Neuroscience ABCs

Neuroscience
ABCs
Human Brain
Physiology Guide
Understand
•Nerve impulses
•Touch & Pain
•Hearing & Vision
•Movement
•Speech & EEG
Robert Lavine, Ph.D. Robert Lavine, Ph.D.

NEUROSCIENCE ABCs
Copyright (c) 2016 by Robert Lavine
All rights reserved, including the rights to publish this book or portions thereof in any form whatsoever.
Neuro Cognitive Publications
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)
The book can be read alone or along

with lectures, longer texts, and exam
reviews, with an emphasis on physiology
that can be accompanied by books on
the anatomy of the nervous system. The
review questions can be skipped. I’ve
provided examples from clinics,
laboratory research, and everyday life.
The goal is to be more readable than
textbooks that emphasize dry
definitions.
The goal is also to provide a foundation

for topics such as pharmacology,
neurology, and current neuroscience
research. A necessary disclaimer is that
the book is not intended to provide
medical advice.
Since neuroscience is complex, you may

be tempted to remember bits and pieces
without mastering the essential
concepts. If you are taking or reviewing
a class in the subject, the best way to
learn is to do something actively with
the subject—that’s why I have included
questions at the end of each chapter and
review tests at the end. But otherwise,
you can skip these practice questions
and go on to the next chapter, or any
other chapter you wish.
Research findings and clinical exam

outlines are inserted in text boxes.
Important brain regions, brain imaging,
research methods, and major
Contents

c
How the nervous system is designed to 1
communicate information and control
activity
2. The Nerve Impulse How nerve cells become excited and 25

conduct impulses
3. Sensory Reception How receptors convert energy from the 47

environment into nerve impulses
4. Synaptic Transmission How signals pass from neuron to neuron 59

and from neuron to muscle fiber
5. Somatic Sensation How the nervous system converts touch, 85

pressure, joint position, temperature, and
painful stimuli into sensations
6. Auditory and How neural mechanisms function in 105

Vestibular Function hearing and balance
7. Vision How the eye works and how the visual 129
system converts light into sensed visual
images
8. Chemical Senses How the nervous system responds to 157

chemicals to produce sensations o f taste
and smell
9. Posture and Movement How motor systems control the skeletal 165
muscles
10. Autonomic Function How sympathetic and parasympathetic 197

activity help control the internal
environment
viii Contents
11. Cerebral Physiology and How cerebral activity relates to 207

Higher Functions wakefulness and sleep, language, mem
ory, and other complex processes
Glossary 233
References 242
Quiz. Yourself 244
Index 251
How the nervous system is designed to communicate

information and control activity_________________
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
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.
put). There are then four types of peripheral

spinal nerve fiber:
Figure 1-3. (A) Diagram of nervous system as seen
from dorsal surface (not to scale), divided into pe
1. Somatic afferent, carrying sensory informa ripheral nervous system (PNS) and central nervous
tion from skin, skeletal muscles, tendons, system (CNS), consisting of spinal cord and brain.
and joints Spinal segments are C = cervical, T = thoracic,
2. Somatic efferent, carrying motor-command L = lumbar, S = sacral, and Co = coccygeal.
Only somatic afferent and efferent fibers are shown.
signals to skeletal muscles
MN. motoneuron (alpha motoneuron controlling
3. Visceral afferent, carrying sensory informa muscle fibers in arm). Ascending fibers shown cany
tion from the visceral organs sensory information from spinal cord to somatic sen
Receptors PNS CNS
& Effectors
Dorsal column
D O R S At
Dorsal root
Dorsal horn (Gray matter)
White matter
Ventral horn (Gray 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.
Table 1-1. Cranial Nerves and Their Major Functions

Nerve Afferent Fiber Functions Efferent Fiber Functions
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
VIII Auditory Hearing

(vestibular) Maintenance of equilibrium
IX Glossopharyngeal Taste from posterior one-third Salivation
of tongue Swallowing
Blood pressure information from
cervical baroreceptors
Chemical information from cervical
chemoreceptors
X Vagus Taste from part of tongue and pharynx Salivation
Viscerai sensation from pharynx, thorax, Swallowing
and abdomen Slowing of heart rate
Blood pressure information from thoracic Secretion and patency of bronchioles
baroreceptors Production of surfactant in lungs
Chemical information from thoracic Secretions and movements of
chemoreceptors abdominal digestive organs
XI Accessory Swallowing
Turning head
Elevating shoulder
XII Hypoglossal Tongue movement
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
at the midline into a left and right half and

have surfaces labeled dorsal (toward the back)
and ventral (toward the front).
Within the CNS are white and gray areas.
The white areas contain bundles of nerve axons
(also called nerve fibers); their color is imparted
by the myelin sheath around the nerve fibers.
There are several anatomical terms for specific
bundles of nerve fibers within the CNS, such as
tract, column, fasciculus, lemniscus, and radia
tion. The gray areas contain the cell bodies of
neurons, their dendrites, and associated
synapses.
In cross-section, the gray areas in the spinal
cord form a butterfly-shaped area in the center
of the cord, surrounded by a ring of white
matter (figure 1-3B). The white matter con
tains bundles of axons passing up and down don). Abbreviations: Cblm. = cerebellum, Thai. =
through the cord connecting areas of the cord thalamus, Hy. = hypothalamus, Corp. Cal. = cor
pus callosum, M = motor cortex, S. = somatic
with each other and with the brain, while the
sensory cortex, V. = visual cortex. Temporal lobe
central gray area contains numerous synaptic not shown. The ventricles contain cerebrospinal fluid.
regions. Thus, a sensory signal may enter the Dashed line shows juncture of spinal cord and brain.
cord through a dorsal-root nerve axon that (1)
travels up to the brain in an ascending column information from the eyes and ears as well as
or tract or (2) enters the gray area to synapse on other sources. Within the core of the brain
other nerve cells. Within the brain, gray areas stem a network of nerve cells called the
consist of cell bodies and synapses organized reticular formation is essential in regulating
into nuclei (clusters) and layers (in the surface wakefulness and sleep, respiration, and car
of the cerebrum and cerebellum). diovascular function.
The brain is the highest level of the CNS. It 2. The cerebellum is a globular structure con
is the organ ultimately responsible for move nected to the dorsal surface of the brainstem
ment, perception, and thought. The brain has by bundles of nerve fibers. Its surface is
four major divisions (figure 1-4): covered with folds, so that it partly resem
bles a ball of twine. A major function of this
1. The brainstem partially resembles an exten organ is to correct errors so that movement
sion of the spinal cord upward into the head. proceeds in a smooth and controlled man
It is composed of the medulla, pons, and ner— so that the hand, for example, can
midbrain. Like the spinal cord, the brain reach for and pick up a cup without spilling
stem contains white bundles of nerve fibers the contents.
(called lemnisci, tracts, and so on) carrying 3. The thalamus and hypothalamus, together
signals both up and down, gray nuclei called the diencephalon, are gray areas con
containing cell bodies and synapses, and the taining nuclei (clusters of cells) located
roots of peripheral nerves— the cranial above the brainstem and near the center of
nerves— arranged in a more complex fash the brain, surrounding the walls of the third
ion than the spinal nerves and carrying ventricle. The thalamus is a major relay
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.
central
sulcus
parietooccipital
fissure
preoccipital
anterior ram us notch
lateral ascending ramus

fissure
posterior ram us
premotor
area motor area
frontal eye som esthetic

field 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.
This fMRI image show a side view of the inside

of the brain during a numerical task. The
occipital lobe in back is to the left, and the
frontal lobe is to the right. The highlighted
region in red shows an activated area in the
parietal lobe.
Adapted from: Cantlon JF, Brannon EM, Carter EJ,

Pelphrey KA (2006) Functional Imaging of
Numerical Processing in Adults and 4-y-Old
Children. PLoS Biol 4(5): e125.
doi:10.1371/journal.pbio.0040125
Copyright: © 2006 Cantlon et al. Creative
Commons Attribution License.
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.
Chemical Transmitter Release Velocity
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
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
Giant Axons from the

Giant Squid
A Giant Squid
(ClipArtist.info, Public Domain)
The giant squid can reach 40 feet long

with a giant axon that can reach 1 mm in
diameter. It allowed British researchers
Hodgkin and Huxley to squeeze out the
cytoplasm, insert a micropipette, and
change the salt solution in the cell. They
could then insert a thin wire to record the
voltage across the membrane.
The membrane has two layers of lipid or

phospholipid molecules (a bilayer).Their
lipid chains face each other and their
polar ends (the small circles in the
diagram) face the ionic solutions outside
and inside the nerve cell. Ion channels
are found crossing this membrane matrix,
allowing the axon to transmit signals.
More about these channels later.
Outside
Inside
Adapted from Depolarizing Pre-Pulse,

Wikipedia, (Creative Commons CC BY S-A 3.0).
and vibrations set up by the predator at a

distance. Similar functions are now carried out
by the human sensory processes— body sensa
tion, vision, hearing, and others. Taking
somatic sensation (body sensation, including
touch and pressure) as an example, we can look
at the sequence of activities that allow us to
sense the outside world so that we can deal with
it.
Figure 1-8. Information arriving at synapses from
two axons is combined by cell body, an example of
Transduction synaptic integration.
Pressure on the skin (after someone hands you a
book, for example) is a common type of sensory reached. At higher levels, some neurons re
stimulus. Specialized receptors in the skin spond as if they detect certain complex features
respond to the pressure with a graded receptor of the stimulus, such as, the angle of a line or
potential (an electrical signal). The transfor the change in pitch of a tone. This feature
mation of one form of energy (mechanical) into detection is one important result of the ability
another (electrical) is called transduction. of synapses to integrate several incoming
Transduction is the fundamental step that signals.
permits the nervous system to react to events in
the outside world. Specificity
Conduction Some receptors in the somatic sensory system

are specialized to process pressure; receptors in
Action potentials are generated at the periph the ear, sound waves; and the rod receptors in
eral (receptor) end of the sensory axon and are the retina, light. In all three cases, the out
then conducted at a characteristic velocity to come is a graded receptor potential. Each
the spinal cord or brain. Because of unpredict receptor is most sensitive to a specific type of
able fluctuations of neuron activity, some infor external energy stimulus (figure 1-9). Each
mation in each axon is lost. This is compen receptor also is connected to a specific sensory
sated for by a number of parallel axons trans system of nerve cells in the brain and spinal
mitting the same message so that information cord and transmits neural signals through that
lost in one axon is transmitted in the others. system. The different sensations produced
(pressure, sound, light) are due to such specific
Integration connections.
Chemical transmitters, released from several
axons, affect the next neuron in the sensory
Motor Processes
system (figure 1-8). This allows the neuron to
integrate (put together) signals from several Skeletal muscle contractions either hold the
sources. Such integration is a function of the body in a relatively fixed posture (sitting,
synapse. standing, waiting for a tennis ball) or cause it to
Conduction along axons and integration at move (writing, lifting, swinging at the tennis
synapses occur at successive levels of the sen ball). Both posture and movement are motor
sory pathway until the cerebral cortex is processes, controlled by somatic efferent nerve
13 Struetore and Function
EXTERNAL RECEPTOR SENSORY

ENERGY SYSTEM
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
1_3)‘ The electrical potential change within muscle

The pathway from the muscle-stretch recep then leads to muscle contraction (a mechanical
tor to the spinal motoneuron to the muscle change). This change from one form of energy
effector is an example of a spinal reflex, which is to another is a form of transduction, which is
an automatic motor response utilizing a circuit the reverse of what occurs in receptors that
built into the spinal cord. The effect of this transduce mechanical into electrical energy.
reflex would be to support the weight of the
book, opposing the force of gravity. The path
Autonomic Nervous System
way from the brain motor centers to the spinal
motoneuron is an example of supraspinal motor Posture and movement are governed by con
control. It governs the strength of the spinal tractions of the skeletal muscles. On the other
reflex and any voluntary, purposeful move hand, the internal state of the body, including
ment, such as raising the book to eye level. heart rate, blood pressure, pupil size, and the
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,
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 willprobablyfind that while most adults

can repeat List 1, containing 4 digits, and List
2, containing 6 digits, they cannot do so for
List 3, containing 11 digits. This is a test for
short-term memory and demonstrates that its
capacity is limited. Long-term memory, which
lasts hours, days, or years, has no clear limits in
its capacity and seems to be organized on dif
ferent principles. Moving material to be
memorized from short-term memory to long
term memory is a most important step. Inter
estingly, this step seems to be associated with
the function of a particular area of the brain—
the hippocampus, a cellular structure within the
temporal lobe.
Memory is closely associated with learning in
human behavior. Both can be demonstrated at
a simple level. If you snap your finger once,
twice, or three times, a child may be startled.
After 20 times you will probably be ignored.
The change in motor response, from startle to
boredom, is evidence that the repeated sounds
have been stored in memory, and the listener
has learned that they are harmless. In other
words, there has been a modification of behav
ior due to experience. Learning and remember
ing lists o f numbers, or even learning
neurophysiology, are also modifications of be-
Readers, congratulations!
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
Review Questions
1. Diagram a neuron, labeling the cell body, dendrites, axon, and synapses.
2. Describe the role of receptors and effectors.
3. List and define the major components of the peripheral nervous system and the
central nervous system.
4. Compare the white and gray areas of the CNS.

5. List in outline form the major structures within the brain.
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.
8. Define transduction in a sensory receptor.

9. Give an example o f how a spinal motoneuron might integrate information from

several sources.
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
How nerve cells become excited and conduct impulses
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 2-2. An unmyelinated axon (left) and class

ical model of its cell membrane (right) shown m
schematic diagram. Figure 2-3. The sodium-potassium pump in the
nerve cell membrane.
metabolic energy to pump out any sodium ion
that diffuses in, and, to a lesser extent, to pump
back in the potassium ion that diffuses out
(figure 2 -3 ).
The excess sodium ion outside the nerve cell
is also kept there because the sodium permea
bility of the membrane is very small in the
resting state. The large anions (amino acids,
protein, organic phosphate, sulfate) remain
inside because they cannot permeate the mem
brane at all.
In the axon, 20% or more of all metabolic
energy is used by the pump for active transport
of sodium ions. The immediate source of this
energy is adenosine triphosphate (ATP). One
If the membrane were permeable only to

potassium ions, potassium ions would diffuse
through channels in the membrane from inside
to outside, down their concentration gradient.
They would leave behind the large anions,
which cannot permeate the membrane. Since
originally both internal and external solutions
are electrically neutral (each has the same
concentration of anions and cations), the slight
excess of potassium ions that develops outside
and the slight excess of anions that develops
inside cause a potential difference across the
membrane. At equilibrium, this potential dif
ference exactly opposes further diffusion of
Figure 2-4. One model of the coupled sodium-potas- potassium ions outward (positive repels posi
shan pump; the pump utilizes energy provided by the tive). The concentration difference of potas
breakdown of ATP.
sium ions is balanced by an opposing potential
model for the pump is a carrier that transports difference. The potential difference can then
sodium ions from inside to outside the cell and be calculated from the concentration of potas
then combines with potassium ions and moves sium ions on the inside, or K +j( and the
concentration of potassium ions on the outside,
from outside to inside (figure 2 -4 ). ATPase
enzymes, which are activated by sodium and or K +<„ according to this formula:
potassium ions, have been isolated in cell
membranes and may be involved in this trans K+
port process. Vjnside — ^outside — V — —61 fog ^ +
Now that we know how the unequal concen
trations of ions are maintained across the 150
membrane, we can describe them in more = -6 1 log —
detail and see how they produce a resting
potential. = -6 1 log 30
= -6 1 x 1.47
The Resting Potential = -9 0 mV1 (inside
negative to outside)
Dependence on K + Concentration Difference
This potential difference is called the Nemst
Inside the nerve cell in the neuronal cyto equilibrium potential for potassium ions, or Eg.
plasm, the potassium ion is highly concentrated Thus, normally:
at 150 mM (all concentrations are approximate
only). These potassium ions are associated with 150
equal concentration of large anions symbolized Ek = -6 1 log ~j~ = -6 1 log 1.47
by A". Outside the nerve cell, in the extracel
lular fluid, potassium ion concentration is low = -9 0 mV.
(5 mM). Sodium chloride concentration is low
inside and high outside the membrane (figure 90
‘- 9 0 mV = volts or slightly less than 1/10 volt.
2-5 ). 1000
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 .
Table 2-1. Common Electrical Symbols

