arithmetic problems
Thought process and structure
productivity, speed, spontaneity of ideas; note incoherence, blocking, perseveration,
loose associations, tangentiality, etc.
Thought content
preoccupations, feelings of detachment from environment or self, grandiosity,
delusions, hallucinations, etc.
Mood and affect
depression, anxiety, euphoria, anger; flat, blunted, labile, or inappropriate affect
Abstract thinking
interpret proverbs such as “Rome wasn’t built in a day”, or “don’t cry over spilt
227 Cerebral Physiology and Higher Functions
Review Questions
2. Diagram and describe a model for the origin of the EEG in cortical neurons and
in subcortical EEG control centers.
3. Diagram normal EEG patterns and describe their association with varying levels
of alertness and sleep.
Cerebral Physiology and Higher Functions
Compare generalized with partial epilepsy. Briefly describe grand mal, petit mal,
focal motor, and psychomotor forms of epilepsy.
es to sensory stimulation.
229 Cerebral Physiology and Higher Functions
8. Compare expressive with receptive aphasia, in terms of both behavior and brain
lesions.
9. Define alexia. Describe its relationship to the angular gyrus and the interaction of
that structure with surrounding areas.
10. List and briefly describe the methods of studying left-right hemisphere special
izations.
230 Cerebral Physiology and Higher Functions
11. List and briefly describe the higher functions for which either a left- or a right-
hemisphere specialization has been reported.
12. Speculate about the probable results of the following experiment With a group of
normal subjects, one word was briefly flashed on a screen in the left visual field,
while simultaneously another word was flashed in the right, after which the sub
jects repeated aloud the word they recognized. Which word was repeated more
often in a series of such word-pairs?
13. Diagram a model for human memory including sensory store, short-term memo
ry, and long term memory.
231 Cerebral Physiology and Higher Functions
14. Prepare a table comparing short-term memory with long-term memory in terms
of capacity, duration, how information enters, and how information is lost
16. Describe the association of amnesia with brain lesions and brain dysfunctions.
14. Prepare a table comparing short-term memory with long-term memory in terms
of capacity, duration, how information enters, and how information is lost
16. Describe the association of amnesia with brain lesions and brain dysfunctions.
233
234 Glossary
Beta waves EEG waves above 13 Hz cortex; it is more responsive to the orientation
Binocular vision Simultaneous use of both eyes to of a stimulus than to the exact position of the
yield a single image stimulus within the receptive field
Bipolar cells One of the five types of retinal cell Complex periodic sound Sound composed of sev
Blind spot Small area in the visual field that is eral frequencies that yield a complicated but
unresponsive to visual stimuli, approximately recurrent vibratory pattern
15 degrees lateral to the fixation point, corre Conduction Movement of action potential along a
sponding to the area of the retina at which the nerve or muscle fiber
optic nerve exits and photoreceptors are absent Cones Photoreceptors responsible for color
Bone conduction Transmission of sound waves to (photopic) vision and high visual acuity
the cochlea through the bones of the skull Consolidation A process of fixating information by
rather than through the middle ear, often used transferring it from short-term to long-term
to determine the nature of a hearing loss memory
Bradykinin A polypeptide released by tissue dam Convergence Bringing together to a common
age, causing pain by activation of C fibers point; applied to ( l) the refraction of light rays;
Brainstem auditory evoked potential Averaged and (2) die synaptic connections of neurons
response to auditory stimulus, arising from Cornea The transparent structure comprising the
brainstem auditory pathways foremost portion of the eye
Broca’s area Part of the frontal-lobe association Corpus callosum Large interconnecting band
cortex, in front of the facial motor area, (commissure) of fibers between left and right
important in language expression cerebral hemispheres
Brodmann’s areas A division of cerebral cortex Cortical column Refers to the cellular organization
into about 50 areas based on cytoarchitecture of the cerebral cortex. In the visual area, cells
Caloric test Test of vestibular function by placing in a column perpendicular to the surface have
warm or cold water in the ear, yielding nystag the same receptive-field axis orientation; also,
mus in the normal state at right angles to the orientation columns are
Central sulcus (sulcus of Rolando) A groove alternating ocular-dominance columns each
between the motor area in the frontal lobe and predominantly influenced by one eye.
the somatic sensory area in the parietal lobe Corticospinal tract Tract descending from cerebral
Cerebral aqueduct Ventricular passage in the cortex through medullary pyramids to spinal
brainstem connecting the third and fourth cord
ventricles Crossed extension Reflex extension of one leg
Cerebrum The main portion of the brain, includ associated with flexor withdrawal of the oppo
ing the cerebral hemispheres, above the brain site leg after a painful stimulus
stem and cerebellum CT scan Computerized tomography scan in which
Chemoreceptors Receptors that are responsive to the head is scanned with a moving x-ray beam
particular molecules either in liquid or gaseous and a series of horizontal sections are recon
form structed with computer processing; it provides
Cholinergic Pertaining to neurons that release an image of brain structure based on tissue-
acetylcholine as a synaptic transmitter density differences
Chorea (Gr. choros, a dance) A movement disor Curare A drug, extracted from a South American
der characterized by irregular, jerky, dancelike, vine, that blocks acetylcholine receptor sites
involuntary movements and thereby interferes with neuromuscular
Clonus (Gr. klonos, turmoil) Involuntary move transmission
ment consisting of alternating contraction and Decibel (dB) Measure of the intensity of a sound
relaxation of a muscle group Deep tendon reflex See tendon jerk reflex
Coactivation Acting together of alpha and gamma Delta waves EEG waves below 4 Hz
motoneurons Dendrite (Gr. dendrites, pertaining to a tree) Pro
Cochlea (L. cochlea, snail shell) Fluid-filled, spi cess or extension of a neuron, containing
rally coiled portion of the inner ear, located postsynaptic sites and capable of receiving
within the temporal bone and containing the neural signals from sensory receptors or other