Symbol Term Common units* Meaning
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
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
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
Figure 2-8. Membrane potential measured in a large The Action Potential

nerve axon. Graphs show potential versus time as it
would appear on an oscilloscope screen. (A) M i-
croelectrode outside axon; the recorded potential dif The action potential, as shown in figure 2-10,
ference = 0; (B) Microelectrode inside axon; the begins when a depolarization goes beyond a
recorded potential difference equals membrane poten certain threshold value— usually about 15 mV
tial of -7 0 mV, the resting potential. (C) Slight from the resting potential (for example, from
(subtkreshold) depolarization and hyperpolarization of
- 7 0 mV to -5 5 mV). A rapid depolarization
the membrane from the resting level
follows and continues until membrane polarity
axon the depolarization disappears completely. is actually reversed; inside becomes briefly
However, signals in the human nervous system positive to outside, overshooting the zero mV
must be transmitted for distances up to almost line by about 30 mV to 40 mV. This is followed
2 m, which is done by means of the action by a rapid repolarization back to resting leveb.
potential. The whole process is called an action potential;
due to its appearance, it may be also called a
spike.
Figure 2-9. A subthreshold depolarization, showing
its decay with distance. The electrodes are about 1 The action potential is conducted along the
mm apart, and all traces start at the time the axon and does not decay with distance. It does
stimulus (S) is applied (arrow). not change in either height or time-course
Figure 2-10. An action potential, originating with a Na+( 15 mM

stimulus applied near the left end of the axon. The ENa = -6 1 log — — = -6 1 lo g -----------
stimulus is stronger, but otherwise the experimental K Na o S 150 mM
preparation is the same as in figure 2-9. Note the
absence of decay with distance. = + 61 mV (inside positive)
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
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.
10-12 mole/cm2 of membrane). Although

sufficient to reverse the membrane poten
tial, this ionic movement is not sufficient to
significantly change the concentration on
both sides. The excess Na+ ions are gradu
ally pumped out.
4. Near the action potential peak, sodium
channels close down again (sodium conduc
tance declines), while potassium channels
open more than in the resting state (potas
sium conductance rises). Both factors cause
repolarization to the resting level and also a
period of slight hyperpolarization (the hy
perpolarization occurs because the increased 0 1 2 3 4
potassium conductance brings the mem msec
brane closer to Er (-9 0 mV) than during

rest). Figure 2-12. Conductance changes for sodium (g\J
and potassium (g*) drawn on the same time scale as
After the onset of the spike there is an the action potential. In the action potential trace, a =
absolute refractory period in which a second spike depolarization, b = peak of action potential, c = re-
polarization, and d = hyperpolarization. Source:
cannot occur, no matter how strong the Adapted from A. L Hodgkin. The Conduction of
stimulus. This is followed by a relative refractory the Nervous Impulse. Springfield, III: Charles C
period, in which a second spike can occur but Thomas, 1964.
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.
10-12 mole/cm2 of membrane). Although

sufficient to reverse the membrane poten
tial, this ionic movement is not sufficient to
significantly change the concentration on
both sides. The excess Na+ ions are gradu
ally pumped out.
4. Near the action potential peak, sodium
channels close down again (sodium conduc
tance declines), while potassium channels
open more than in the resting state (potas
sium conductance rises). Both factors cause
repolarization to the resting level and also a
period of slight hyperpolarization (the hy
perpolarization occurs because the increased 0 1 2 3 4
potassium conductance brings the mem msec
brane closer to Er (-9 0 mV) than during

rest). Figure 2-12. Conductance changes for sodium (g\J
and potassium (g*) drawn on the same time scale as
After the onset of the spike there is an the action potential. In the action potential trace, a =
absolute refractory period in which a second spike depolarization, b = peak of action potential, c = re-
polarization, and d = hyperpolarization. Source:
cannot occur, no matter how strong the Adapted from A. L Hodgkin. The Conduction of
stimulus. This is followed by a relative refractory the Nervous Impulse. Springfield, III: Charles C
period, in which a second spike can occur but Thomas, 1964.
few milliseconds have passed (the sodium con

ductance mechanism becomes inactivated).
The channels through which ions move
across the membrane have recently been the
focus of much attention [14,21]. These chan
nels are thought to consist of proteins extend
ing from one side of the membrane to the
other. Treating the membrane with certain
drugs and enzymes has shown that there are
separate channels for Na+ and for K+ . Let us Ap
consider the N a+ channel. During the resting
state, the channel is effectively blocked by a
closed, protein gate structure. Depolarization Figure 2-13. Conduction of the action potential by
apparently causes a rearrangement within the local circuits in an unmyelinated axon. (A) The
action potential (AP) has depolarized a section of
membrane of charged or dipolar particles as
axon, while the section to its right is still in a resting
sociated with these gates (for example, the state (inside negative). The arrows show local cir
positively charged groups of protein molecules cuits of ionic current. (B) An instant later, the sec
might be driven away from the channel, open tion to the right has been depolarized and is now
ing it to the passage of the positive sodium generating an action potential, while the previously
active section to its left has become repolarized.
ions). This rearrangement of charge within the
membrane produces a minute but measurable in voltage difference across the membrane
current called the gating current and is as (figure 2-13). In unmyelinated axons this active
sociated with opening the gate to sodium. Thus region acts to depolarize the next adjacent
the tiny gating current is followed by a large segment of membrane by means of local cir
ionic current due to the sodium influx across cuits. Local circuits are simply ionic currents in
the membrane. extracellular fluid, axoplasnm, and membrane.
The different ion channels can be selectively If they are blocked— by suspending a short
blocked by certain drugs. For example, let- length of axon in air— conduction will cease.
rodotoxin, a poison obtained from fish, blocks In myelinated axons (figure 2 -14) local cir
the sodium channels that allow sodium to move cuits can only travel across the membrane at
inward during depolarization, and thereby pre the nodes between the myelinated, low-
vents the generation of an action potential. conductance intemodes. This makes for faster
Similarly, tetraethylammonium blocks the potas conduction as each small segment of membrane
sium channels. In addition, an inward move does not have to undergo the process of
ment of calcium through calcium channels
contributes in varying amounts to the action Figure 2-14- Saltatory conduction in a myelinated
potential in nerve and muscle cells, and certain axon.
drugs have been found to block these channels.
Conduction o f the Action Potential
At some segment of axon membrane, depolari

zation to threshold voltage can occur with
adequate stimulation. This segment generates
an action potential, marked by a brief reversal
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.
Figure 2-15. Conduction velocity of myelinated ax

ons in meters per second (M.P.S.) plotted against Recording o f the Action Potential
axon diameter. From Gasser, H.S. Ohio Journal of
Science 41:145-159, 1941. Reprinted with permis We have described how action potentials are
sion. recorded with a microelectrode inside the nerve
Conduction
velocity
(m/sec)
Diameter (microns)
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
Figure 2-16. Types of action potential recordings.

(A) Intracellular (electrode tip inside nerve cell).
(B) Extracellular (electrode tip outside nerve cell).
(C) Compound action potential (electrode placed
outside compound nerve). In B and C, nerve under
neath reference electrode (rectangular plate) has been
crushed so that it does not also generate an action
potential, which would make recording more difficult
to interpret.
yer
Review Giant Axons to

Sodium Channels
A Giant Squid
(ClipArtist.info, Public Domain)
The giant squid can reach 40 feet long

with a giant axon that can reach 1 mm in
diameter. It allowed British researchers
Hodgkin and Huxley to squeeze out the
cytoplasm, insert a micropipette, and
change the salt solution in the cell. They
could then insert a thin wire to record the
voltage across the membrane.
The membrane has two layers of lipid or

phospholipid molecules (a bilayer).Their
lipid chains face each other and their
polar ends (the small circles in the
diagram) face the ionic solutions outside
and inside the nerve cell. Ion channels
are found crossing this membrane matrix,
allowing the axon to transmit signals.
More about these channels later.
Outside
Inside
Adapted from Depolarizing Pre-Pulse,

Wikipedia, (Creative Commons CC BY S-A 3.0).
VOLTAGE-GATED SODIUM
CHANNEL STRUCTURE
No, the sodium, channels are not just little

doorways. They are proteins, composed of
amino acids, embedded in the nerve
membrane, with an opening that serves as
a selectivity filter, only allowing sodium
(Na+) ions through. The voltage-gated
channels have a gate portion that can open
and close, controlled by a voltage sensor
that responds to the membrane potential.
The side-view above shows part of a

voltage-gated sodium channel (Namadurai,
S. and others, 2015). This cartoon shows
an ion channel or pore surrounded by
helical-shaped peptides of the channel's
alpha subunits embedded in the
membrane.
The peptides are divided into domains (D)

and subunits (S).The authors think that
sodium channels in human nerve
membrane resemble this structure,
although they derived this image from
studies of bacterial membrane.
Adapted from Namadurai S, Yereddi NR, Cusdin FS,

Huang CL-H, Chirgadze DY, Jackson AP. (2015). A
new look at sodium channel b subunits .Open Biol. 5:
140192.http://dx.doi.org/10.1098/rsob.140192
Review Questions
1. Describe the sodium-potassium pump in the nerve membrane.
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.
Review Questions
equation.
rent, capacitance.
Review Questions
equation.
rent, capacitance.
5. Define conductance of the membrane and relate this to membrane resistance and
ionic permeability.
6. Describe the technique for measuring the membrane potential of a neuron.
7. Define resting potential and state a typical value for this in a neuron.
8. Define depolarization, repolarization, and hyperpolarization.

9. Describe the spatial distribution and time-course of subthreshold changes in

membrane potential
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.
12. Describe the sequence of changes in membrane conductance (or permeability) to

sodium and potassium ions that occurs on excitation of a neuron.
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.
15. Describe die absolute and relative refractory periods.
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?
17. D raw a diagram to illustrate the local current flow that accompanies the passage
of a nerve impulse along an axon.
18. Describe saltatory conduction.
19. In the Department of Physical Medicine, the conduction velocity of peripheral

nerves is measured. A 28-year-old man was found to have a conduction velocity
in the motor-nerve fibers of his arm of 31 m/sec, below the norm of 50 to 65
m/sec. Describe one possible cause of the showing.
20. Describe the relationship between the diameter of a myelinated nerve fiber and
its conduction velocity.
17. D raw a diagram to illustrate the local current flow that accompanies the passage
of a nerve impulse along an axon.
18. Describe saltatory conduction.
19. In the Department of Physical Medicine, the conduction velocity of peripheral

nerves is measured. A 28-year-old man was found to have a conduction velocity
in the motor-nerve fibers of his arm of 31 m/sec, below the norm of 50 to 65
m/sec. Describe one possible cause of the showing.
20. Describe the relationship between the diameter of a myelinated nerve fiber and
its conduction velocity.
21. Describe a compound action potential and how it differs from a single-axon
action potential
22. State the all-or-none law for nerve.
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?
21. Describe a compound action potential and how it differs from a single-axon
action potential
22. State the all-or-none law for nerve.
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
How receptors convert energy from the environment into

nerve impulses
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
nerve ending and decaying rapidly with

Metssrwr's corpusde distance away from the point of stimulation.
This spatial decay is like that described for
subthreshold depolarization of a nerve axon
in chapter 2 (see figure 2 -9 ).
2. The receptor potential is graded in am
plitude, in that a more intense stimulus
produces a greater degree of depolarization,
as shown (figures 3 -2 B and 3-2C ). This
applies to a wide range of stimulus strength,
but at a certain point the degree of depolar
Figure 3—1. Mechanoreceptors m hairless skin, such
as the palm or fingertips. Pacinian corpuscles, Meiss- ization reaches a plateau, and further in
ner's corpuscles, and free nerve endings are shown. creases of stimulus intensity have no
additional effect.
puscle, like that of other mechanoreceptors 3. The receptor potential can be summated: if
studied, has these general characteristics: two weak stimuli are separated by only a
short time interval, their receptor potentials
1. Whether or not it generates an action will sum to produce a larger depolarization.
potential, the receptor potential itself is a 4. The receptor potential has no refractory
local depolarization, largely confined to the period in that, unlike an action potential,
one receptor potential can immediately fol
Figure 3-2. Pacinian corpuscle and its axon. Recep
low another, with no delay.
tor potentials and action potentials are recorded with
one electrode on the axon and the other m the solu 5. The receptor potential appears to be due to
tion surrounding the corpuscle. Th = threshold. increased ionic permeability to sodium ions
Only response to stimulus onset, not offset, is shown. and possibly other ions. Although the mech
Stimulus
intensity
-, + 4 0
0 Membrane
potential
- Th C">V)
- -7 0
anism is not known definitely, the mechan

ical stimulus to the membrane may cause a RAPIDLY ADAPTING
movement of its protein components, open

ing ion channels and allowing increased I ------------------------
SLOWLY ADAPTING
ionic permeability.
Initiation o f Action Potential. Although a very

small stimulus will initiate only a small receptor STIMULUS
potential with no action potential (figure 3 -

2B or 3 -2C ), a slight increase in the stimulus J I__
strength will cause a receptor potential large U _________1_________ L
0 100 200
enough to cross threshold and trigger an action
potential (figure 3-2D ). The trigger point for msec
the action potential— that is, the region with
the highest sensitivity (lowest threshold) for Figure 3-3. Rapidly adapting and slowly adapting re-
starting an action potential— is the first node ceptors are compared. Only action potentials (spikes)
of Ranvier. Once begun at this point, the of the receptors are shown, with an identical
action potential or nerve impulse will be stimulus given to both types of receptor.
conducted along the nerve axon to the central
nervous system. In contrast, there are other receptors that are
What kind of signal does the nerve impulse slowly adapting, which continue firing nerve
provide the central nervous system? As you impulses for the duration of the stimulus. Both
know, the amplitude and shape of the action cases are illustrated in figure 3 -3 .
potential do not change with stimulus inten Thus the pacinian corpuscle is not well
sity. All that can change is whether an action suited to transmit information about a con
potential occurs in the first place and the tinued mechanical stimulus. Instead, it is best
timing and frequency of the nerve impulses as suited to transmit information about a rapidly
they follow one another. The occurrence of an changing or phasic mechanical stimulus, which
action potential provides a signal that the can be produced by a vibratory movement of an
stimulus is strong enough to cross threshold; experimental probe. In everyday life, a vibra
the timing and frequency of the impulses tory stimulus can be produced by a tuning fork
depends on the stimulus and on the characteris placed over the skin, by touching an auto
tics of the receptor, especially its adaptation. mobile engine, or by moving the hand across a
corrugated surface. If the strength of the vibra
Rapid Adaptation. The pacinian corpuscle is a tion is high enough, each vibratory cycle
rapidly adapting receptor. A quick mechanical triggers one or more nerve impulses, which are
stimulus above threshold causes a receptor then transmitted from the pacinian corpuscle
potential leading to an immediate burst of one, to the central nervous system (figure 3 -4 ).
two, or three nerve impulses. But the receptor
potential quickly dies away; it rapidly adapts to Other Rapidly Adapting (Phasic) Mechanorecep-
the continued stimulus. As the receptor poten tors. The fingertips, obviously one of the areas
tial dies away, the nerve impulses also cease. of the body with the most acute sense of touch,
Even if the indentation is maintained, there contain other rapidly adapting mechanorecep-
will be no more impulses during the inden tors in addition to the pacinian corpuscles.
tation. These receptors are called Meissner’s corpuscles
ment. Bending the hair activates the receptor,

but holding it in one position has no effect.
Example 2: The Stretch Receptor
The stretch receptors in lobsters and other
crustaceans are easily studied because their
large cell bodies are outside the central nervous
system and send dendrites to adjacent muscle
fibers. A microelectrode can be inserted into
Figure 3—4. A vibratory stimulus to a fingertip, the cell body in order to record both the
plotted against time (lower right), and the action receptor potential generated in the dendrites
potentials initiated in a pacinian corpuscle in response and the action potentials generated in the axon
to that stimulus (upper right). The timing of the (figure 3 -5 ). This receptor transduces a me
spikes reflects the timing of the stimulus.
chanical change— stretch of the muscle— into
a depolarizing receptor potential. As in the
(figure 3 -1 ). Meissner’s corpuscles are most pacinian corpuscle, the receptor potential is (1)
sensitive to lower rates of vibration (e.g., about a local depolarization confined to the vicinity
30 Hz) while pacinian corpuscles are sensitive of the nerve ending without spreading signifi
to higher rates (e.g., about 300 Hz). Thus, cantly into the axon; (2) graded in amplitude
each type of receptor is tuned to a part of the with increased stimulus strength (as shown in
spectrum of vibration and movement speeds figure 3 -5 ); (3) capable of summating; (4)
found in nature. without a refractory period; and (5) due to
On the back of the arm and on other areas of increased ionic permeability. If the receptor
skin, hair-follicle receptors, with nerve endings potential is above threshold (figure 3-5D ),
coiled at the base of hair follicles, are also action potentials will be generated at the initial
rapidly adapting and most sensitive to move- segment of the axon (near the point at which it
leaves the cell body), just as the pacinian
corpuscle generates action potentials at its first
Figure 3-5. A crustacean stretch receptor stimulated node of Ranvier.
by increasing its length by the amount A L Both
Unlike the pacinian corpuscle, however, the
receptor and action potentials are recorded with a
microelectrode in the cell body. T h = threshold. stretch receptor is slowly adapting. That is,
Compare with figure 3-2. during a maintained stimulus (a constant
Receptor potential
/ ^
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
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
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.
5. Define adaptation as applied to receptors and contrast slowly adapting or tonic

receptors with rapidly adapting or phasic receptors.
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.
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
How signals pass from neuron to neuron and from neuron to

muscle fiber ________________________
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
muscle fiber, so no muscle action potential

occurs.
When an action potential arrives at the
nerve ending, it causes the release of about 300
packets of acetylcholine, causing a full-sized
end-plate potential (EPP, figure 4 -4 ). The end-
plate potential has a time-course similar to that
of the miniature end-plate potential, but its
much larger depolarization (over 30 mV) is
enough to cross threshold and trigger a muscle
Figure 4-4■ End-plate potential (EPP), leading to a action potential.
muscle action potential. If action potential were not What events are behind the end-plate poten
triggered at threshold, the EPP would continue as
shown by dashed line before returning to resting po tial? When the action potential invades the
tential Arrow: arrival of impulse in motor nerve nerve ending, it allows calcium ions to enter
ending. through the nerve membrane (figure 4 -5 ). The
calcium entry (possibly by changing the electri
cal charge on the vesicles) causes the vesicles to
(MEPPs) occur randomly and apparently are fuse with the nerve membrane, open, and
due to the spontaneous release of acetylcholine release their contents, a process called exocy-
from vesicles into the synaptic cleft. A single tosis. The acetylcholine, once released, diffuses
packet of acetylcholine, possible representing across the narrow synaptic cleft.
the contents of one vesicle, causes a depolariza A synaptic delay of about 0.5 msec intervenes
tion of about 0.4 mV. These small depolariza between depolarization of the axon terminal
tions are far below the threshold value for the and the onset of the end-plate potential. This
interval includes the time required for presy
Figure 4-5. Release of chemical transmitter (acetyl naptic release of the acetylcholine, its diffusion
choline) from vesicles in the motor nerve ending. across the synaptic deft, and its effects on the
(A) The nerve action potential allows calcium ions
to cross the presynaptic membrane and to enter the postsynaptic membrane.
nerve ending. (B) Vesicles then fuse with presynaptic At the postsynaptic membrane, the acetyl
membrane, open, and release chemical transmitter choline reacts with molecular sites called acetyl
contents into synaptic cleft Chemical transmitter choline receptors. This reaction opens membrane
molecules then diffuse across to combine with recep channels to both sodium and potassium ions.
tor sites on postsynaptic membrane; the chemical fit
between transmitter and receptor is shown Both ions then diffuse across the muscle mem
schematically. brane, seeking their electrochemical equilibria.
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
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
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
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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n
E
T h resh o ld
R e stin g Potential
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
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,
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.
tree shown in figure 4 -1 3 has five possible

Presynaptic Inhibition
synaptic sites, site 1 being closest to the cell
body and site 5 being most distant. When Postsynaptic inhibition acts via an effect on the
excitatory synaptic inputs (indicated by e) and postsynaptic membrane (the IPSP). In presy
inhibitory synaptic inputs (indicated by i) naptic inhibition, on the other hand, the
occur at various combinations of sites, the inhibitory neuron synapses onto the presy
computed synaptic potential (or EPSP) re naptic terminal of an excitatory neuron (figure
corded at site 1 shows that the inhibitory input 4 -1 4 ), and its effect is to reduce the amount of
has little effect when significantly distal to the transmitter released by the presynaptic terminal
excitatory input (A). When both inputs are at with each nerve impulse. This reduces the size
the same middle site (B), the recorded EPSP is of the resulting EPSP without causing any
transmitters are acetylcholine and norepineph

rine (and its close relative epinephrine). A
number of other compounds are presumed to be
transmitters on the basis of several experimen
tal criteria, but they are not yet accepted by all
observers as proven transmitters; these pre
sumed transmitters include dopamine, seroto
nin, and the amino acids GABA and glycine.
More recently discovered are the enkephalins
and endorphins, which are peptides that appear
to attach themselves to the same receptor site
as the drug morphine and may share some of its
effects as a pain-relieving (analgesic) sub
stance. The structures of these compounds are
Figure 4-14. Presynaptic inhibition, involving axoax-
anal synapses of inhibitory axon (I) onto excitatory shown in figure 4-15.
axon terminal (E).
direct change of postsynaptic membrane con Acetylcholine (ACh)

ductance and also without affecting the re Acetylcholine is synthesized in neurons by a
sponse of the postsynaptic cell to other synaptic reaction between acetyl coenzyme A (CoA)
inputs. Thus, presynaptic inhibition may func and choline, catalyzed by the enzyme choline
tion in the CNS to reduce the effectiveness of acetylase:
particular synaptic inputs onto a neuron, with
out altering the effects of other inputs. choline
The structural basis of presynaptic inhibition Acetyl CoA + Choline * Acetylcholine
acetylase
is thought to be the synapse of an inhibitory
axon (I in the diagram) onto an excitatory
axon terminal (E), corresponding to the axoax- The transmitter is stored in vesicles in the
onal synapses observed under the electron presynaptic endings and then released when an
microscope. The inhibitory axon releases a action potential sweeps over the endings (figure
chemical transmitter that binds with receptor 4 -1 6 ). After release, the transmitter diffuses
sites in the excitatory axon terminal. If the across the synaptic deft to combine with
excitatory terminal is invaded by a nerve acetylcholine receptor sites on the postsynaptic
impulse within a short time interval it releases side.
fewer vesicles of its chemical transmitter than it The transmitter released into the synaptic
normally would. cleft is quickly removed or inactivated, allow
ing the transmitter secreted later to have a clear
effect. In the case of acetylcholine, this re
moval and inactivation is accomplished by (1)
Chemical Transm itters diffusion of acetylcholine away from the sy
Chemical transmitters are substances that are napse and (2) enzymatic breakdown (hydroly
released by a presynaptic nerve ending after its sis) into acetic acid and choline, under the
stimulation, and cross the synaptic cleft to influence of the enzyme acetylcholine esterase
cause either excitation or inhibition in the (cholinesterase). The products of the break
postsynaptic cell. The best-known chemical down are then actively taken up by the presyn-
HCHpVCH-CHj-NHj
9 PHi
o ^ -o -C H r C H -N ^ a ^ H O -U o H
CH3
NOREPINEPHRINE
ACETYLCHOLINE
HO- -ch-ch2-n h -ch 3

H O -ip V C H ^ -N ^
HO OH
H O ~k^
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
Figure 4-15. Major chemical transmitters and pre

sumed transmitters.
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.
aptic terminal and recycled to produce more

Norepinephrine (NE)
acetylcholine.
Cholinergic neurons secrete acetylcholine as a Norepinephrine (also called noradrenaline) is
transmitter; they include the following (figure synthesized by a series of reactions involving
4-17): several enzymes as shown in figure 4-18. Like
acetylcholine in other nerve terminals,
1. Motoneurons (both in the spinal cord and in norepinephrine is stored in vesicles to be
the cranial motor nuclei) and probably other released during a nerve impulse (figure 4-19).
neurons in the CNS Inactivation of the released transmitter (clear
2. Preganglionic fibers of the autonomic system ing the synapse so it can respond to the next
(both sympathetic and parasympathetic) impulse) occurs by means of (1) diffusion of
and those that innervate the adrenal norepinephrine away from the synapse; (2)
medulla (an organ with a similar embry- enzymatic breakdown to an inactive metabo
ologic origin to the sympathetic ganglia) lite, involving the enzyme catechol-O-methyl
3. Postganglionic fibers of the parasympathetic transferase (COMT); only a small amount of
system norepinephrine is inactivated by this pathway;
4. Some specific postganglionic fibers of the and (3) active re-uptake by the presynaptic
Figure 4-17. Distribution of transmitters acetylcholine those arising in certain brainstem regions
(ACh), norepinephrine (NE), and epinephrine (E). (locus ceruleus and reticular formation)
There is evidence linking the CNS effects of

terminal, in which the norepinephrine can be norepinephrine with such complex functions as
recycled into storage vesicles and released alertness, learning, sleep, and mood.
again, or changed to an inactivate metabolite
under the influence of the enzyme monoamine
oxidase (MAO). Dopamine
Adrenergic neurons or secretory cells are those Dopamine is the precursor of norepinephrine in
that secrete norepinephrine (noradrenaline) as neurons that secrete norepinephrine. Certain
a transmitter; they include the following (figure identified neurons, however, secrete dopamine
4-17): directly. Important examples are some neurons
in the basal ganglia whose degeneration is
1. Postganglionic fibers of the sympathetic sys related to the movement disorder of Parkin
tem (with the exception of a few described son’s disease. Dopamine-secreting neurons also
under acetylcholine) project to the cerebral cortex and limbic system
2. Secretory cells in the adrenal medulla, and may be involved in emotional responses.
which are analogous to postganglionic
neurons of the sympathetic system and se
Serotonin
crete a mixture of norepinephrine and
epinephrine Serotonin (5-hydroxytryptamine or 5-HT) may
3. Some neurons in the CNS, particularly be an inhibitory transmitter secreted by neurons
dence suggests that they are CNS transmitters.

PHENYLA LAN IN E GABA may be released as a transmitter by
inhibitory intemeurons in the brain (for exam
Hydroxylase
ple, in the cuneate nucleus of the somatic
sensory pathway, in cerebellar cortex, and in
i r cerebral cortex) and in the spinal cord (specif
ically, those intemeurons responsible for pre
TYR O SIN E
synaptic inhibition). Picrotoxin, a convulsion-
inducing drug, may act by blocking the inhibi
Hydroxylase
tory effect of GABA. Glycine may mediate
the effects of intemeurons that cause IPSPs in
' r
spinal motoneurons. Strychnine, another con
DOPA vulsion-inducing drug, may act by antagoniz
(D ihydroxyphenylalanine)
ing the inhibitory effects of glycine.
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,
NOREPINEPHRINE (NE) AS A
TRANSMITTER
-S Y N T H E S IS
PRESYNAPTIC
CELL
Action potential
Inactive metabolite
II
SYNAPTIC CLEFT Inactive metabolite
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.
Glutamate as an excitatory transmitter causing EPSPs;

GABA as an inhibitory transmitter causing IPSPs
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.
Chapter 4
Synaptic Transmission
Postsynaptic Potential Action Potential
Chemically-gated channel Voltage-gated channels
Vm Cm G k & (jNa Vm Cm Gk Gnh

Na © channels Na ©
ACh
OUt 0 Receptor I channel

lipid
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
Review Questions
1. Diagram a neuromuscular junction, labeling the presynaptic terminal, synaptic vesi

cles, and post-synaptic membrane Diagram a neuron-neuron synapse similarly.
2. Describe the sequence of events during neuromuscular transmission.

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.
6. Describe changes in membrane potential and ionic conductance associated with

EPSPs and IPSPs.
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.
11. Relate the characteristics of postsynaptic potentials to summation.
12. Compare postsynaptic inhibition with presynaptic inhibition.
13. State the transmitters at the following synapses: (1) neuromuscular junction
(skeletal muscle), (2) autonomic ganglia, (3) sympathetic effectors, and (4)
parasympathetic effectors.
14. Describe the effect of curare on neuromuscular transmission.

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.
16. Explain why an anticholinesterase agent is sometimes given to counteract the

residual effects of tubocurarine after surgery.
86 Somatic Sensation
reduced in certain conditions, for example,

when the peripheral nerve is constricted by
connective tissue (in the median nerve this is
often associated with carpal tunnel syndrome).
Physiological Classification of Sensory-Nerve

Fibers
Large peripheral nerves, such as the sciatic,

have been removed from animals and
Figure 5-1. Testing conduction velocity in a sensory stimulated at the peripheral end by an electrical
nerve. Stimulus applied over digital nerves elicits pulse (figure 5—2). Near the other end, a
compound action potential, which can be recorded recording electrode (a metal wire) picks up the
from skin over median nerve. S, stimulus. (Method electrical signs of the compound action poten
described in Smorto, M.P., and Basmajian, J.V.
tial. Its reference electrode is placed a few
[32].)
centimeters away, over a section of nerve that
150 mm. The conduction velocity, then, is has been crushed; this ensures that no nerve
impulses activate the reference electrode, so
velocity = distance/time = 150 mm/3 msec that it can provide an inactive baseline from
= 50 m/sec which to measure activity under the recording
electrode. Unlike an individual action poten
The conduction velocity may be significantly
tial in a single nerve fiber, the compound
action potential increases in size as stimulus
Figure 5-2. Recording the compound action potential intensity is increased. This occurs because more
of a peripheral nerve. Inset: Diagram of the experi
mental method S, stimulating electrode; R, recording and more nerve fibers are being activated, and
electrode (reference electrode is at right end). the compound action potential represents a
msec
summation derived from these individual

responses. A
Also unlike the individual action potential,

a compound action potential may have several
DIAM.
peaks, because the nerve contains several /tm
8 - - - m - ,
groups of fibers with different conduction ve 4
r • jB f e s g a i a i K : =
locities. Imagine that the nerve fibers are 0 ■ -------------------
----------------w
runners in a race. As they cross the finish line 0 25 50 100
(the recording electrode), the number arriving C O N D U C T IO N V E L O C IT Y M /SEC .
at any given fraction of a second after the
starting gun are recorded on a chart. The next
Figure 5-3. Typical conduction velocity (in m/sec)
fastest group arrive bunched together and form a and fiber diameter (m |xm) for different groups of
peak at the beginning of the chart. The next afferent nerve fibers.
fastest group come in a few milliseconds later
and form a second peak, and two or three more Ill are small-diameter myelinated fibers, and
groups of stragglers provide the later peaks. The Group IV are unmyelinated fibers. The rela
first and largest peak represents activity in the tionship of classification, conduction velocity,
Group I fibers, the fastest conducting of all, with and fiber diameter is diagramed in figure 5-3.
a velocity of around 100 m/sec. The next peak Furthermore, and most important in study
represents activity in the Group II fibers, not ing somatic sensation, the different fiber groups
quite as rapidly conducting at about 50 m/sec. have different functions. The Group I fibers
Peaks due to Group III (around 25 m/sec) and carry signals from muscle and tendon receptors,
Group IV (about 1 m/sec) follow. which are important for controlling posture and
The peaks in a compound action potential movement; they will be discussed in chapter 9.
have also been classified alphabetically. In The Group II fibers carry sensations from
general, Group I corresponds to the A-alpha receptors for touch, pressure, vibration and
peak; Group II to A-beta; Group III to A- joint position and movement, and are discussed
delta; and Group IV to C. In addition to below. The other two groups, which carry
conduction velocity, the fiber groups differ information about pain and temperature (as
structurally under a microscope: Group I are well as crude touch), will be discussed later in
the largest-diameter myelinated fibers, Group this chapter. Table 5 -1 summarizes the types of
II are large-diameter myelinated fibers, Group afferent fiber and their major functions.
Table 5—1. Afferent Fiber Types and Their Major Functionsa