auditory sense organ, the organ of Corti neurons
Complex cells One type of cell in the visual Denervation supersensitivity Increased sensitivity
235 Glossary
to chemical transmitters seen in muscles, Epileptic focus Discrete area of cortex from which
neurons, and glands following denervation epileptiform EEG activity originates
Depolarization Change of membrane potential Equilibrium potential Membrane potential at
from resting level toward zero or inside which there is equivalent movement of ions in
positivity both directions across the membrane, leading to
Dichotic Pertaining to both ears an equilibrium; may be predicted from the
Divergence Spreading apart from a common point; Nemst equation
applied to (1) the refraction of light rays and (2) Eustachian tube A tube that connects the middle
synaptic connections between neurons ear with the nasopharynx and permits equaliza
Dominant hemisphere The cerebral hemisphere tion of air pressures on both sides of the
most essential for language function, generally tympanic membrane
the left hemisphere Evoked potential Potential change in the CNS
Dopamine A chemical transminer at some sy evoked (elicited) by a sensory stimulus, gener
napses, and a precursor of chemical transminers ally recorded with a large electrode
(epinephrine and norepinephrine) at other syn Excitatory postsynaptic potential (EPSP) The po
apses tential change generated by excitatory synaptic
Dorsal column—medial lemniscus system The action in the postsynaptic neuron, tending to
somatic sensory pathway that includes dorsal excite the postsynaptic neuron
columns, medial lemniscus, VPL thalamic nu Expressive aphasia A language disorder charac
cleus, and somatic sensory conex, and is con terized by defective speech expression
cerned with discriminative touch and pressure, Extrafusal fibers (L. fusus, spindle) In skeletal
vibration and lcinesthesis muscle, the contractile fibers that are external
Dysarthria (Gr. arthroun, uner distinctly) Diffi to the muscle spindles and make up the bulk of
culty in aniculating words the muscle mass
Dysmetria (Gr. metron, measure) Inability to Extrapyramidal Supraspinal motor areas and their
judge distances during voluntary movement descending tracts that are not part of the
Edinger-Westphal nucleus In midbrain, parasym pyramidal tracts
pathetic component of oculomotor nucleus, Facilitation Increased effect of a stimulus on ex
governing pupillary constrictor and ciliary mus citatory synaptic transmission, when another
cles of the eye excitatory stimulus is presented (1) shortly
Efferent (L. ex, out + ferre, to bear) Referring to before or (2) simultaneously
axons that conduct impulses away from the Fast pain Earliest pain sensation, associated with
central nervous system, or the impulses conduction in A-delta fibers
themselves Feedback Pertains to a system in which some of
Electroencephalogram (EEG) Electrical activity of the output is returned to the input (Examples:
the brain, usually as recorded with electrodes on hearing oneself speak; baroreceptor response to
the scalp blood pressure changes due to cardiovascular
Electromyogram (EMG) Electrical activity arising activity)
from muscle activity Fixation point A point at which one stares to
Endolymph The fluid filling the scala media of the immobilize the eye
cochlea and the semicircular canals, similar in Flaccidity Weakness of muscles; lack of muscle
composition to intracellular fluid tone
Endorphin A polypeptide found in brain and Flexor withdrawal reflex Limb withdrawal reflex
pituitary with actions similar to the opiate drugs mediated by nociceptors
End-plate The postsynaptic region of skeletal mus Focal length Distance from center of lens to point
cle membrane at which light rays from a distant point con
End-plate potential (EPP) The depolarization re verge
corded across muscle end-plate membrane fol Focal motor epilepsy (partial motor seizure) A
lowing a nerve impulse in the presynaptic type of partial epilepsy with motor activity in a
motoneuron particular part of the body
Enkephalin Peptide composed of five amino acids Fovea (fovea centralis) (L. fovea, a pit or depres
and found in brain, with actions similar to the sion) The depression in the center of the
opiate drugs macula lutea of the retina; the area of sharpest
236 Glossary
vision, upon which the fixation point is nor brought into focus behind the retina; also called
mally focused farsightedness
Frequency Rate of repetition; the number of cycles Hyperpolarization Change of membrane potential
made by a sine wave within a second (cycles per from resting level toward a more-negative value
second or hertz) (inside of cell referred to outside)
G Conductance; the ability of ions to pass through a Hypertonus Excessive muscle tone, with increased
solution or across a membrane within an electric resistance to being stretched by an external
field source
GABA (gamma-aminobutyric acid) A probable Hypotonus Diminished muscle tone, with de
chemical transmitter in the CNS creased resistance to being stretched by an
Gamma motoneurons Small, myelinated neurons external source
innervating intrafusal muscle fibers Hz (hertz) Cycles per second (c/s)
Gating A mechanism by which synaptic transmis Inhibitory postsynaptic potential (IPSP) The po
sion in ascending tracts associated with pain tential change generated by inhibitory synaptic
sensation may be modulated; associated with action in the postsynaptic neuron, tending to
the Gate Theory of Melzack and Wall inhibit the postsynaptic neuron
Generalized epilepsy A form of epilepsy affecting Initial segment Short, unmyelinated segment of
widespread areas of brain, with seizure activity axon just distal to the axon hillock; often the
that is bilaterally symmetrical; includes grand site of impulse origination
mal and petit mal Integration Process by which a neuron combines
Generator potential Graded, localized electrical several excitatory and inhibitory synaptic inputs
potential change in a sensory receptor when before generating one or more nerve impulses
properly stimulated; also called a receptor Intention tremor Tremor that is made apparent or
potential aggravated by attempting a coordinated, volun
Glycine An amino acid and probable chemical tary movement
transmitter in the CNS Internal capsule A thick band of white matter in
Golgi tendon organs Receptors located in muscle the cerebral hemisphere, containing nerve fi
tendons and signaling muscle tension bers that connect the cerebral cortex with sub-
Graded O f differing magnitudes cortical structures, such as the thalamus
Grand mal epilepsy (French, major sickness) A and the spinal cord
type of generalized epilepsy with loss of con Internal environment The physiological condi
sciousness, convulsive movements, and postic tions within the body
tal (after the fit) depression; EEG initially Intemeuron (intrinsic neuron) A neuron whose
shows trains of spikes, then spikes and slow short processes remain within a local region of
waves, and, finally, low-amplitude slow activity the CNS
Gyrus A fold or ridge in the cerebral or cerebellar Intrafusal fibers (L. fusus, spindle) Thin, special
cortex ized muscle fibers within a muscle spindle,
Habituation The gradual waning of a response containing muscle stretch receptors
with repeated stimulation, not due to adapta Inverse muscle stretch reflex A reflex, mediated
tion of receptors or general fatigue by Golgi tendon organs, .which causes a release
Helicotrema Apical end of cochlea to which basi of tension in a muscle when its tension reaches
lar membrane does not extend; area through high levels
which scala vestibuli and scala tympani merge Iodopsin Violet-colored visual pigment found in
Hemianopia (Gr. hemi, half + ope, vision) Blind the retinal cones
ness in half die visual field; can involve one or Iris Colored, ring-shaped portion of eye, located
both eyes behind comea, containing the muscles by
Hippocampus A structure consisting of primitive, which the pupil is constricted and dilated
three-layered cortex, at the medial edge of the Ischemia Deficiency of blood supply to a part of
temporal lobe and bordering part of the lateral die body due to reduction of flow through a
ventricle; part of the limbic system blood vessel
Horizontal cells One of the five types of retinal Isoproterenol Sympathomimetic agent, which ac
cell tivates beta receptors
Hyperopia (Gr. hyper, beyond -I- ope, vision) Jacksonian epilepsy A partial motor epilepsy
Visual refractive error in which light rays are originating in the precentral motor cortex and
237 Glossary
spreading along the motor area with accom plate membrane due to spontaneous release of a
panying motor activity in corresponding parts of unit (quantum) of transmitter from the presyn
the body aptic motoneruon
Kinesthesis (Gr. kinesis, movement + aesthesis, Mitral cells The olfactory bulb cells upon which
perception) The sense of muscle movement and the nerve fibers from the olfactory receptors
position synapse and whose axons form the olfactory
Latency (latent period) The time interval between tract
a stimulus and a particular neural or muscular Monosynaptic Having only one synapse in the
response CNS
Lateral fissure (fissure of Sylvius) A deep groove Motor cortex In the frontal lobe, precentral gyrus
in the cerebral cortex, partly serving as a or Brodmann’s area 4; origin of corticospinal
boundary for the temporal lobe tract
Lateral spinothalamic tract A spinal tract ascend Muscarinic Pertaining to cholinergic receptors
ing to the thalamus, containing nerve fibers that are activated by muscarine
responding to pain and temperature stimuli Muscle spindle Structure found in skeletal muscle,
Law of specific nerve energies Principle that every containing intrafusal muscle fibers and recep
sensory neuron reacts to only one kind of tors for the muscle stretch reflex
stimulus and gives rise to only one kind of Muscle stretch reflex Reflexive muscular contrac
sensation. Under abnormal circumstances such tion elicited by passive longitudinal stretching
a neuron can be excited by other stimuli, but it of the muscle
still produces the same sensation; also called Muscle tone Steady-state muscle contraction
Muller’s law or the law of specific irritability when at rest or when passively stretched by an
L-dopa (L-dihydroxyphenylalanine) A metabolic external force, of reflex origin
precursor of dopamine, given as a therapeutic Myelin sheath A sheath formed of layers of
agent in Parkinson’s disease Schwann or glial cell membranes surrounding
Limbic system (L. limbus, border) An intercon myelinated axons, with high resistance and low
nected system including the cingulate and capacitance
parahippocampal gyri, the hippocampal forma Myopia Visual refractive error in which light rays
tion, and parts of die amygdala, hypothalamus are brought into focus in front of the retina; also
and thalamus, concerned with visceral and called nearsightedness
emotional responses and with memory Myotatic reflex (Gr. mys, muscle + temein, to
Locus cenileus (L. locus, place + coendeus, dark stretch) The muscle stretch reflex
blue) A small group of pigmented nerve cells Narcolepsy A clinical syndrome characterized by
alongside the floor of the fourth ventricle attacks of uncontrollable sleep in inappropriate
Long-term memory (long-term store) A hypothet situations, often with rapid onset of REM
ical store or repository of information that is sleep
retained for a long period, often years Near response Reflex ocular adjustments occurring
Macula lutea (L. macula, a spot + lutea, yellow) A when gaze is shifted from a far to a near object,
spot at the posterior pole of the eye, containing including accommodation and pupillary con
the fovea centralis at its center striction
Mechanoreceptor Sensory receptor that responds Neocerebellum (Gr. neos, new) The newest part
to mechanical stimuli, such as pressure and of the cerebellum to appear phylogenetically,
displacement receiving major input from the cerebral cortex
Mental-status examination An evaluation of a via the pons, and helping to regulate voluntary
patient’s mental state, often in standardized movements
form, including assessments of orientation, Neuromuscular junction Synapse between moto
memory, delusions, hallucinations, emotional neuron and muscle fiber
state, and cooperation Nicotinic Pertaining to those cholinergic receptors
Middle-ear deafness Deafness resulting from inter that are activated by nicotine
ference with the acoustic transmission of sound Nociceptors (L. nocere, to injure + capere, to
within the middle ear; also called conductive receive) Sensory receptors that are responsive
hearing loss to potentially injurious stimuli
Miniature end-plate potential (MEPP) The small Nodes Nodes of Ranvier; areas of exposed axonal
depolarization recorded across the muscle end- membrane between myelinated sections
238 Glossary
Non-REM (NREM) sleep Stages of sleep without Parkinson’s disease (paralysis agitans) A motor
rapid eye movements; slow-wave sleep disorder characterized by a slowing of voluntary
Nonspecific thalamic nuclei A group of thalamic movement, masklike face, and resting tremor
nuclei, including the midline and intralaminar Partial epilepsy A form of epilepsy with seizures
nuclei, that project to widespread areas of beginning locally; focal seizures
cerebral cortex and exercise a control over their Perilymph Fluid filling the seala vestibuli and scala
rhythmic electrical activity; the nuclei and tympani, nearly identical in composition to