Typical Conduction Velocity
Group Type Description and Range (in msec) Major Functions
Group I A-alpha Largest-diameter myelinated 100 (72-120) Muscle feedback from
primary muscle stretch
receptors and Golgi tendon
organs
Group II A-beta Large-diameter myelinated 50 (36-72) Touch, pressure,
vibrations, and kinesthesis
Group III A-delta Small-diameter myelinated 25 (6-36) Fast pain, temperature,
and crude touch
Group IV C Unmyelinated 1 (0.5 -2 ) Slow pain, temperature,
and crude touch
*Source: Adapted from W.D. Willis and R.G. Grossman. Medical Neurobiology, 2nd ed. St. Louis: The C.V . Mosbv Co
1977.
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
CEREBRAL HEMISPHERES
AVUEG
Somatic sensory cortex
Thalamus
(VPL nucleus)
Medial lemniscus
MIDBRAIN
Dorsal column nucleus
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
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
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
D orsal colum n fib e rs'
D orsal colum n n u c'leu s

Figure 5 -7 . A possible consequence of a lesion to the of parietal lobe association cortex. This area
right parietal association cortex. has importance in the perception of bodily
sensations. Anatomically, it receives direct or
In both cases, intemeurons within the cuneate
indirect projections from the postcentral gyrus.
nucleus inhibit the neurons surrounding the
One indication of this function is given by the
most excited neuron and allow it to stand out.
clinical observation that damage to the poste
The amount o f surround inhibition can also
rior parietal association cortex on the right
be affected from higher centers in the brain.
hemisphere (figure 5 -7 ) often leads to defec
For example, nerve fibers have been traced
tive perception of sensations from the opposite
from the somatic sensory cortex to the cuneate
(left) side of the body. The patient may fail to
nucleus. Activation of the descending fibers
recognize the left side of his body as his own; he
can affect the inhibitory intemeurons and alter
may foil to dress it, shave the left side of his
the degree of surround inhibition in the
face, or comb the left side of his hair. He may
cuneate nucleus. We can speculate that atten
also fail to perceive objects to the left in his
tion directed toward a fingertip might increase
field of vision. A reasonable conclusion from
the amount of surround inhibition and increase
this syndrome is that the posterior parietal
the ability to differentiate between closely
association cortex normally functions to inte
spaced stimuli. Similar mechanisms occur in
grate somatic and visual sensation into a com
the gracile nucleus, the VPL thalamic nucleus,
plete body image. Similar lesions on the left
and the postcentral gyrus.
hemisphere may produce similar defects on the
Functions o f Posterior Parietal Association Cortex. right side of the body, although often less
Posterior to the postcentral gyrus is a large area marked and mixed with language disorders,
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)
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.
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.
Pharmacological Relief of Pain thalamic nuclei [2], structures that function in

the control of consciousness (chapter 11) as
Drugs given for die relief of pain fall into
well as in the neural response to painful stimuli.
several main categories.
Non-narcotic analgesics, such as aspirin and
acetaminophen, may be useful in relatively Psychophysiological Relief of Pain
minor painful conditions such as headache.
There is evidence that they act peripherally to Even in animals, the response to pain is not
reduce the synthesis of prostaglandin, a sub simply a matter of stimulus intensity alone;
stance that sensitizes free nerve endings to human contact and the presentation of food
painful stimuli. have diminished the response greatly, under
Local anesthetics, such as procaine, are used certain conditions. In humans, pain has been
to block nerve conduction either in the periph altered or relieved by several psychological and
eral nerve, the dorsal roots, or the spinal cord psychophysiological procedures.
itself, depending on where the drug is applied. Placebos are pills or other medications con
Consciousness is not normally affected. taining only presumably inactive ingredients,
Narcotic analgesics, such as morphine and such as sugar. Susceptible individuals have
codeine, are opiate drugs. They tend to subdue reported less pain after taking a placebo medi
the affective reaction to the pain, even though cation— in one test, nearly half the number of
the painful sensation may remain. They may patients who reported less pain after taking a
also produce sedation, euphoria, and anxiety narcotic analgesic. The power of suggestion
reduction. Opiate drugs have been studied by induced by both the physician and the drug is
radioactive labeling of their antagonist nalox probably behind the placebo effect.
one. The labeled antagonist has been found to Suggestion and hypnosis are also effective in
bind with opiate receptor sites in the dorsal selected individuals. Some patients have
hom of the cord, the reticular formation, and undergone dental work or even major surgery
the nonspecific thalamic nuclei, and also in the under hypnosis without anesthesia. Hypnosis is
hypothalamus and amygdala, areas associated also sometimes used to alleviate the subjective
with emotional and autonomic response pat experience of pain, including that of terminal
terns. The opiate refctptors also act as binding illness. Such treatment requires special training
sites for met-enkephalin, a five-amino-acid and the recognition that individuals differ in
peptide found in pituitary tissue, and for endor hypnotic susceptibility.
phin, a polypeptide in human blood with the Experimentally, a warning prior to an elec
same five-amino-acid sequence. Enkephalin tric shock has been found to reduce the
was found to have analgesic effects when intensity of the response. Music or white noise
injected into rat brain, suggesting that it may (like that from an electric fan) coupled with
act as an endogenous analgesic substance. the suggestion that this kind of sound reduces
Tranquilizjng and antidepressant drugs may pain, has been found to reduce the pain
have favorable results in some patients by response experimentally and during dental pro
relieving the affective reaction to pain. cedures, a phenomenon called audio analgesia.
G eneral anesthetics, such as enflurane, In clinical practice, relaxation training and
halothane, and nitrous oxide, may have a anxiety reduction have been found to reduce
specific analgesic effect, and used during pain intensity in some instances. Both methods
surgery at adequate levels, they also act to are incorporated in various systems of natural
lower the level of consciousness. At appropri childbirth education; factual information about
ate doses these agents have been found to childbirth, relaxation training, and breathing
diminish the activity of neurons in the brain exercises cause a decrease in anxiety and an
Chapter 5
Somatic Sensation: Dorsal Column System
L R
Stimulus
Peripheral Nerve
Group Type Velocity Diameter Function (Modality)
I A alpha 100 m/s 16 p Muscle length & tension
n A beta 50 8 Touch, pressure, vibration, kinesthesis

Schematic diagram of superior view of dorsal column pathway in spinal cord and brain (top), and peripheral
nerve fibers involved (bottom). 3, 1 ,2 = Brodmann’s areas, RP = receptor potential, AP = action potential,
PN = peripheral nerve. Only representative values o f velocity and diameter are shown in table..
hapters
Somatic Sensation: Spinothalamic System

L R
Group Type Velocity Diameter Function (Modality)
ITT A5 25 m/s 4p Fast pain, temp.,crude touch
IV C 1 1 Slow pain, temp., crude touch
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.
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.
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.
5. Define surround inhibition for such a neuron. Describe the synaptic mechanisms
involved.
6. Define and describe the somatotopic mapping on the postcentral gyrus.
7. Describe the function of tire posterior parietal association cortex.
8. List the structural and functional differences between the DC-ML and the lateral
5. Define surround inhibition for such a neuron. Describe the synaptic mechanisms
involved.
6. Define and describe the somatotopic mapping on the postcentral gyrus.
7. Describe the function of tire posterior parietal association cortex.
8. List the structural and functional differences between the DC-ML and the lateral
9. List the differences between fast and slow pain.
10. Define referred pain. Describe a possible mechanism for it.
11. Define gating. Describe a possible mechanism for it
12. List and briefly describe pharmacological and psychophysiological means of pain
suppression.
6. Auditory and Vestibular Function
How neural mechanisms function in hearing and balance
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
can have the same fundamental, but with a

different combination of harmonics).
In addition to pure tones and complex
periodic sounds, another type of sound is called
noise. Noise has no recurring periodicity and is
composed of random combinations of numer
ous frequencies. Vacuum cleaners produce good
examples of noise.
Sound Conduction Through the Middle Ear

The middle ear (figure 6 -4 ) is an air-filled
Figure 6-1. Sound waves in front of an oscillating space between the eardrum or tympanic mem
loudspeaker cone, at one instant in time. brane and the inner ear or cochlea. The three
small middle-ear bones or ossicles (malleus,
incus, and stapes) connect the tympanic mem
so on; the higher frequencies are called har brane (eardrum) with the oval window (a
monics or overtones of the fundamental fre membrane in the cochlea). The middle ear has
quency F. These harmonics lend a quality (or three major functions: amplification, protec
timbre) that makes voices and musical instru tion, and equalization. These functions may be
ments unique (a middle C on a piano sounds expressed by the mnemonic APE.
quite different from a middle C on a cello; both
Amplification. The middle ear amplifies sound-
induced vibrations so that when they reach the
Figure 6-2. A microphone (center) converts sound oval window they are strong enough to over
pressure waves into electrical voltage, which con be come the impedance of the fluids that fill the
displayed as it changes m time (lower right). The cochlea. These fluids impede (or resist) vibra
sound shown is a pure tone (sine wave). T = the tions due to sound, just as a moving paddle
period of the sine wave (the duration in seconds of
each cycle). The vertical distance between peaks is a meets more resistance in water than in air.
measure of amplitude. Fortunately, the amplification of sound vibra-
sounds elicit reflex contractions in the two

1 cycle middle-ear muscles, tensor tympani (inserted
I i Amplitude on the malleus) and stapedius (inserted on the
stapes). The combined contraction increases
the rigidity of the ossicular chain and reduces
Time transmission, mainly for lower-frequency
sounds (below about 1,000 Hz). Since the
latency of the middle ear reflex is about 50 to
150 msec, there is no protection against very
brief, sudden sounds, but the cochlea is pro
vided some protection against sustained loud
sounds, which might be attenuated by as much
as 40 dB.
D
Equalization. The eustachian tube connects the
middle ear to the pharynx and is opened during
swallowing, yawning, and sneezing. At those
E times, it provides an open passageway between
the middle ear and the atmosphere which
allows steady pressure levels on both sides of
Figure 6 -3. Sound pressure plotted against time. (A) the eardrum to equalize. Such equalization is
A pure tone, of frequency F els or Hz. (B) A pure necessary for the most efficient transmission of
tone, of frequency 2F Hz- (C) A complex periodic
sound. During air travel, if cabin pressure
sound, which can be derived from the combination of
A and B. (D) A complex periodic sound from a becomes appreciably higher or lower than
musical instrument. (E) Noise. mddle-ear pressure and the eustachian tube is
not opened, sound transmission across the
middle ear is reduced, and pain may result as
tions that takes place across the middle ear is
well.
enough to overcome this impedance. The
vibrations are transmitted by the mechanical
linkage from the tympanic membrane through Bone Conduction
the ossicles to the footplate of the stapes in the
oval window. Most of the amplification is due In addition to the middle-ear pathway, sound
to the decrease in area between the tympanic can also be conducted to the cochlea directly
membrane and the footplate of the stapes. The through the bones of the skull, a process called
effective area of the stapes footplate has been bone conduction. When you hear yourself speak
estimated as only 1/14 the area of the tympanic or hum, it is through a combination of air
membrane (figure 6 -5 ). Since pressure equals conduction (via the air, tympanum, and mid
force per unit area, this results in an amplifica dle ear) and bone conduction (directly from the
tion (by about 14 times) of the pressure. A oral cavity through the skull to the cochlea).
small additional contribution may be made by The reason your voice sounds different on a
the lever arrangement of the ossicles. tape recorder is that only air conduction is
involved. Bone conduction can be tested by
Protection. The middle-ear muscles protect the placing a vibrating tuning fork or other object
cochlea against sustained loud sounds. Loud on the head.
Tensor tympani Stapedius

•muscle n N N n n . musde v'O'-X
Oval window
Basilar membrane
Tympanum
Round window
Outer earl Middle earj Inner ear (Cochlea)
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
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
Transduction in Auditory Receptors

The hair cells, located on the basilar membrane
and within the organ of Corti, are the auditory
"'■' m ■- En velope maxim um
receptors. A tonal stimulus causes maximal up-
and-down motion at a particular location along
the basilar membrane. As seen in cross section
(figure 6 -8 ), both the hair cells that rest on the D irection o f traveling w av e
basilar membrane and the stiff tectorial mem

Neuronal Responses in the Auditory System
Anatomy o f the Auditory Pathway. Auditory

nerve fibers have their cell bodies in the spiral
ganglion, and their peripheral endings synapse
on the hair cells (figure 6 -8 ). Their central
endings synapse in the medulla at the cochlear
nuclei. The auditory pathway ascends through
several nuclei in the brainstem and thalamus to
the cerebral cortex (figure 6 -9 ).
Auditory Nerve Fiber Responses. Auditory nerve

fibers end at synaptic structures at the base of
the hair cells. Receptor potentials in the hair
cells somehow elicit action potentials in the
nerve fibers. A stimulus of a particular fre
quency will cause maximal movement at a
particular place along the basilar membrane,
which will activate a group of auditory nerve
fibers that innervate that area of basilar mem
brane. Thus another aspect of the place theory
of hearing is that a given sound frequency will be
represented in the CNS by a particular group of
activated nerve fibers.
The electrical response of individual auditory
nerve fibers can be studied using tonal stimuli
and microelectrode recording techniques. For
Figure 6-7. Displacement envelopes for the stimulus
frequencies shown beside each curve; only the posi example, as shown in figure 6-10, a fiber might
tive half of the envelopes are shown. From Bekesy, respond to low-intensity tones of 1,600 Hz,
G. Experiments in Hearing. New York: McGraw- while low-intensity tones above and below that
Hill Book Co., I960. Reprinted with permission. frequency have no effect. The best frequency of
this fiber is 1,600 Hz, and the frequency
lear tube; their vertical motion is converted corresponds to the place innervated by the fiber
into a horizontal shearing force exerted on the on the basilar membrane. The fiber can also be
hairs of the hair cells. The hair cells behave excited by adjacent frequencies of greater in
like specialized mechanoreceptors, transducing tensity, particularly on the low-frequency side.
the horizontal force into a receptor potential. A plot of all the frequency-intensity combina
The receptor potential from numerous hair tions that yield above-threshold responses is
cells is the probable origin of an electrical called the tuning curve of the fiber; it resembles
wave, called the cochlear microphonic (CM), the receptive field of a receptor on the skin or
which can be recorded in or near the cochlea. the retina. The tuning curves of two represen
With a pure tone (sine wave) stimulus, the tative fibers are shown in figure 6-10; if a larger
cochlear microphone is a sine wave of identical sample of timing curves were shown, they
frequency, similar to the electrical output of a would cover the entire range of audible
microphone exposed to the same sound. frequencies. Increased sound intensity, corres-
I l l Auditory and Vestibular Function
Reissner’s m em brane
■Scala media (endolymph)

Scala vestibuli
(perilym ph)
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
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
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.
mately 70 to 80 dB SPL, and a pneumatic drill

Figure 6-12. Localization of a transient sound. Path
lengths of the sound are shown in centimeters. is about 120 dB SPL. Auditory discomfort
Sound-pressure waves as they arrive at both ears are
(tickle, touch, or pain sensations in the ear) is
shown in simplified form: note time delay At and
amplitude difference between left (L) and right (R)produced by intensities of 120 to 140 dB SPL.
ears. Within the frequency range of greatest sen
sitivity (1,000 to 4,000 Hz), we can discrimi
nate between tones only 2 to 6 Hz apart. The
Decibels calculated on this basis are called human speaking voice overlaps this frequency
decibeIs sound-pressure level (dB SPL). In the range, extending from 200 Hz (slightly below
most-sensitive hearing range, if the threshold middle C) to 4,000 Hz.
of hearing is at a sound pressure, P, equal to Po,
then the threshold sound intensity is expressed
as: Audiometry: The Measurement of Hearing. An
audiometer is a device used for testing hearing
P by providing tones at various frequencies to
20 log — = 20 log 1 = 0 dB SPL
Po each ear separately, generally via earphones.
At each frequency, intensity is varied until the
For frequencies on either side of the 1,000 to threshold of hearing is found. Results are
4,000 Hz range, greater sound intensities are plotted in the form of an audiogram, a graph of
required to reach the threshold of hearing. hearing loss as a function of frequency (figure
The range of sound intensities present in 6-14), in which zero dB is commonly defined
everyday life is' immense. Although under ideal as the average threshold intensity for each
conditions, a tone of zero dB SPL can just be frequency, obtained from a group of normal
heard, normal conversation ranges from 40 to subjects. Hearing loss is then defined as the
60 dB SPL, heavy traffic on a city street or number of decibels above this value required to
amplified music on a dance-floor is approxi reach the threshold of hearing for the particular
counducted through the air, transmitted