their cortical projections are often called the cerebrospinal fluid
generalized or diffuse thalamocortical system PET scan Positron emission tomography scan; a
Norepinephrine (NE, noradrenaline) Chemical computerized radiographic technique that
transmitter, synthesized from amino-acid pre shows differences in metabolic activity in vari
cursor, released by postganglionic sympathetic ous areas of the brain
neurons, adrenal medulla and some neurons in Petit mal epilepsy (French, minor sickness) Gen
die CNS eralized nonconvulsive seizures generally
Nystagmus Involuntary, repetitive eye movement characterized by brief loss of consciousness and
in a horizontal, vertical, rotatory (or mixed) 3-per-second spike-and-wave EEG activity
direction Phase locking The preferential response (firing) of
Olfactory bulb and tract The bulb, located on the auditory neurons to one particular phase of the
inferior face of the frontal lobe, is the site of stimulating sound wave
synapse for fibers from the olfactory epithelia Phasic (Gr. phasis, an appearance) Brief or inter
and the origin of the olfactory nerve (tract) mittent
Olfactory sensory cells The receptor organs for Phenothiazine drugs A group of drugs, including
the sense of smell chlorpromazine, given as therapeutic agents in
Opiate drugs A group of drugs, including mor schizophrenia (and other conditions)
phine, codeine, and heroin, that relieve pain as Phenylephrine Sympathomimetic drug that acti
well as causing euphoric and other CNS effects vates alpha receptors
Optic disk Point of entrance on retina of optic Photopsin See iodopsin
nerve and blood vessels Photoreceptors Sensory receptors that respond to
Organ of Corti The sense organ of hearing, located light— the rods and cones of the eye
on the basilar membrane within the cochlea, Pitch The property of a sound ranging from high to
and containing sensory hair cells and auditory low and determined by the frequency of the
nerve endings sound waves
Ossicles The three small, interconnecting bones Place theory Theory that the response to a given
(maleus, incus, and stapes) of the middle ear frequency arises from the place of maximal
which form a system of mechanical levers for vertical movement of the basilar membrane for
efficient transmission of sound energy from the that frequency, the orderly arrangement of such
tympanum to the oval window places along the basilar membrane, and the
Oval window Small, membrane-covered opening associated neural responses
into the inner ear, nearly closed by the foot Plantar reflex A reflex elicited by stroking the
plate of the stapes, and the point at which outer aspect of the sole with a blunt object, the
sound energy is transmitted into the inner ear normal adult response being flexion of toes and
Pacinian corpuscle Mechanoreceptor, sensitive to foot
light pressure and vibration, widely distributed Polysynaptic Having several synapses in the cen
throughout the body tral nervous system
Paleocerebellum (Gr. palaios, ancient) A primi Positive supporting reaction A reaction to cutane
tive part of the cerebellum, receiving input ous stimulation of the palms of the hands or
from the spinocerebellar and other sensory soles of the feet, involving an extensor thrust
tracts, that helps to control posture and Postcentral gyrus A fold or ridge immediately
locomotion behind die central sulcus, containing the pri
Parasympathetic nervous system Portion of the mary somatic sensory area
autonomic nervous system, with efferent nerve Postganglionic neurons Autonomic neurons
fibers originating in certain cranial nerves and whose cell bodies are in the autonomic ganglia
sacral segments of the spinal cord and whose axons terminate upon effector organs
Paresthesia Abnormal sensation, such as tingling Postsynaptic neuron The neuron that is excited or
239 Glossary
inhibited by another neuron across the synaptic the impingement of an appropriate stimulus will
cleft cause the response of a neuron in the sensory
Post-tetanic potentiation An enhanced neuronal pathway
response to a standard stimulus for a prolonged Receptor (1) The peripheral terminal of a sensory
period of time following a tetanic (rapidly neuron or a specialized cell such as a rod or
repeated) stimulus cone, that responds to various environmental
Precentral gyrus A fold or ridge immediately in stimuli and transduces them into electrical
front of the central sulcus, containing the signals; (2) A molecule that combines with a
motor area specific chemical substance, such as a transmit
Preganglionic neurons Autonomic neurons with ter
cell bodies in the CNS and whose axons Receptor potential Membrane potential change in
terminate in the autonomic ganglia a receptor, caused by an appropriate stimulus;
Premotor cortex In frontal lobe, Brodmann’s area generally a depolarization
6, anterior to motor cortex
Reciprocal inhibition Arrangement of neuronal
Presbyopia (Gr. presbys, old + dps, eye) Hyper connections such that excitation of one group
opia secondary to loss of accommodative power
of neurons produces inhibition of the antagon
and elasticity of lens, often accompanying aging
istic group
Presynapdc inhibition Inhibition arising from the
depolarization and consequent reduction of Referred pain Pain sensed as coming from one site
transmitter output of presynaptic nerve end but really originating in another site, generally
ings, resulting from the action of axoaxonal an internal organ
synapses Reflex Involuntary response to a stimulus, involv-
Presynaptic neuron The neuron that excites or ing receptors, afferent impulses to the central
inhibits another neuron across the synaptic deft nervous system, efferent impulses from the
Pretectal nucleus A small nucleus in the mid- central nervous system, and an effector re
brain, receiving afferents from the retina and sponse. Reflexes are predictable and common to
involved in the pupillary light reflex normal members of the species at the same
Principal neuron A neuron that sends a long axon developmental stage
from one brain region to another Refraction (1) The bending of light as it passes
Psychomotor epilepsy A type of partial epilepsy from one medium into another; (2) The process
combining a disordered mental state and com of determining and correcting refractive errors
plex motor activity, generally associated with of the eye
an epileptic focus in the temporal lobe Refractive surface Surface at which light is bent
Pupil The opening in the center of the iris through Refractory period (1) Absolute. The period of time
which light is admitted to the retina following a nerve impulse during which another
Pupillary light reflex Constriction of the iris in nerve impulse cannot be generated; (2) Relative.