160 through the tympanic membrane and the mid-
140 Discomfort
ffi
2 dle-ear ossicles to the cochlea. Bone conduction
120 Feeling threshold '
© refers to sound conducted from a vibrating
3 100
to
source placed on the mastoid or other bone,
© 80
Q.
60
conducted directly through the skull to the
T3
C
3 40 cochlea. Although bone conduction is less
O
CO 20 efficient, the pattern of vibration set up in the
Hearing threshold
0 cochlea is the same as that for air conduction.
I
o o
i
o
i____ J __i
CM ■» O 8 Hearing loss falls into two major categories.
T“ ii (1) Conduction or middle-ear deafness is indi
Frequency (Hz) 5 8'
cated when air conduction is impaired but bone
conduction is normal. This could be due, for
Figure 6—13. Hearing sensitivity. Frequency and in example, to calcium deposits making ossicles
tensity range of ordinary speech indicated by the rigid, a condition called otosclerosis, which can
dashed outline.
often be treated surgically. The normal bone
conduction indicates that the cochlear receptor
ear being tested. These are relative units, mechanisms and the neural pathway is func
which should not be confused with dB SPL. tional. (2) Sensorineural deafness is indicated
Hearing may be tested for two types of sound when both air conduction and bone conduc
conduction. Air conduction refers to sound tion are impaired and involves impaired func
tion either in the cochlea or in the auditory
Figure 6-14. An audiogram of a patient with hearing pathway. For example, certain antibiotics may
impairment, for stimulation of one ear. Bone con cause degeneration of cochlear hair cells; con
duction is normal (near the 0 dB hearing level), but tinued exposure to an intense high-pitched
air conduction is not, suggesting some degree of con
sound could lead to degeneration of hair cells in
duction deafness.
the corresponding part of the basilar membrane
(near the oval window) and hearing loss speci
FREQUENCY IN HERTZ
250 500 1000 2000 4000 8000
fically for the frequencies in that sound; and
N ORMAL 0
neural lesions in the brainstem can interrupt
0
►
— : » the auditory pathway.
10 10
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
5i 5.8 5.9 7.6 9.2

1.9 3.0 4.1
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
(M EDULLA) C O M P LE X (PO NS) (T H A L A M O
(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
Bulletin #X740L77 Prepored by E. Gross and J . J . Stockord, M .D . ou //vcv- w w .. u .* .a .
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
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).
potential, called the brainstem auditory evoked

response or simply the brainstem evoked re The Semicircle Canals During Rotation
sponse, is composed of a sequence of low-
The vestibular apparatus consists of three
amplitude (less than one microvolt) potentials
semicircular canals, plus the utricle and sac
during the initial 10 msec following click
cule. The semicircular canals resemble three
stimulation. These waves have been found to
hollow rings, each in a different plane approxi
be altered in various neurological disorders
mately perpendicular to the other two (figure
affecting the brainstem auditory pathway. On
6-16). They function to detect rotation, or,
this basis, wave I is thought to represent
more properly, rotational acceleration. Each
activity in the auditory nerve and wave V
canal is filled with fluid (endolymph). In each
activity in the midbrain portion of the auditory
canal, an enlarged region (the ampulla) con
system. The brainstem auditory-evoked re
tains a sensory structure consisting of receptors
sponse has been found useful in evaluating
(hair cells) whose sensory hairs protrude into a
hearing loss and suspected lesions of the brain
gelatinous cap (the cupola). If the head rotates
stem. It also demonstrates the rapid sequence of
in the plane of the canal, the enclosed fluid,
events in normal hearing: transmission from
having inertia, lags behind the canal and bends
earphones to inferior colliculus takes only
the hairs in a direction opposite to the move
about 6 msec.
ment. For example, if the head rotates to the
right (clockwise as viewed from above), the
hairs in the horizontal canals are bent in a
The Sense of Balance
counterclockwise direction. It may help to
The sense of balance is closely related to the think of the hairs as passengers in a whirling
sense of hearing: its sensory organ, the vestibu ring at an amusement park: rotation of the ring
Slow phase le ft (fo llo w e d by corrective saccode {fast phasel right)
Endolym ph moves to left,

stim uloting am pulla
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
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.
The Saccule and Utricle During

Linear Acceleration
Like the semicircular canab, the saccule and
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.
2. Explain the three functions of the middle ear.
3. Compare bone conduction and air (or ossicular) conduction.

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.
5. Describe how sound frequency corresponds to the location of envelope peaks of

the travelling waves.
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).
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.
5. Describe how sound frequency corresponds to the location of envelope peaks of

the travelling waves.
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).
8. Describe auditory nerve-fiber responses, using the terms bestfrequency and tuning
curve.
9. Define phase locking of auditory nerve fibers.
10. Describe two mechanisms of sound localization.
11. Explain why a unilateral lesion in the auditory cortex does not lead to complete
deafness in the contralateral ear.
12. What is the frequency range of normal hearing?
13. Define threshold of hearing. How does it differ across frequencies?
14. Define dB.
15. Describe a brainstem auditory-evoked potential and its clinical use.
16. Compare middle-ear deafness and sensorineural deafness

12. What is the frequency range of normal hearing?
13. Define threshold of hearing. How does it differ across frequencies?
14. Define dB.
15. Describe a brainstem auditory-evoked potential and its clinical use.
16. Compare middle-ear deafness and sensorineural deafness

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.
19. Describe the normal stimulus to the saccule and utricle.

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
Figure 7-1. Diagram for finding your blind spot.
Figure 7-3. Right eye viewed from the front.

Refraction
In a camera (figure 7 -5 ), the lens focuses light in figure 7-6B. The top prism bends the light
onto the film. In the eye, the optical system, downward, the middle section lets the light pass
composed of cornea—aqueous humor—lens- through without bending, and the bottom prism
vitreous humor, focuses light onto the retina. bends the light upward. At a certain distance
In both cases, focusing of light is based on past the prisms the light waves come together
refraction. and intersect.
A simple example of refraction is found in a
triangular glass prism (figure 7-6A ). As light
waves enter the glass, they slow down and also Figure 7-4. Right eye as viewed from above, in
bend. This slowing and bending of the light is horizontal section. The x and the arrow are objects
called refraction. Now consider two prisms with a in the outside world (visual field) focused on the
fovea and nearby retina, respectively. As in a cam
rectangular piece of glass in between arranged as era, the image is reversed. Not drawn to scale.
Figure 7-2. Right eye os viewed from above, in

horizontal section, while fixated on x. When the cir
cle is focused on the region shown it cannot be seen.
Simplified diagram is not drawn to scale.
131 Vision
Refractive power (diopters) = Ilf (meters)
The refraction itself occurs only at the interface

between two different substances. In the case of
the simple lens, the first refractive surface is the
interface between air and the front glass sur
face, and the second is the interface between
the rear glass surface and air. In a camera, the
Figure 7-6. Prisms illustrate refraction. (A) Trian

gular prism. (B). Three prisms combined. (C)
Biconvex lens.
Figure 7-5. A camera viewed from above. Its lens

focuses light into the film, just as the optical system
of the eye focuses light onto the retma (compare
with figure 7-4).
The three-prism arrangement can be con

sidered a crude first step toward developing a
biconvex lens (figure 7-6C ). In the lens, as in
the prisms, the top section bends the light rays
downward, the center section allows them to
pass straight through, and the bottom section
bends them upward. In the lens, the light rays
come together, or converge, to form a point of
light at a certain distance from the lens. If the
light originates at a distant point, the distance
from the lens at which the light converges is
called the focal length of the lens. The greater
the ability of the lens to bend light (its
refractive power), the shorter its focal length.
In fact, the refractive power (in diopters) is
equal to 1If, where f is the focal length in
meters:
132 Vision
distance between the lens and the film surface

is adjusted to equal the focal length when
distant objects are being photographed. The
object is then focused on the film. Each point
of light in the object forms (ideally) a corres
ponding point of light on the film, and these
points on the film together constitute an image
of the object. Compared to the object, the
image is reversed in direction and upside-down.
A good camera contains a compound lens,
composed of several lenses cemented together
and thus several refractive surfaces. Similarly,
the eye has several refractive surfaces as follows:
air / cornea / aqueous humor / lens / vitreous

humor
Most of the refraction actually takes place at

the first refiactive surface, between air and the
anterior of the cornea. The major contribution
of the lens is to provide adjustable refraction.
A distant object (say, a star or a penlight
Figure 7-7. Common refractive errors and their cor
from 50 feet away) will form a point focus on rection. (A) Myopic eye. (B) Hyperopic eye. (C)
the retina, approximately 24 mm behind the Emmetropic eye (normal refraction).
cornea of a normal, relaxed eye. A simplified
optical model of such an eye has a focal length
of about 17 mm; its refractive power is 1/0.017
meters or approximately 59 diopters. apart) the light waves and allows them to come
to a focus on the retina.
In hyperopia (figure 7-7B ), distant objects
from an image behind the retina; again, at the
Refractive Errors
retina itself, the image is blurred. Hyperopia
Difficulty in clearly focusing images can be due may be called farsightedness because the prob
to three major sources of refractive error: lem is worse for near than for faraway objects. It
myopia, hyperopia, and astigmatism. arises because the eye is too short for the
In myopia (figure 7 -7 A), distant objects refractive power of the lens. Hyperopia can be
form an image in front of the retina; at the corrected with a convex lens, which converges
retina itself, the image is blurred. People with the light waves and allows them to come to a
myopia are also called nearsighted because they focus on the retina.
have difficulty seeing distant objects clearly, In astigmatism, the curvature of the cornea
while nearby objects can often be seen very (or other refractive structure) may be greater in
well. Myopia arises because the distance be one direction (e.g., vertically) than in another
tween cornea and retina is too long for the (e.g., horizontally). The image of a point then
refractive power of the eye. It can be corrected becomes an ellipse on the retina. In the
with a concave lens, which diverges (spreads example given, the problem can be corrected
133 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
Sym pathom im etic substances P a ra s y m p ath o m im etic substances

(dilate pupil) (c o n s tric t p u p il)
Amine oxidase inhibitors (cocaine) Cholinesterase inhibitors (erserine,

Noradrenaline-like substances Prostigmin, DFP)
(Paredrine, amphetamines) Acetylcholine-like substances (pilocarpine)
I Noradrenaline I
Sympathetic Parasympathetic
innervation innervation
Sym patholytic substances Parasympatholytic substances

(constrict pupil) (dilate pupil)
Dibenamine, Priscol, hexamethonium Atropine, homatropine, scopolamine
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
Sym pathom im etic substances P a ra s y m p ath o m im etic substances

(dilate pupil) (c o n s tric t p u p il)
Amine oxidase inhibitors (cocaine) Cholinesterase inhibitors (erserine,

Noradrenaline-like substances Prostigmin, DFP)
(Paredrine, amphetamines) Acetylcholine-like substances (pilocarpine)
I Noradrenaline I
Sympathetic Parasympathetic
innervation innervation
Sym patholytic substances Parasympatholytic substances

(constrict pupil) (dilate pupil)
Dibenamine, Priscol, hexamethonium Atropine, homatropine, scopolamine
Dilator Sphincter
136 Vision
sake of simplicity, be studied as a function of opsin. The retinene is enzymatically reduced to

input to one eye only (monocular vision) and vitamin A. The photochemical reaction causes
without reference to color. Binocular and color a visible bleaching of the deep purple rhodopsin
vision will be discussed later. to a colorless form. At the same time, the
reaction gives rise to a transduction from light
Transduction in the Retina to electrical energy: a receptor potential occurs
in the photoreceptors.
The optical apparatus of the eye focuses on the This receptor potential is most unusual. In
retina an image of the visual field, reduced to most receptors, it consists of depolarization,
the size of a postage stamp and inverted, as is due to an increased conductance to sodium and
the image focused on the film of a pocket other small ions. In the photoreceptors, how
camera. The varying intensities of light in this ever, it consists of a hyperpolarization— the
image are absorbed by the photoreceptors of interior of the cell becomes more negative—
the eye, called rods and cones. Specifically, the due to a decreased conductance to sodium.
light is absorbed by a visual pigment in the This hyperpolarization develops in a cell that,
photoreceptors; in the case of the rods, this in comparison with other receptors, is already
pigment is called rhodopsin. In the presence of partially depolarized, having a resting potential
light, a photochemical reaction takes place of -2 0 to -4 0 mV. The amount of hyperpolari
(figure 7-11) in which rhodopsin breaks down zation depends on the intensity of light: a weak
to a colorless pigment, retinene, plus a protein, light stimulus might cause a hyperpolarization
of only 2 mV (from —20 to -2 2 mV), while a
Figure 7-11. The photochemical reaction occurring in brighter stimulus might cause a hyperpolariza
the rods. Lumrrhodopsin and metarhodopsin are inter tion of 15 mV (from —20 to -3 5 mV).
mediate chemical forms produced by the action of
light on rhodopsin. Source: Reprinted with permission
Synaptic Actions in the Retina
from Brown, K.T., Physiology of the retina. In
Mountcasde, V.B. (Ed.), Medical Physiology The rods and cones synapse with two other cell
(12th edL). VoL 2. St Louis: The C.V. Mosby
types: horizontal cells and bipolar cells (figure
Co., 1968. Adapted from Wald, G., in Magoun,
H.W. (Ed.), Handbook of Physiology. Baltimore: 7-12). The presence of vesicles in the presy-
Williams & Wilkins, 1959. naptic region suggests that a release (or reduced
136 Vision
sake of simplicity, be studied as a function of opsin. The retinene is enzymatically reduced to

input to one eye only (monocular vision) and vitamin A. The photochemical reaction causes
without reference to color. Binocular and color a visible bleaching of the deep purple rhodopsin
vision will be discussed later. to a colorless form. At the same time, the
reaction gives rise to a transduction from light
Transduction in the Retina to electrical energy: a receptor potential occurs
in the photoreceptors.
The optical apparatus of the eye focuses on the This receptor potential is most unusual. In
retina an image of the visual field, reduced to most receptors, it consists of depolarization,
the size of a postage stamp and inverted, as is due to an increased conductance to sodium and
the image focused on the film of a pocket other small ions. In the photoreceptors, how
camera. The varying intensities of light in this ever, it consists of a hyperpolarization— the
image are absorbed by the photoreceptors of interior of the cell becomes more negative—
the eye, called rods and cones. Specifically, the due to a decreased conductance to sodium.
light is absorbed by a visual pigment in the This hyperpolarization develops in a cell that,
photoreceptors; in the case of the rods, this in comparison with other receptors, is already
pigment is called rhodopsin. In the presence of partially depolarized, having a resting potential
light, a photochemical reaction takes place of -2 0 to -4 0 mV. The amount of hyperpolari
(figure 7-11) in which rhodopsin breaks down zation depends on the intensity of light: a weak
to a colorless pigment, retinene, plus a protein, light stimulus might cause a hyperpolarization
of only 2 mV (from —20 to -2 2 mV), while a
Figure 7-11. The photochemical reaction occurring in brighter stimulus might cause a hyperpolariza
the rods. Lumrrhodopsin and metarhodopsin are inter tion of 15 mV (from —20 to -3 5 mV).
mediate chemical forms produced by the action of
light on rhodopsin. Source: Reprinted with permission
Synaptic Actions in the Retina
from Brown, K.T., Physiology of the retina. In
Mountcasde, V.B. (Ed.), Medical Physiology The rods and cones synapse with two other cell
(12th edL). VoL 2. St Louis: The C.V. Mosby
types: horizontal cells and bipolar cells (figure
Co., 1968. Adapted from Wald, G., in Magoun,
H.W. (Ed.), Handbook of Physiology. Baltimore: 7-12). The presence of vesicles in the presy-
Williams & Wilkins, 1959. naptic region suggests that a release (or reduced
137 Vision
could send information via a bipolar cell to a

ganglion cell, which would respond with a
depolarization (an EPSP) and a series of action
potentials.
At the same time, the adjacent ganglion cell
could respond with a hyperpolarization (an
IPSP) and a cessation of action potentials. The
inhibition of the adjacent ganglion cell could be
due to an inhibitory action of one or both of the
laterally-transmitting cells (the horizontal and
amacrine cells), and is an example of lateral
inhibition, similar to the mechanism in somatic
sensation. In brief, the outcome would be that
illumination of one receptor would cause excita
tion of the directly connected ganglion cell and
inhibition of surrounding ganglion cells. If we
record from a single ganglion cell while moving
the light stimulus, the ganglion cell would be
excited by illumination of a directly connected
receptor and inhibited by illumination of a
Figure 7-12. General scheme of cell types and sy
naptic connections in the retina. R = rods, B = neighboring receptor.
bipolar cells, G = ganglion cells, H = horizontal In reality, the ladder model is an oversim
cells, and A = amacrine cells, light falling on the plification; a number of photoreceptors can
receptive components of the rod causes synaptic and converge to synapse on a single bipolar cell,
spike potentials in ganglion cells as shown below.
and a number of bipolar cells can converge on a
After Dowling [6].
single ganglion cell. In addition, most stimuli
release) of a chemical transmitter is involved in illuminate more than one photoreceptor. This
the transmission of information from the leads to the more general result of a retinal
photoreceptors. The bipolar cells relay infor ganglion cell being excited by illumination of
mation to the ganglion cells, which in turn its more directly connected receptors and in
send information in the form of nerve impulses hibited by illumination of surrounding recep
along their axons, which bundled together tors.
make up the optic nerve. Sideways transmis Experimentally, most studies of ganglion
sion of information is handled by the horizontal cells have been done one cell at a time, while
cells, which connect adjacent photoreceptors illuminating various spots on the retina. For a
and bipolar cells, and by the amacrine cells, single ganglion cell, illumination of only a
which connect adjacent bipolar cells and gan small circular area of the retina, generally
glion cells. Thus, the basic circuit of the retina overlying the cell, causes either excitation or
can be visualized as two rungs of a ladder, with inhibition; this area is called its receptive field
the sides composed of the sequence of photo- (figure 7-13). In the visual system as a whole,
receptor-bipolar-ganglion cells and the rungs the receptive field of a neuron is “the area of
composed of horizontal cells and amacrine retina from which the discharges of that neuron
cells. can be influenced” [6]; this area of retina
With this simplified model in mind, we can corresponds to a particular area of the visual
see how a single illuminated photoreceptor field. Stimuli outside the receptive field have
138 Vision
no effect, although the cell may continue

Light S tim u lu s R e c e p t iv e Field N eural R e s p o n s e discharging spontaneously. For a ganglion cell,
illumination confined to the center of the
receptive field (presumably stimulating the
more directly connected photoreceptors) may
cause excitation (figure 7-13A and C) while
illumination confined to the concentric sur
round area may cause inhibition (the firing rate
may be reduced below the spontaneous level;
figure 7-13B). Thus the receptive field is
composed of an excitatory center and an
inhibitory surround. In other retinal ganglion
cells, the receptive field is reversed, with an
inhibitory center and an excitatory surround.
When the entire receptive field is illuminated
(figure 7-13D ), the surround response reduces
the center response.
Visual Acuity and Retinal Receptive Fields

LIG H T
Visual acuity— the ability to see detail— can be
measured by the Landolt C test, which tests the
Figure 7-13. Receptive field of a retinal ganglion cell ability to see the small gaps in the rings (figure
with excitatory center and. inhibitory surround.
7-14a). The person stands a fixed distance
Neural spike responses shown on right.
from the chart and is asked to indicate the
direction of the gaps in successive characters.
For measuring acuity, the size of the gaps is
Figure 7 - 14- Tests for visual acuity: (a) Landolt-C; translated into a visual angle. Under good
(b) and (c) two examples of letters from Snellen conditions, a person should be able to detect a
test. The gap or space, s, subtends a known angle gap that subtends an angle of one minute of arc
at the eye. Source: Davson, H., and Eggleton,
(1/60 of a degree). The gap in a small printed c
M.G. Principles of Human Physiology (14th edL).
Philadelphia: Lea & Febiger, 1968. Reprinted with in this text subtends such an angle at a distance
permission. of approximately 3.4 meters. The familiar Snel-
139 Vision
len Chart, with a series of letters on each line,

is used in a similar manner: a person with
normal vision, standing 20 feet from the chart,
can just recognize the letters on the line
marked 20 feet. The gaps in the letters (figure
7—14b and 7-14c) will then subtend a visual
angle of one minute; and the person will be said
to have a visual acuity of 20/20. These tests are
commonly given to test the optical function of
the eye but may also be used to test the visual
pathway.
Visual acuity can be compared with two-
point tactile discrimination. We have seen that
the two-point threshold is smallest on parts of Figure 7-15. Graph of visual acuity (solid line),
the body (fingertips, tongue) having small against visual angle (from the fixation point). Cone
concentration (dotted line) and rod concentration
receptive fields and a high density of receptors (dashed line) plotted against distance from fovea in
per square centimeter. Similarly, visual acuity degrees. Vertical axis shows relative units only.
varies over different parts of the visual field
and, thus, over corresponding parts of the
tory surround is an example of lateral inhibi
retina, and the variation can be related to
tion. As in somatic sensation, it causes an
changes in retinal receptive fields.
exaggerated neural response to edges. In vision,
The variation in acuity can be illustrated by
the edge effect can be seen in a series of strips
fixating a single letter on the printed page with
like those prepared by photographers testing
one eye covered; letters at increasing distances
exposure time in the darkroom (figure 7-16).
to either side will be more difficult to recognize.
Although each strip really consists of a con
The principle illustrated is that visual acuity is
stant light intensity, subjectively, it appears as
good at the fixation point but falls off rapidly
if the darker strip becomes darker near its
toward the periphery. This relationship is illus
border with the lighter strip, and the lighter
trated in the graph of figure 7-15.
strip becomes lighter near its border with the
Visual acuity is related to the size of the
darker strip. This can be explained by noting
receptive-field centers of retinal ganglion cells:
that the ganglion cell with a receptive field
small receptive-field centers are required to see
completely illuminated by the lighter strip has a
fine detail. The smallest receptive-field centers
slightly lower firing rate, and therefore leads to
occur in the foveal region and consist of one to
less perceived light intensity, than the ganglion
three photoreceptors only. With increased dis
cell having less of its inhibitory surround
tance from the fovea, the size of the receptive-
illuminated due to the impingement of the dark
field centers grows to several degrees, as a
border.
number of photoreceptors converge to activate
a single ganglion cell.
Rod and Cone Vision
The two kinds of photoreceptors, rods and
Edge Enhancement and Retinal Receptive-Field
cones, differ in several ways.
Organization
Distribution. The fovea itself contains only
The organization of ganglion-cell receptive cones. With distance from the fovea, as meas
fields into an excitatory center and an inhibi- ured in terms of visual angle (degrees from
140 Vision
distribution and light sensitivity of the rods. He

found that a dim star could be seen only if
viewed obliquely, a short distance from the
fixation point. This was later attributed to the
ability of rods rather than cones to respond to a
dim object, and to the presence of rods only a
short distance from the fovea rather than
within the fovea itself.
Dark adaptation is due in part to the resynth
esis of visual pigment (figure 7-11). In the
dark, the retinene and opsin in the rods
become converted into light-sensitive rhodop-
sin (in the cones a similar process leads to
Figure 7-16. Strips of stepwise increasing light inten synthesis of the cone pigment, photopsin). In
sity, demonstrating accentuation of edges (also called addition, large quantities of vitamin A are
the Mach band effect). Superimposed receptive fields converted to retinene, which is then available
of retinal ganglion cells illustrate a possible mecha for resynthesis of additional visual pigment.
nism for this effect Receptive field on left has less When the photoreceptors contain a higher
illumination of its inhibitory surround (minuses),
therefore its cell fires at higher rate than cell on concentration of rhodopsin and photopsin due
right to this resynthesis, they are more sensitive to
light. Without vitamin A, light sensitivity is
fovea), the concentration of cones per unit area poor, particularly in dim light, giving rise to
declines, and up to a point, the concentration night blindness.
of rods increases (figure 7-15). In the periphery Color sensitivity. The color vision we have in
of the retina, the photoreceptors are only rods. bright light is due to the cones. Rod vision
Light sensitivity. Vision in moderately bright allows us to see black, white, and shades of
light is a product of both rods and cones. Vision gray, as we do in very dim light.
in very dim light is due only to rods and In addition, the rods and cones respond to
requires a substantial delay for dark adaptation overlapping but not identical parts of the color
to occur. A good example of dark adaptation spectrum: the rods do not respond to pure red
occurs when you walk from a bright lobby into light (the longest wavelengths in the visible
a dark movie theater. Initially you can see very spectrum), while the cones do. Thus, a pure
little and only gradually does light sensitivity red light stimulus is a handy method for
increase enough to allow you to see well. When experimentally activating cones without rods.
this process is measured in the laboratory, it In one practical application, night-flying pilots
turns out that the time-course of the adaptation can avoid the 40-minute wait for dark adapta
is composed of two curves: the first reaches a tion by wearing red goggles for some time
plateau in 10 minutes and is due to dark before takeoff. Since only red light passes
adaptation of the cones, and the second through the goggles to the eyes, the rods are
reaches a plateau at a greater level of light not activated and adapt just as they would in
sensitivity in about 40 minutes and is due to darkness. After removing the goggles, the pilot
adaptation of the rods. After this 40-minute can immediately see in the dim light of the
delay, vision in dim light is essentially rod night sky using rod vision.
vision. Visual acuity. The greatest visual acuity (the
The astronomer Kepler made an interesting ability to resolve details separated by a small
observation that can be related to both the visual angle) is due to cone vision and is found
141 Vision
Table 7-1. Characteristics of Rod and Cone Vision

Characteristics Measurement Cones Rods
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
Light Stimulus Receptive Field Neural Response
A Spot in excitatory area
a
B Bar at optimal orientation
C Bar at sight angle to

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
Block of visual cortex
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
wavelength region (455 millimicrons); the

green-sensitive cones have maximal sensitivity
in the moderate-wavelength region (535 mp,);
and the red-sensitive cones have maximal
sensitivity in the long-wavelength region (575
mp.). Light stimuli often activate two or more
of these cone types at once. For example, a
yellow light of 560 mp, will activate both green
and red cones; the corresponding neural activ
ity leads to the perception of yellow. If pure
green and pure red light are made to fall upon
the same area of the retina (this can be done,
for example, by projecting a green and red
spotlight on the same part of a white screen) a
yellow color will result. Such additive combi
nation of colors should not be confused with
Figure 7-22. Sensitivity curves of blue-, green-, and the subtractive combination that occurs when
red-sensitive cones. B = blue, G = green, Y = paints are mixed.
yellow, O = orange, and R = red. Adapted from
Various forms of partial color blindness are
R .L Gregory. Eye and Btain: The Psychology of
Seeing (3rd ed.). New York: World University Li found in a small portion of the population. One
brary, McGraw-Hill Book Co., 1977- form, called protanopia, is explained by a lack of
red cones; another form, called deuteranopia, is
explained by a lack o f green cones. Both may
columns. Two of these ocular dominance areas, be associated with the tendency to confuse red
representing both right and left eyes, can be and green. Tritanopia, explained by a lack of
accommodated within about 800 p.. In addition blue cones, is rare. Also rare is total color
to monocular neurons, a large number of blindness, which can generally be explained by
orientation-specific neurons are binocular. Al the presence of only the pigment rhodopsin in
though they can be excited by either eye both rods and cones. People with this condi
separately, they are excited to a greater extent tion can perceive only various shades of gray.
by stimulation of approximately corresponding The three types of cones converge onto
locations on both retinas. The behavior of retinal ganglion cells and give rise to several
these neurons may provide a physiological basis types of response characteristics. For example,
for binocular vision. in one type, the center of the receptive field is
excitatory, and the surround is inhibitory,
regardless of the wavelength of the light
Color Vision stimulus. In another type, however, the center
and surround respond differently to different
In dim light most vision is by means of the rods colors: the center may be excited by red and
and involves little or no color vision. In bright inhibited by green, while the surround may be
light, cone function makes possible color vi excited by green and inhibited by red. A third
sion. Three different kinds of cones, each type of receptive field is differentially sensitive
containing a pigment most sensitive to light of to yellow and blue. These receptive-field types
a particular color, have been found in the of retinal ganglion cells represent further re
human retina (figure 7-22). The blue-sensitive finement of color processing in the visual
cones have maximal sensitivity in the short- system.
147 Vision
Specialized visual pathways for M and P cells
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
Table 8-1. Examples of Substances Eliciting the Four Taste Qualities
Sweet Sour Salty Bitter
Glucose Citric acid Sodium chloride Quinine

Fructose Acetic acid Potassium chloride Nicotine
Saccharin Tartaric acid Ammonium chloride Caffeine
D-leucine Hydrochloric Magnesium chloride Sodium citrate
acid
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.
■ ''' b i t t e r ' ' - n ix ■

ii
SOUR; SOUR
Smell
! n. VII
While the sense of taste responds to chemical
\ SALTY J
substances already in the mouth, smell re
SW EET sponds to chemical substances in the air, which
may emanate from a distant source. Thus
160 Chemical Senses
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.
Structure Odor Sensations and Sensory Responses

Olfaction is a scientific term for the sense of To elicit an odor sensation, a substance must be
smell. The olfactory sensory cells are located in in gas or vapor form and then soluble in the
an area at the top of the nasal cavity (figure 8 - mucous layer covering the olfactory epi
3). The sensory cells together with supporting thelium. The qualities and classes of odor are
cells form an olfactory epithelium. The sensory more numerous and less agreed on than those
cells have cilia projecting down into the mu for the sense of taste. The intensity of an odor
cous layer on the epithelial surface, and give off increases with the concentration of the odorous
axons that pass upward through holes in the substance. Odor sensation is notably subject to
cribriform plate, a section of the ethmoid bone adaptation: a faint scent of perfume may disap
of the skull, into the olfactory bulb. There they pear altogether after a few minutes of exposure.
synapse with axons of the large mitral cells, A slow negative wave can be recorded from
which are the principal cells in this synaptic the olfactory epithelium, in response to an
161 Chemical Senses
odorant stimulus, and reflects receptor poten

tials in the olfactory sensory cells. The wave
increases with concentration of the stimulus.
Like the taste fibers, individual olfactory-
receptor cells are stimulated by several sub
stances, more by some than by others. Action
potentials recorded from individual nerve fibers
in the olfactory tract either increase or decrease
in frequency in response to specific odorant
stimuli.
The synaptic organization of the olfactory
bulb has received much experimental attention
[31]. Intemeurons called granule cells seem to
lack axons; instead, their dendrites synapse
upon the dendrites of the mitral cells. Another
unusual feature is that the granule cells act by
means of graded potentials rather than action
potentials. Their effect is to inhibit the mitral
cells; this can be compared to the effect of
amacrine cells on ganglion cells in the retina.
163 Chemical Senses
Review Questions
1. Describe the taste receptors and their neural connections.
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.
4. Describe the olfactory receptors and their neural connections.