response to light stimulating the retina The period of time, following the absolute
Pure tone The auditory signal generated by a sine refractory period, during which another nerve
wave impulse can be generated only with a stronger-
Pyramidal cell A roughly pyramid-shaped cell in than-normal stimulus
the cerebral cortex, whose axon generally car Repolarization Return of membrane potential
ries output signals from that area of cortex from a depolarized state, such as the peak of the
Pyramidal tract See corticospinal tract action potential, toward the resting potential
Raph£ nucleus A nucleus in the midline of the Resting potential The steady membrane potential
brainstem reticular formation, containing a found during the resting state, generally -5 5
high concentration of serotonin and influenc mV to —90 mV (inside of neuron referred to
ing sleep outside)
Rapid eye movement (REM) sleep A stage of sleep Resting tremor Tremor that is present at rest but
characterized by rapid horizontal eye move disappears or diminishes with voluntary
ments, desynchronized EEG, lowered muscle movement
tone, and frequent reports of dreams Reticular formation (L. reticulum, a net) A net
Receptive aphasia A language disorder charac work of neurons and synapses arranged in a
terized by defective understanding of speech column extending throughout the brainstem in
Receptive field Area on skin or retina upon which its central region, below the ventricle
240 Glossary
Retina Innermost layer of eyeball, containing the primary visceral afferent fibers of facial (VII),
photoreceptors glossopharyngeal (IX), and vagus (X) nerves
Retinal ganglion cells Final retinal cell layer, the Soma Cell body
axons of which comprise the optic nerve Somatic sensory Denoting sensations from the
Retrograde amnesia Defective memory for a period limbs and the body, exclusive of the viscera
preceding a brain lesion or other event Somatotopic mapping The orderly representation
Reverberating circuit A closed loop of synaptic- of the body on the surface of the somatic-
ally-connected neurons around which nerve sensory cortex or other level of the somatic-
impulses circulate repetitively after an appropri sensory system
ate input Spasticity Hypertonicity
Rhodopsin The purple visual pigment found in the Specific thalamic nuclei A group of thalamic
rods nuclei projecting to specific cortical areas (e.g.,
Rods Most-numerous photoreceptors, responsible the lateral geniculate nucleus, projecting to
for scotopic (night) and peripheral vision and primary visual cortex)
containing the visual pigment rhodopsin Spike ( 1) An action potential; (2) A spike-shaped,
Round window Small membrane-covered opening sharp wave in an EEG
from middle ear to inner ear, below oval Split-brain patients Patients in whom the corpus
window callosum and other commissures between the
Saccule Smaller of two sadike structures in the cerebral hemispheres have been split for
vestibular apparatus; functions similarly to ut therapeutic reasons
ricle Striatum Part of the basal ganglia, including the
Saltatory conduction (L. saltare, to jump) Con caudate nucleus and putamen
duction along a myelinated axon, in which the Substantia nigra A midbrain nucleus with motor
action potential jumps fiom node to node functions; its dark appearance is due to melanin
Semicircular canal Part of the vestibular apparatus pigment
responding to angular acceleration. There are Sulcus A groove in the cerebral cortex
three in each labyrinth, situated at approximate Summation Adding together of subthreshold po
right angles to each other tentials, either spatially or temporally
Sensorineural deafness Deafness secondary to dys Superior colliculus Midbrain structure involved in
function of the organ of Corti or the auditory ocular accommodation for viewing near objects
nerve pathway and in reflex movements of the head and eyes
Sensory store A hypothetical store or repository of for following objects in motion
information that remains for some milliseconds Supraspinal Above the spinal cord, pertaining to
after a sensory stimulus the brain
Serotonin (5-hydroxytryptamine or 5-HT) A Surround inhibition The effect of stimulation of
probable chemical transmitter an annular (ringlike) area of the skin or retina
Servomechanism A device that automatically surrounding an excitatory receptive field, lead
guides a movement by comparing actual to ing to inhibition of a neuron in the somatic
desired performance and correcting errors sensory or visual system, respectively
Short-term memory (short-term store) A hy Sympathetic nervous system Portion of the auto
pothetical store or repository of information nomic nervous system, with efferent nerve
that remains, in general, for a period of seconds fibers originating in thoracic and upper lumbar
Simple cells One type of cell found in the visual segments of the spinal cord
cortex; the receptive field of each has a specific Synapse (Gr. synapto, to join) The site at which
position and orientation two neurons approach each other across a
Sine wave A curvilinear waveform consisting of microscopic gap and at which one neuron
continuous equal alterations of polarity at a set excites or inhibits the other
rate Synaptic cleft The microscopic space between
Slow pain Later pain sensation, associated with presynaptic and postsynaptic membranes
conduction in C fibers Synaptic potential Change in membrane potential
Snellen test A test of visual acuity in postsynaptic membrane, caused by synaptic
Solitary tract Medullary tract composed of the transmission
241 Glossary
Synaptic vesicles Small sacs in presynaptic nerve Tubocurarine A neuromuscular blocking agent,
terminals, bounded by membrane and contain the active principle of curare, that competes
ing chemical transmitter with acetylcholine for receptor sites in the
Synchronized EEG waves Relatively high- muscle end-plate
amplitude and low-frequency EEG waves Tuning curve Graph of threshold sound intensity
Taste buds The sense organs of taste (gustation) versus frequency for a neuron in the auditory
Temporal lobe epilepsy A type of partial epilepsy system
arising in the temporal lobe and generally Tympanic membrane (tympanum; eardrum)
associated with psychomotor seizures Cone-shaped membrane that closes the end of
Tendon jerk reflex A phasic muscle stretch reflex the external auditory canal (meatus) and to
elicited by tapping a tendon with a reflex which the first ossicle, the maleus, attaches
hammer, thereby producing a sudden stretch of Utricle Larger of two sadike structures in the
the muscle vestibular apparatus, responding to gravita
Thermoreceptors Sensory receptors that respond tional pull and acceleration
to changes in temperature Visual acuity A measure of the ability of the eyes
Theta waves EEG waves between 4 and 7 Hz to resolve detail, distance vision being tested
Threshold (1) The smallest depolarization elicit with a Snellen chart and near vision with Jaeger
ing an action potential; (2) The smallest sen cards
sory stimulus that can be detected Visual angle The angle subtended between two
Threshold of hearing The softest sound that can be lines drawn from the eye to the extremities of
detected 50% of the time, for a specified the object in view
frequency Visual field That area which can be seen by a fixed
Tonic (Gr. tonikos, tension, tone) Pertaining to a eye
chronically active state Vitreous humor The fluid behind the lens of the
Tonic neck reflex (TNR) A reflex response in eye, also called the vitreous body
which forcibly turning the head to one side Volley theory Theory that frequency information
produces extension of the ipsilateral limbs and is coded by the timing of auditory action
flexion of the contralateral limbs potentials
Transducer Structure or device that converts one Wernicke’s area Part of the temporal-lobe associa
form of energy into another (e.g., auditory hair tion cortex, behind the auditory cortex, impor
cells and microphones both convert sound- tant in language comprehension; may also
induced mechanical vibrations into electrical include adjacent parietal area
potential changes)
242 References
The following questions are in several formats. The pacinian corpuscle and the hair follicle receptor
C ircle T or F for True-False questions,./!// in the are both
blanks where indicated, and circle the letter before A. rapidly-adapting
the one best answer for multiple-choice questions. B. slowly-adapting
C. nociceptors
T F The resting potential is largely due to D. proprioceptors
potassium conductance and the potassium
concentration ratio across the nerve membrane. Like the synaptic potential, the receptor potential:
A. can travel over several cm without loss of
T F In a nerve action potential, there is a amplitude
simultaneous increase in sodium and potassium B. has an absolute refractory period
conductance. C. has a relative refractory period
D. is graded
T F The potassium channel can be blocked by
tetraethylammonium (TEA). T F In neuromuscular transmission, the binding
of acetylcholine with postsynaptic receptor sites
If die absolute refractory period o f a neuron is 5 precedes the occurrence of a muscle end-plate
msec, its highest possible discharge frequency is: potential (EPP)
action potentials or spikes per second.
T F Presynaptic inhibition requires
A myelinated nerve fiber with a diameter of 9 hyperpolarization o f the postsynaptic cell body.
microns has a conduction velocity o f about
meters/second. The inhibitory postsynaptic potential in a spinal
motoneuron is associated with:
During the relative refractory period: A. increased sodium conductance
A. no stimuli can elicit an action potential B. hyperpolarization
B. stimuli above the normal threshold can elicit an C. depolarization
action potential D. no change in ion conductance or membrane
C. stimuli below the normal threshold can elicit an potential
action potential
D. only stimuli at the normal threshold can elicit an For the following 2 questions, match each item with
action potential the correct observation below and fill in the blank.:
_Referred pain
The action potential: Gating
A. can be slowed in conduction velocity by
demyelinating disease A. Striking the elbow can lead to tingling that
B. is mainly useful for conducting neural messages appears to come from the little finger..
over short distances only because it dies away after a B. Vibration and massage can sometimes reduce
few millimeters pain sensation from a nearby area.
C. is graded in size depending on the size of the C. A small area on the fingertip can be the receptive
stimulus field for a cutaneous mechanoreceptor.
D. can occur in receptors but not in axons D. A myocardial infarction may be associated with
pain that appears to come from the left arm.
A compound action potential was recorded over a
patient's median nerve at the wrist, 5 msec after Which o f the following is TRUE?
electrical stimulation o f the index finger, 150 mm A. Type A-delta fibers have a conduction velocity
away. The conduction velocity is: of about 25 m/sec and are responsible for fast pain.
A. 150 m/sec
B. Type A-beta fibers have a conduction velocity of A lesion of parietal association cortex, limited to
5 m/sec and are responsible for the inverse stretch areas 5 and 7, on die right side, can lead to:
reflex. A. loss o f two-point tactile discrimination on the left
C. Type C fibers have a conduction velocity o f 100 side o f the body
m/sec and are responsible for slow pain. B. receptive aphasia
D. Type A-alpha fibers have a conduction velocity C. neglect of the left side o f die body and the left
of about 100 m/sec and are responsible for visual fields
temperature sensation. D. visual field loss in the left half of the visual fields
of both eyes
Aspirin is thought to exert its analgesic effect mainly
at which site? Three years prior to admission, a patient noted the
A. receptor loss o f pain and temperature sensation in die left foot
B. peripheral nerve axon and lower calf. A lesion in which o f the following
C. spinal cord synapses areas o f the cord is the most likely cause?
D. reticular formation A. Right posterior column
B. Left lateral spinothalamic tract
Pain sensation: C. Right lateral spinothalamic tract
A. may be reduced by analgesics D. Left anterior horn
B. involves the more slowly conducting nerve
fibers A normal afferent nerve fiber in a human peripheral
C. may be elicited by exercising a muscle which is nerve conducts an impulse over a distance of 100 mm
deprived o f blood flow in 2 msec. Its normal sensory stimulus would be
D. all of the above most likely to be:
A. tension exerted on a tendon
Which of the following is false? B. pain
A. Right parietal lesions can lead to left-sided C. temperature change
neglect D. light touch
B. Two-point tactile thresholds are smaller for the
back than for the finger T F Spinal motoneurons and inhibitory
C. Pressure on a peripheral nerve can cause intemeurons both receive afferent input from spinal
paresthesia sensory nerve fibers and from descending tracts from
D. Position sense is mediated by the dorsal the brain.
(posterior) column system
For the following 2 questions, match the type of
A painful or very hot stimulus to the hand activates lesion with a physical finding below.
A. unmyelinated afferent C fibers in peripheral Lower motor neuron lesion
nerve _U pper motor neuron lesion
B. the flexor withdrawal reflex
C. the reticular formation A. Delayed conduction in sensory nerve fibers
D. all o f the above B. Ankle clonus
C. Flaccid paralysis
Which of the following is true o f pain sensation? It: D. Left-sided neglect
A. may be elicited by stretching the ureter
B. involves the more slowly conducting nerve For the following 2 questions, match the reflex
fibers response with a characteristic below.:
C. may be elicited by tissue damage Inverse stretch reflex
D. all of the above Flexor withdrawal reflex
In deafness due to intense, prolonged noise: Placing atropine in the left eye would have what
A. bone conduction from a tuning fork would be effect on the pupillary light reflex elicited by shining
superior to air conduction a light in the left eye?