163 Chemical Senses
Review Questions
ally adaptive.

163 Chemical Senses
Review Questions
ally adaptive.

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?
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
How motor systems control the skeletal muscles
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
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
MUSCLE STRETCH REFLEX
EXTRAFUSAL
MUSCLE FIBER '______________ J . A L P H A , MOTONEURON
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
Table 9-1. Alpha and Gamma Efferent Fibers Compared

Type of Efferent Fiber Typical Diameter Typical Velocity Destination
Alpha 14 p. 80 m/sec Extrafusal

muscle fibers
Gamma 4 H 25 m/sec Intrafusal
muscle fibers
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
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
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-
INVERSE STRETCH REFLEX

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-
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
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.
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).
This method was used to compare

musicians who play stringed
instruments, those who play
keyboards, and non-musicians.
Nerve fibers in a human brain, shown by

DTI method .By Thomas Schultz (Own
work) [GFDL
(http://www.gnu.org/copyleft/fdl.html), CC-
BY-SA-3.0 (http://creativecommons.org)
The results suggested that nerve
cells can adapt to the demands of
playing a musical instrument, like a
guitar or violin, by developing more
myelin-insulated nerve fibers running
alongside each other as they
descend from the motor cortex to the
spinal cord (Ruber and others,
2015),
Is it possible that some of this

difference was inborn and inclined
the musicians to take up their
instrument in the first place? That
question is still unanswered.
Furthermore, in patients recovering

from stroke, white matter changes in
descending motor tracts correlated
with improvements in motor
impairment after a treatment course
of intensive physical therapy
combined with a new method of
noninvasive stimulation of the motor
cortex, called transcutaneous direct
cortical stimulation or tDCS (Zheng
and Schlaug, 2015).
References:
Rüber, T. and others, (2015), Cerebral

Cortex 25(6). Differential Adaptation of
Descending Motor Tracts in Musicians.
Zheng, X. and Schlaug, G. (2015).

Frontiers in Human Neuroscience, 30.
White matter changes in descending motor
tracts correlate with improvements in motor
impairment after undergoing a treatment
course of tDCS and physical therapy
These three descending pathways are often

called extrapyramidal pathways because they
M O TOR C O R T E X
travel outside the pyramidal tracts. In the
spinal cord, the rubrospinal tract travels along
side the lateral corticospinal tract, while the
reticulospinal and vestibulospinal tracts are
located more ventral and medial.
B R A IN ST E M
M O TO R A R E A S
Function. The major function of these path
ways is to gather information from other
sources, including motor cortex, cerebellum,
basal ganglia, and vestibular nerve, and use it
in the control of spinal motoneurons.
RF
The effects of the descending fibers in the
extrapyramidal tracts are largely exerted on
motoneurons indirectly through spinal inter
r r neurons. Both alpha and gamma motoneurons
are affected, and both excitatory and inhibitory
M U SC LE synaptic responses (EPSPs and IPSPs) have
been shown.
©1 © The best-understood effect is on the upright
posture. Standing upright involves contraction
ALPHA M O TON EU RO N
of the antigravity muscles, including those that
extend the leg and hold the head up, and, in
humans, those that flex the arm (in four-legged
animals the forelimb extensors are antigravity
Figure 9-7. Brainstem motor areas and the descend muscles). The stretch reflexes of these muscles
ing tracts that carry their output. R = red nucleus, are enhanced by activation of a large part of the
V = vestibular nuclei, RF = reticular formation; reticular formation and of the vestibular nuclei,
their descending tracts are, respectively, rubrospinal, at least in animals. Many of the descending
vestibulospinal, and reticulospinal tracts, and may be
grouped together as the extrapyramidal tracts or fibers at the level of the internal capsule or
extracorticospinal tracts. These tracts, labeled 3, in midbrain, originating in the motor cortex and
fluence alpha motoneuron along with corticospinal basal ganglia, normally function to keep the
tract, 2, and muscle stretch afferents, 1. brainstem motor areas under control. The loss
of these fibers after internal capsule or midbrain
cortex, cerebellum, and basal ganglia, lesions is followed by a release of function in
among other sources. It gives rise to the which the facilitating effects of brainstem
reticulospinal tracts, which descend in the motor areas are unchecked, leading to an
spinal white matter. exaggeration of the standing posture, hyperac
3. The vestibular nuclei, located in the pons, are tive tendon jerk reflexes in the antigravity
sensory nuclei of the vestibular nerve that muscles, and spasticity. The particular posture
also have a motor function. Their inputs are that results varies with the level of the lesion.
from the vestibular organs and the cerebel At the internal capsule level, the arms tend to
lum, and they give rise to the vestibulospinal be flexed. At the midbrain level, the arms tend
tracts, which descend in the ventral and to be extended. In both cases, the legs tend to
medial portions of the spinal white matter. be extended.
The Neurological Examination
Testing of reflexes is a part of the neurological examination, a systematic analysis of

nervous system function designed to detect neurological disorders and find their origin
Here is a brief, partial outline of the neurological exam ination
1. Mental status (see Chapter 11)

Orientation: time, place, situation, self
Attention: subtract serial sevens
Learning and memory
Memory: immediate, recent, remote
Language: comprehension, speech, repetition,
reading, writing, nam ing
Calculations
Information
Abstractions
Constructions
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
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 ).
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 ).
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
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:
Structure. The cerebellum is a large, roundish

structure protruding from the dorsal surface of 1. Dysmetria— a tendency to overshoot or
the brainstem at the pontine level. Like the undershoot a target, for example, knocking
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
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
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
ALPHA M OTON EURON
—V--
PN S CNS
Figure 9-11. A schematic diagram showing the over

all plan of the motor systems, combining figures 9 -
7, 9 -8 , and 9-10 (same abbreviations). Three ma
jor influences on alpha motoneuron activity are
shown: (I) Muscle stretch afferents (Group la fi
bers), from muscle stretch receptors. (2) Cortico
spinal fibers, from motor cortex. (3) Descending fi
bers in the rubrospinal, vestibulospinal, and reticulo
spinal tracts (excrapyramidal or extracortico-
spinal tracts), from brainstem motor areas.
speech is known to be significant in organizing

speech output.
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.
3. Describe the sequence of neuronal events on the muscle stretch reflex.
4. Describe the effects of activity in the gamma motoneurons.
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.
7. Define muscle tone, flacddity, spasticity, and clonus. Relate each term to spinal
reflexes.
8. Diagram and describe the inverse muscle stretch reflex.

9. Diagram and describe the flexor withdrawal reflex.
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).
11. Define reciprocal inhibition as applied to spinal reflexes.
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.
15. List the sequence of events in a voluntary 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.
18. Define the movement disorders of chorea, athetosis, and ballismus.

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
How sympathetic and parasympathetic activity help control the

internal environment
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
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
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
Chapter 10
Autonomic Nervous System
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.
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
to exercise, may enhance the diversion of blood Autonomic Denervation

flow to muscle.
Effector organs depend on their autonomic
innervation in a complex way. For example,
Gastrointestinal Tract after the cutting (denervation) of the sym
pathetic postganglionic nerves to a group of
Parasympathetic stimulation causes increased
blood vessels, the normal, tonic constriction of
glandular secretion, peristaltic activity, and
the blood vessels will be lost, and they will
sphincter relaxation in the gut; thus, the
dilate. After some time, however, a certain
digestive processes are generally enhanced.
degree of constriction will return, because of
Sympathetic activity has opposite effects on
the activity of the vascular smooth muscle
perjstaltic and sphincter activity, which to
itself. At that time, the effects of sympathetic
gether with its effect on gastrointestinal blood
stimulation produced by injecting norepin
supply can reduce or shut down the digestive
ephrine or a drug with similar effects will be
processes.
greatly enhanced, producing more constriction
than under normal conditions. This effect is
The Eye called denervation supersensitivity and is attrib
uted to an increase in the number of receptor
As described in chapter 7, parasympathetic
sites after denervation.
stimulation constricts the pupil while sym
pathetic stimulation dilates it. Parasympathetic
stimulation also constricts the ciliary muscle,
leading to accommodation for near vision.
These parasympathetic effects are functions of
cranial nerve III (oculomotor).
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.
Review Questions
1. Outline the functional anatomy of the sympathetic and parasympathetic nervous

systems.
2. Give two examples of autonomic reflexes. Describe the influence of sensory input
and of supraspinal control.
Review Questions
1. Outline the functional anatomy of the sympathetic and parasympathetic nervous

systems.
2. Give two examples of autonomic reflexes. Describe the influence of sensory input
and of supraspinal control.
3. Prepare an outline showing the organization of chemical transmission in the auto

nomic nervous system, including the sites of aceylcholine and norepinephrine
secretion, and of nicotinic, muscarinic, alpha, and beta receptors.
4. Briefly describe the effects of sympathetic and parasympathetic stimulation on

the cardiovascular system, the gastrointestinal tract, the eye, and the metabolism
of glucose.
5. Define denervation and denervation sensitivity.

11. Cerebral Physiology and
Higher Functions
How cerebral activity relates to wakefulness and sleep,
memory, language, and other complex processes______________
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
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
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
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
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
Table 11-1. Some Characteristics of Four

Types of Epilepsy
Generalized
Grand mal Loss of consciousness
Tonic and clonic muscle contractions
EEG showing repetitive spikes or spike-
and-wave complexes throughout
cortex
Probable origin in brainstem reticular
formation and/or nonspecific thalamic
nuclei
Petit mal Brief loss of consciousness
Minor motor effects
EEG showing spike-and-wave com
plexes, symmetrical throughout cor
Figure 11-6. EEG samples during an attack of tex, at about 3 per second
grand mol (A) and petit mol epilepsy (B). High-am- Probable origin in nonspecific thalamic
plitude, repeated spikes are recorded during the tonic nuclei
phase of a grand mol attack but are frequendy ob Partial
scured by EMG artifact.
Focal motor Consciousness retained
Clonic muscle jerks limited toparticular
parts of the body
EEG showing spikes in motor cortex
Probable origin in epileptic focus in
consciousness, falls down, and has convulsive motor cortex
muscle contractions, both continuous (tonic) Psychomotor Consciousness altered to variable extent
and jerky (clonic). An EEG recorded during Automatic motor activity often organ
ized into complex acts, and possible
the seizure shows high-amplitude, high- mood changes, perceptual distortions,
frequency spike potentials or sometimes spikes and sensory hallucinations
followed by slow waves, throughout the cortex EEG usually shows spikes or slow waves
in temporal lobe (but other locations
(figure 11-6A ). The probable origin is the are possible)
brainstem reticular formation and/or the non Probable origin in epileptic focus in
specific thalamic nuclei, which regulate the temporal lobe
level of consciousness and influence electrical
activity of the entire cortex, as well as (in the
case of the reticular formation) influence mus foci in these nuclei have produced a similar
cle contraction. These characteristics are sum syndrome.
marized in table 11 - 1 , along with those of
other forms of epilepsy. Partial (Focal, Local) Epilepsy:
Petit MaL This involves a brief loss of con Focal Motor. A typical seizure might start
sciousness, usually lasting less than 60 seconds, with an involuntary jerk of a finger, while at
with few or no motor signs, and can be so the same time abnormal spike discharges appear
inconspicuous as to go unnoticed. It is often in the hand area of the contralateral motor
found in children. The very characteristic EEG cortex (the epileptic focus). If the epileptic
pattern consists of spike-and-slow wave com activity spreads to adjacent areas of the motor
plexes repeated at about three per second, sym cortex, motor activity spreads along corre
metrical throughout the cortex (figure 11-6B). sponding parts of the body (e.g., from finger to
The probable origin is in the nonspecific hand to arm to the whole side of the body); this
thalamic nuclei; artificially induced epileptic phenomenon is known as Jacksonian March.
Consciousness may be retained if epileptic ments, such as buttoning and unbuttoning a

activity remains limited to the motor cortex. In jacket. An epileptic focus is generally found in
some cases, however, epileptic activity spreads the temporal lobe, so that temporal lobe
from the motor cortex to the nonspecific epilepsy sometimes refers to the same syndrome.
thalamic nuclei and/or the brainstem reticular
formation, leading to a generalized seizure with Evoked Potentials
loss o f consciousness as in grand mal epilepsy.
The characteristics listed in table 11-1 apply to Evoked potentials are voltage changes within
focal motor epilepsy that does not become the EEG that are evoked or elicited by sudden
generalized. sensory stimuli, such as electrical stimulation of
Psychomotor. The psycho- refers to bizarre the finger, a click, or a light flash. To use the
feelings, mood changes, or hallucinations; a first example, an electrical pulse applied to the
patient may experience feelings of unreality, right index finger so as to give a mild shock
feelings of anger or fear, or hallucinations, such sensation sets up nerve impulses in the median
as events from the past that seem real. The nerve that continue up the spinal cord and are
motor of psychomotor refers to automatic move- relayed from the thalamus to the left somatic
sensory cortex (postcentral gyrus). The cortical
response recorded by an EEG electrode over
Figure 11-7. A human somatosensory evoked poten
tial (SEP), recorded with a scalp electrode over the that area of the scalp is minute compared with
hand projection area of the left postcentral gyrus with the spontaneous EEG waves. However, if the
reference electrode on left ear, while right median stimulus is repeated about 100 times and the
nerve is stimulated. Evoked potential (right) repre voltage values at corresponding time intervals
sents average of 120 responses; P25, P45, and after each stimulus are averaged over those
P100 are positive peaks at 25, 45, and 100 msec
after stimulus onset, respectively. Circled area repetitions, the cortical sensory response will
around scalp electrode is area where P25 is found at summate and be enhanced, while the spon
maximal amplitude. Source: Adapted from Goff, taneous EEG waves will tend to cancel each
G.D., Matsumiya, Y., Allison, T., et aL The scale other out. (This technique is known as signal
topography of human somatosensory and auditory averaging: the sensory response is considered a
evoked potentials. EEG Clin. Neurophysiol.
42:57-76, 1977; and Buchsbaum, M.S., Lavine, signal to be extracted from a background of
R.A., Davis, G.C., et aL Effects of lithium on EEG noise).
somatosensory evoked potentials and prediction of The resulting waveform is called an averaged
clinical response in patients with affective illness. In evoked potential (figure 11-7). It may be com
Cooper, T.B., Gershon, S., Kline, N.S., and pared with evoked potentials recorded directly
Schou, M. (Eds.), International Lithium Confer
ence: Controversies and Unresolved Issues. Law- from the cortex of human or animal subjects. A
renceville, N.J.: Excerpta Medico, ICS Series, positive wave that peaks at a latency of 25 to 30
1979. msec after the electrical stimulus is thought to
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
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.
Reading and the brain. Brain imaging by

fMRI has revised the traditional model of
reading, although there is still disagreement
about details. In a revised view, the angular
gyrus is thought to be involved in detecting
visual forms more general than just letters,
while recognizing letters is attributed to a
lower, more ventral region between the
occipital and temporal lobes called
the occipito-temporal sulcus (Dehain, S.
(2009), Reading and the Brain, New York:
Penguin Books, pp. 70-71). It’s also called
the Visual Word Form Area, within
Brodmann’s Area 37.This area receives
information from the visual cortex during
reading, detects letters, and sends its output
to temporal lobe areas that detect words,
sentences, and their meaning, and
associates written syllables to their
sounds. Current research is looking at the
exact location, function, and connections of
these neural regions.
The visual word form area is adjacent to or

overlaps the face area, a group of neurons
specialized to respond to different faces.
The face area is in the fusiform gyrus and
typically more prominent in the right
hemisphere.
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
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.
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
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
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
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
to a single stimulus during a period ranging up stimulation, is retained (remembered) from