B. an operation on the middle-ear ossicles could A. Pupillaiy constriction in the right eye
correct the deafness B. Pupillary constriction in the left eye
C. the primary cause of the hearing loss is damage C. Absence of the reflex response in both eyes
to the cortex D. Absence of the reflex response in the left eye
D. there is damage to the organ o f Corti
Which of die following is true? T F A patient whose hearing is normal but who
A. in accommodation die suspensory ligaments cannot understand speech may have a lesion in
allow the lens to become less convex Wernicke's area on the left hemisphere.
B. pupillary dilation is blocked by atropine in the
eye T F In focal motor epilepsy, abnormal neural
C. the greatest ability to adapt to dim light is in the activity occurs in die brainstem reticular formation
fovea centralis and die entire cerebral cortex.
D. die blind spot in the visual field corresponds to
the optic nerve head in the retina T F Alexia is due to a lesion o f the arcuate
fasciculus.
Accommodation for near vision involves which of
the following: Which of the following statements is FALSE?
A. the pupils dilate A. Brief lapses o f consciousness in a child with a
B. sympathetic nerve fibers to the eye are activated spike-and-wave EEG at 3 per second is characteristic
C. the front surface o f the lens becomes rounder of petit mal epilepsy.
D. all of the above B.. Seizure activity limited to the hand and arm with
no alterations o f consciousness is characteristic of
Acetylcholine secreted by cranial nerve III causes: psychomotor epilepsy.
A. pupillary dilation only C. Seizure activity moving from one part of die
B. accommodation only body to another is characteristic of Jacksonian march
C. accommodation and pupillary constriction epilepsy.
D. pupillary constriction only D. Loss of consciousness with tonic-clonic seizures
is characteristic of grand mal epilepsy.
Shining a flashlight in the right eye normally causes
the pupil to: Closing the eyes often leads to which o f die
A. constrict in the right eye only following EEG patterns?
B. constrict in the left eye only A. beta waves
C. dilate in the right eye only B. alpha waves
D. constrict in both eyes C. spike-and-wave rhythm
D. blocking o f alpha waves
Acetylcholine secreted by cranial nerve III causes:
A. pupillary dilation only Consciousness may readily be lost with damage to
B. accommodation only the:
C. accommodation and pupillary constriction A. motor cortex
D. pupillary constriction only B. dorsal columns
C. cerebellum
Which of the following is true? D. brainstem reticular formation
A. in accommodation the suspensory ligaments
allow die lens to become less convex Motor seizures limited to the left hand with no loss of
B. pupillary dilation is blocked by atropine in the consciousness are often due to lesions of which
eye cortical area?
C. the greatest ability to adapt to dim light is in the A. Wernicke's area on the left hemisphere
fovea centralis B. Striate cortex on the right hemisphere
D. die blind spot in the visual field corresponds to C. Motor cortex on the right hemisphere
the optic nerve head in the retina D. Broca's area on the left hemisphere
T F Petit mal epilepsy is characterized by motor If a patient can understand speech, and has no oral
seizures and a spike-and-wave EEG pattern at 10 per paralysis, but speaks with difficulty and not fluently,
second. he may have a lesion in:
A. Wernicke's area on the left hemisphere
B. Striate cortex on the right hemisphere
C. Motor cortex on the right hemisphere
D. Broca's area on the left hemisphere
Grateful acknowledgment is made to the following sources for permission to reprint material copyrighted or
controlled by them:
Figure 1-5: "Lateral views of the left cerebral hemisphere," by Murray L. Barr, reprinted from The Human Nervous System:
An Anatomic Viewpoint, 1979, Lippincott-Raven Publishers.
The following figures by John C. Eccles are reprinted from The Understanding ofthe Brain, 1973, McGraw-Hill:
Figure 2-1, "Peripheral synaptic transmission," and figure 4-6, "Experiment demonstrating concentration of
acetylcholine receptors near motor nerve terminals."
Figure 6-7: "Displacement envelopes..." by George Von Bekesy, reprinted from Experiments in Hearing, 1960,
McGraw-Hill.
Figure 6-15: "Far-field recording of auditory brain stem responses latencies measured in human subjects," reprinted from
"Detection and Localization of Occult Lesions with Brainstem Auditory Responses," by J.J. Stockard, J.E. Stockard, and
F.W. Sharbrough, by permission from Mayo Clinic Proceedings, Vol. 52,1977.
Figure 6-17: "Physiologic nystagmus produced by rotating the head..." by Andrew J. Gay, Nancy M. Neman, John L.
Keltner, reprinted from Eye Movement Disorders, 1974, C.V. Mosby Company. Reprinted by permission of the authors.
Figure 7-10: "The eye, including central nervous system control of eye movements," by Gerald Westheimer,
reprinted from Medical Physiology, Vol. 2,1968, C.V. Mosby Company.
The following figures by Murray L. Bair are reprinted from The Human Nervous System: An Anatomic Viewpoint, 1979,
Lippincott-Raven Publishers:
Figure 10-1: "The sympathetic nervous system," and Figure 10-2: "The parasympathetic nervous system..."
Figure 11-2: "The International 10-20 electrode system," by H.H. Jasper, reprinted from Electroencephabgraphy and Clinical
Neuropltysiology, Vol. 10,1958, with kind permission from Elsevier Science Ltd., Bay 15K, Shannon Industrial Estate, Co.
dare, Ireland.
Figure 11-8: "Broca's area..." by N. Geschwind, reprinted from "Specializations of the Human Brain," Scientific American,
Vol. 241, September 1979.
Figure 11-9: "Lateral specialization in the surgically separated hemispheres," by R.W. Sperry, reprinted from The
Neurosciences: Third Study Program, edited by F.O. Schmitt and F.G. Worden, 1974, by permission of MTT Press.
Contents of this book include revisions of Human Brain Function: An Introduction to Clinical Neurophysiology by Robert
Lavine, 19%, ISBN 0536598223. This book was previously published by: Pearson Education, Inc.
Mental disorders are com mon. You may have a friend, • How genes and the environm ent affect the brain
colleague, or relative with a mental disorder, or perhaps
you have experienced one yourself at some point. Such • The basic structure o f the brain
Tire Working Brain
disorders include d ep re ssio n , a n x ie ty d is o rd e rs , b ip o la r
• How different parts of the brain com m unicate and work
d iso rd e r, a tte n tio n d e fic it h y p e ra c tiv ity d is o rd e r (ADHD),
with each other
and many others.
Brain Basics in Real Life • How changes in the brain can lead to mental disorders,
Some people w ho develop a mental illness m ay recover such as depression.
com pletely; others may have repeated episodes of illness
with relatively stable periods in between. Still others live with
symptoms of mental illness every day. They can be moderate,
Brain Research or serious and cause severe disability.
N EX T P A G E > 1
\Z * * 7
I V I I I I I National Institute of Mental Health
Brain Basics
The Growing Brain
In s id e t h e B r a in : N e u r o n s & N eu ra l C ir c u its Each neuron is enclosed by a cell membrane, which separates
the inside contents o f the cell from its surrounding environm ent
Neurons are the basic w orking unit o f the brain and nervous and controls w hat enters and leaves the cell, and responds to
The Growing Brain system. These cells are highly specialized for the function of signals from the environm ent; this all helps the cell m aintain its
conducting messages. balance with the environment.
A neuron has three basic parts: Synapses are tiny gaps between neurons, where
The Working Brain m essages move from one neuron to another as chemical
• Cell body w hich includes the nucleus, cytoplasm, and cell or electrical signals.
organelles. The nucleus contains D N A a n d information that
the cell needs for growth, metabolism, and repair. Cytoplasm The brain begins as a small group o f cells in the outer layer
Brain Basics in Real Life is the substance that fills a cell, including all the chem icals of a developing embryo. As the cells grow and differentiate,
and parts needed for the cell to w ork properly including neurons travel from a central “birthplace” to their final
small structures called cell organelles. destination. Chemical signals from other cells guide neurons
in form ing various brain structures. Neighboring neurons make
• Dendrites branch off from the cell body and act as a neuron’s connections with each other and with distant nerve cells (via
Brain Research point o f contact for receiving chemical and electrical signals axons) to form brain circuits. These circuits control specific
called impulses from neighboring neurons. body functions such as sleep and speech. The brain continues
m aturing well into a person’s early 20s. Knowing how the
• Axon which sends impulses and extends from cell bodies to
brain is w ired and how the normal brain’s structure develops
meet and deliver impulses to another nerve cell. Axons can
Glossary range in length from a fraction of an inch to several feet.
and m atures helps scientists understand w hat goes wrong in
mental illnesses.
More Information
< P R E V IO U S P A G E N EX T P A G E > 2
\Z * * 7
I V I I I I I National Institute of Mental Health
Brain Basics
Introduction
The Growing Brain
T h e C h a n g in g B r a in — E f f e c t s o f G e n e s a n d E pigenetics is the study o f how e nvironm ental fa cto rs can
t h e E n v ir o n m e n t affect how a given gene operates. But unlike gene m utations,
epigenetic changes do not change the code fo r a gene. Rather,
The Growing Brain There are m any different types of ceils in the body. W e say they effect w hen a gene turns on or off to produce a specific
that cells differentiate as the embryo develops, becoming more protein. S cientists believe epig e ne tics play a m ajor role in
specialized fo r specific functions. Skin cells protect, m uscle m ental disorders and the effects of m edications. Some, but
cells contract, and neurons, the most highly specialized cells not all m utations and epigenetic changes can be passed on
Tire Working Brain of all, conduct messages. to future generations.
Every cell in our bodies contains a complete set o f DNA. DNA, Further understanding o f genes and epigenetics may one
the “recipe of life,” contains all the information inherited from day lead to gen e tic testing fo r people at risk fo r m ental
Brain Basics in Real Life our parents that helps to define w ho we are, such as our looks disorders. This could greatly help in early detection, more
and certain abilities, such as a good singing voice. A gene is tailored treatments, and possibly prevention of such illnesses.
a segm ent o f DNA th a t contains codes to m ake proteins and
other important body chemicals. DNA also includes information
to control which genes are expressed and when, in all the cells
B rain R esearch of the body.
< P R E V IO U S P A G E N EX T P A G E > 3
\Z * * 7
I V I I I I I National Institute of Mental Health
Brain Basics
Introduction
The Working Brain
N e u r o tr a n s m itte r s
Everything we do relies on neurons com m unicating with synapse fo r the receiving neuron to bind onto, leading
Tire Growing Brain one another. Electrical impulses and chemical signals carrying to more normal mood functioning.
messages across different parts o f the brain and between the
< P R E V IO U S P A G E N EX T P A G E > 8
Mil National Institute of Mental Health
Brain Basics
Links
Introduction
N atio n a l In s titu te o f M ental H ealth (NIMH)
Publications, science news, and funding inform ation related to Federally-supported research on mental disorders
M e d lin e P lu s
Hie Working Brain C onsum er health information provided by the National Library of Medicine
Brain Research
Glossary
Links
More Information
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Brain Basics
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Introduction
N a tio n a l I n s titu te o f M e n ta l H ea lth (NIMH)
Science, Writing, Press & Dissem ination Branch
. . 6001 Executive Boulevard
H ie Growing Brain Room 8 184, m s c 9663
Bethesda, MD 20892-9663
Brain Research
Glossary
Links
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Postsynaptic receptors either regulate sodium (Na+) and other ionic channels in the
nerve membrane (for ionotropic receptors) or affect metabolic processes from DNA to
RNA and proteins in the postsynaptic nerve cell (for metabotropic receptors).
ACh. Acetylcholine, or ACh, acts in both the CNS and in the autonomic nervous
system, where it is released in the autonomic ganglia and also by parasympathetic fibers
onto end organs. For example, it can slow the heart rate by acting on the SA node of the
heart. After release into the synapse, it is broken up by the enzyme acetylcholinesterase
and is also subject to reuptake into the presynaptic terminal for resynthesis.
Serotonin. Serotonin. or 5-HT, acts on the brain to affect mood, alertness, and the
sleep cycle. The raphe nucleus in the brainstem sends neurons that branch out to
secrete serotonin at several brain and spinal cord regions. SSRIs or Specific Serotonin
Reuptake Inhibitors are drugs that are thought to act as antidepressants by regulating
serotonin at synapses.
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