to several minutes afterward is increased. This minutes to hours. This habituation is paral
effect is called post-tetanic potentiation. It has leled by a decline in the EPSP in the
also been found in various parts of the brain, motoneuron that supplies the gill muscle. The
and is a mechanism by which changes in EPSP decline was attributed to a depletion of
neuron excitability could be retained for a short transmitter from the presynaptic terminal of
period. the sensory neuron [20 ].
Short-term memory has been experimentally In mammalian nervous systems, habituation
disrupted by various treatments that disrupt has been found for polysynaptic spinal reflexes,
neural activity. These include electrical shock such as the flexor withdrawal reflex. Both a
to the brain (electroconvulsive shock), chem decline in presynaptic transmitter release, as in
ically induced depolarization of cortical Aplysia, and a buildup in the activity of
neurons (spreading depression), and reduction inhibitory intemeurons have been suggested as
of temperature (hypothermia) and oxygen sup possible mechanisms. Single neurons within
ply (anoxia). the brain have also been found to habituate to
Consolidation of information from short repeated stimuli. Most of the neurons sampled
term to long-term memory is often attributed to within the hippocampus appear to habituate, as
more stable biochemical and structural changes do a number of neurons sampled within the
in neurons, including altered synthesis of RNA brainstem reticular formation and the cerebral
and protein, and anatomical changes in sy cortex.
napses. In animals such as fish and mice, the
drugs actinomycin D, which interferes with
RNA synthesis, and puromycin, which inter
feres with protein synthesis, appear to block a
consolidation stage without interfering with
short-term or long-term memory itself. Particu
larly interesting, in view of the role of the
hippocampus in human memory, are the
changes found in synaptic structure when a
pathway w ithin the hippocampus was
stimulated repetitively at high frequency [31].
Not only did post-tetanic potentiation occur,
but dendritic spines (small knobs on dendrites
which contain postsynaptic terminals) in
creased in size. Such a change could lead to
more effective spread of EPSPs into the postsyn
aptic cell.
One of the simplest examples of modified
behavior or learning is the decline of the
response to a regularly repeated stimulus, a
change called habituation (although there are
additional criteria for habituation [20]). In an
invertebrate (Aplysia), distinguished by its
enormous nerve cells, habituation of a simple
behavior, the withdrawal of a gill after sensory
The Mental Status Examination
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
Other topics in the Mental Status Examination include the following.
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
Review Questions
1. Describe the method of recording the EEG.
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
Define REM and NREM sleep.
Compare generalized with partial epilepsy. Briefly describe grand mal, petit mal,
focal motor, and psychomotor forms of epilepsy.
es to sensory stimulation.
7. Define aphasia and language-dominant hemisphere.
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.
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.
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
15. Define anterograde amnesia and retrograde amnesia.
16. Describe the association of amnesia with brain lesions and brain dysfunctions.
17. Give examples of possible physiological mechanisms underlying short-term and

long-term memory.
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
15. Define anterograde amnesia and retrograde amnesia.
16. Describe the association of amnesia with brain lesions and brain dysfunctions.
17. Give examples of possible physiological mechanisms underlying short-term and

long-term memory.
Glossary Anterograde amnesia Defective memory for a
period following a brain lesion or other event
Word origins in Latin (L.) or Greek (Gr.) are Aphasia (Gr. phasis, speech) A partial or complete
given only where particularly helpful for under loss of the ability to express or comprehend
standing or remembering the term. language (including speech, writing, and signs),
generally due to brain pathology; partial loss is
also called dysphasia
Acetylcholine (ACh) Chemical transmitter, syn Aqueous humor The fluid in the front part of the
thesized from acetic acid + choline, released by eye
vertebrate motoneurons, preganglionic sym Archicerebellum (Gr. arche, beginning) A primi
pathetic and parasympathetic neurons, and tive part of the cerebellum, including the
probably other neurons in the central nervous flocculonodular lobe, that helps maintain equi
system librium
Action potential Brief change in membrane poten Astigmatism (Gr. stigma, point) Refractive error
tial that is conducted along axon or muscle fiber caused by defective curvature of the refracting
without decay surface (e.g., cornea), the radius of curvature
Adaptation Diminished response of a sensory being greater in one plane than in another
neuron to a maintained stimulus perpendicular to it
Adiadochokinesis (Gr. diadochos, succeeding + Athetosis (Gr. athetos, not fixed) A movement
kinesis, motion) Inability to perform movements disorder characterized by slow snakelike, wri
that are rapid and alternating thing, involuntary movements, often of fingers
Adrenergic Pertaining to neurons that release and toes
norepinephrine (noradrenaline) as a synaptic Atropine Anticholinergic drug acting at parasym
transmitter pathetic effectors
Affect Emotion, outwardly expressed Audiogram A graph showing hearing threshold as
Afferent (L ad, to + ferre, carry) Referring to a function of frequency
axons that conduct nerve impulses toward the Averaged evoked potentials CNS response to a
central nervous system, or the impulses them sensory stimulus obtained by averaging the EEG
selves data points at corresponding time intervals
Air conduction Transmission of sound waves to following a number of stimuli, thereby cancel
the cochlea via the external auditory canal and ling those potential oscillations with no fixed
the middle ear relation to the stimulus while enhancing those
Alexia (Gr. lexis, word) Loss of reading ability time-locked to the stimulus
Alpha blocking Disappearance of alpha waves after Axon Process or extension of a neuron, capable of
eyes are opened or other sensory stimulus is conducting nerve impulses to other cells; a
presented nerve fiber
Alpha motoneurons Rapidly-conducting my Axon hillock Raised cone-shaped area of the cell
elinated neurons innervating the extrafusal fi body at which the axon originates
bers of skeletal muscles Babinski’s sign Upgoing of the big toe in response
Alpha receptor A type of adrenergic receptor to stroking sole of foot, generally after a lesion
Alpha waves EEG waves between 8 and 13 Hz of descending motor pathways
occurring when the subject is relaxed with eyes Ballismus (Gr. ballismos, a jumping about) A
closed movement disorder characterized by violent,
Amacrine cells One of the five types of retinal cell flinging, involuntary movements, associated
Amnesia Loss of memory, of varying extent with lesions of the subthalamic nucleus
Amplitude Height of a wave or signal; a measure of Basilar membrane Membrane extending longitu
its magnitude dinally through cochlea, separating the scala
Analgesic (Gr. angesis, pain) Pain-relieving media and the scala tympani and upon which
Angular gyrus Part of the parietal lobe association the organ of Corti rests; forms pan of the
cortex, important in the comprehension of cochlear partition
written words Best frequency Determined from a tuning curve;
Annulospiral nerve ending Receptor that signals the frequency at which a particular neuron in
stretch of a muscle; also termed primary stretch the auditory system has the lowest threshold
receptor Beta receptors A type of adrenergic receptor
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
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Mosby Co., 1977.
QUIZ YOURSELF B. 50 m/sec
C. 30 m/sec
PARTI D. 15 m/sec
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
A. Disappears after infancy

B. Depends on Golgi Tendon Organ
C. Hyperactive in upper motor neuron lesions
D. Associated with crossed extension response
Which o f the following parts of the body has the

largest representation on the motor cortex (area 4)?
A. face
B. lower leg
C. upper back
D. lower back
A lesion o f extracorticospinal tracts in the left

hemisphere can lead to which o f the following?
A. Increased tendon jerk reflexes with normal
muscle tone in the right left.
B. Increased tendon jerk reflexes and muscle tone in
the right leg.
C. Decreased tendon jerk reflexes with normal
muscle tone in the right leg.
D. Decreased tendon jerk reflexes and muscle tone
in the right leg.
E. Increased tendon jerk reflexes and muscle tone in
the left leg.
T F A drug which blocks dopamine receptors in

the basal ganglion can produce parkinsonian
symptoms.
Which o f the following is true?

A. the best stimulus for annulospiral receptors is
increased tension
B. lower motor neuron lesions are associated with
clonus
C. gamma efferent stimulation increases the
sensitivity of the muscle stretch reflex
D. muscle stretch reflex afferent fibers belong to
Group II
In the brain-stem auditory evoked response, Wave I
QUIZ YOURSELF
at about 1.5 msec after a click stimulus probably
PART II represents:
A. auditory nerve activity
The following questions are in several formats. B. REM sleep
C ircle T or F for True-False questions, fill in the C. auditory cortex activity
blanks where indicated, and circle the letter before D. a spike response in grand mal epilepsy
the one best answer for multiple-choice questions.
An absence of both air conduction and bone
T F Each taste receptor cell responds to only one conduction on one side is consistent with:
type of taste stimulus. A. a puncture of the eardrum
B. middle ear deafness
Which cranial nerve carries sensory information C. sensorineural deafness
about sweet substances on the tip of D. wax in the outer ear
the tongue?
A. nerve V The function of the semicircular canals is:
B. nerve VII A. detection o f high-frequency sounds
C. nerve IX B. reducing transmission to the cochlea
D. nerve XII C. detection o f rotation
D. detection o f straight-line acceleration
T F Inability to hear via bone conduction E. amplification o f sound waves
indicates middle ear deafness.
The difference in area between the tympanic
T F One o f the functions o f the middle ear is to membrane and the footplate of the stapes in the oval
equilibrate the air pressure across the tympanic window is responsible for:
membrane by means of the eustachian tube. A. reducing transmission o f loud sounds to the
cochlea
The function o f the middle ear reflex is: B. a small part o f the pressure amplification across
A. protection of the cochlea from sudden and brief the middle ear
loud sounds C. most of the pressure amplification across die
B. protection of the cochlea from sustained loud middle ear
sounds D. equilibration o f air pressure across the tympanic
C. amplification o f high frequency tones membrane
D. amplification o f low frequency tones
T F In accommodation, the suspensory
Physiological nystagmus during rotation to the ligaments allow the lens to become more convex.
person's right:
A. involves slow eye movements to the right. T F The greatest visual acuity is at the periphery
B. requires sensory input via the optic nerve. o f the visual field.
C. is due to a lesion in the vestibular organ or nerve.
D. can appear similar to the caloric response if the T F Cones function to provide black and white
patient's head is positioned appropriately. vision in dim light.
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
A spike and slow-wave pattern in the EEG at 3 per

second is characteristic of which form o f epilepsy?
A. petit mal
B. grand mal
C. focal motor
D. psychomotor
Motor seizures limited to the left hand with no loss of

consciousness are often due to lesions of which
cortical area?
A. Wernicke's area on the left hemisphere
B. Striate cortex on die right hemisphere
C. Motor cortex on the right hemisphere
D. Broca's area on the left hemisphere
A loss in visuospatial ability, as tested by copying a

complex figure or constructing a design in blocks, is
often due to damage in:
A. primary visual cortex
B. parietal association cortex
C. temporal association cortex
D. prefrontal cortex
Copyright Acknowledgments
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.
Cover art adapted from iStock.(c), iStock _ 000073913101
“Neurotranmitters” © CC BY-SA 3.0
NIMH "Brain Basics” www.nimh.nih.gov. downloaded September 2016

Supplementary Updates
Brain Basics. This overview from the National Institutes of Mental Health describes
important brain regions and processes with an emphasis on neuroscience applied to
mental health. It goes on to describe methods of brain imaging such as magnetic
resonance imaging (MRI) and provides a list of information resources.
Major Neurotransmitters Overview. This brief overview describes several of the

major neurotransmitters and their actions.
\Z * * 7
I V I I I I I National Institute of Mental Health
Brain Basics
Introduction
Introduction
W elcom e. Brain Basics provides inform ation on how the Brain Basics will introduce you to some o f this science,
brain w orks, how mental illnesses are disorders o f the brain, such as:
and ongoing research that helps us better understand and
Tire Growing Brain treat disorders. • How the brain develops
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.
Through research, we know that mental disorders are brain

disorders. Evidence shows that they can be related to changes
Glossary in the anatomy, physiology, and chem istry o f the nervous
system. When the brain cannot effectively coordinate the billions
of cells in the body, the results can affect many aspects of life.
Links Scientists are continually learning more about how the

brain grows and w orks in healthy people, and how normal
brain developm ent and function can go awry, leading to
mental illnesses. 4
More Information
N EX T P A G E > 1
\Z * * 7
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.
Scientists have already begun to chart how the brain develops

Links over tim e in healthy people and are w orking to com pare that
w ith brain developm ent in people mental disorders. Genes and
environmental cues both help to direct this growth.
More Information
< P R E V IO U S P A G E N EX T P A G E > 2
\Z * * 7
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.
As we grow, we create new cells, each with a copy of our

original se t o f DNA. S om etim es th is copying process is
Glossary imperfect, leading to a gene mutation that causes the gene
to code for a slightly different protein. Some mutations are
harmless, some can be helpful, and others give rise to
disabilities or diseases.
Links
G enes a ren ’t the only dete rm in a nts of how our bodies
function. Throughout our lives, our genes can be affected by
the environment. In medicine, the term environment includes
not only our physical surroundings but also factors that can
More Information affect our bodies, such as sleep, diet, or stress. These factors
may act alone or to g e th er in com plex w ays, to change the
way a gene is expressed or the way m essages are conducted
in the body.
\Z * * 7
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
r brain and the rest o f the nervous system . W hen a neuron is

activated a sm all difference in electrical charge occurs. This
unbalanced charge is called an action potential and is caused by
• Dopamine— mainly
involved in controlling
m ovem ent and aiding the
Tire Working Brain the concentration o f ions (atoms or molecules with unbalanced flow o f information to the
charges) across the cell membrane. The action potential travels front o f the brain, which
very quickly along the axon, like when a line of dom inoes falls. is linked to thought and
emotion. It is also linked to reward system s in the brain.
Brain Basics in Real Life Problem s in producing dopam ine can result in Parkinson’s
When the action potential reaches the end o f an axon, most
neurons release a chemical message (a neurotransmitter) which disease, a disorder that affects a person’s ability to move as
crosses the synapse and binds to receptors on the receiving they w ant to, resulting in stiffness, trem ors or shaking, and
neuron’s dendrites and starts the process over again. A t the end other sym ptom s. Some studies suggest that having too little
Brain Research of the line, a neurotransm itter m ay stim ulate a different kind of dopam ine or problem s using dopam ine in the thinking and
cell (like a gland cell), or may trigger a new chain o f messages. feeling regions o f the brain may play a role in disorders
like s c h iz o p h re n ia or a tte n tio n d e fic it h y p e ra c tiv ity
Neurotransmitters send chemical m essages between neurons. d is o rd e r (ADHD).
Glossary Mental illnesses, such as depression, can occur w hen this
• Glutamate— the most
process does not w ork correctly. C om m unication between
common neurotransmitter,
neurons can also be electrical, such as in areas of the brain that
glutamate has many
control movement. When electrical signals are abnormal, they
roles throughout the
Links can cause trem ors or symptom s found in Parkinson’s disease.
brain and nervous
system. Glutam ate is an
• Serotonin— helps control
excitatory transm itter: when it is released it increases the
many functions, such
chance that the neuron will fire. This enhances the electrical
as mood, appetite,
More Information and sleep. Research
flow am ong brain cells required for normal function and
plays an im portant role during early brain development.
shows that people
It may also assist in learning and memory. Problem s in
with d e p re ssio n often
making or using glutam ate have been linked to many
have lower than normal levels o f serotonin. The types of
mental disorders, including a u tis m , o b s e s s iv e c o m p u ls iv e
m edications most com m only prescribed to treat depression
d is o rd e r (OCD), s c h iz o p h re n ia , and d e p re s s io n .
act by blocking the recycling, or reuptake, of serotonin by
the sending neuron. As a result, more serotonin stays in the
< P R E V IO U S P A G E N EXT PA G E > 4

a
National Institute of Mental Health
Brain Basics
Introduction
The Working Brain
B ra in R eg io ns Anterior cingulate cortex (ACC)— the AC C has many
different roles, from controlling blood pressure and heart
Just as many neurons w orking together form a circuit, many rate to responding when we sense a mistake, helping us feel
Tire Growing Brain circuits working together form specialized brain systems. We motivated and stay focused on a task, and m anaging proper
have many specialized brain system s that w ork across specific emotional reactions. Reduced ACC activity or dam age to
r brain regions to help us talk, help us make sense of w hat we

see, and help us to solve a problem. Some of the regions most
this brain area has been linked to disorders such as ADHD,
schizophrenia, and depression.
Tire Working Brain com monly studied in mental health research are listed below.
■ Hippocampus— Helps create and file new memories. When
• A m ygdala— The brain’s “fe a r hub,” w hich activates our the hippocam pus is dam aged, a person can't create new
natural "fight-or-flight” response to confront or escape from memories, but can still rem em ber past events and learned
skills, and carry on a conversation, all which rely on different
Brain Basics in Real Life a dangerous situation. The am ygdala also appears to be
parts of the brain. The hippocam pus may be involved in
involved in learning to fe a r an event, such as touching a
hot stove, and learning not to fear, such as overcom ing a mood disorders through its control of a m ajor mood circuit
fe a r of spiders. Studying how the am ygdala helps create called the hypothalam ic-pituitary-adrenal (HPA) axis.
m em ories of fe a r and safety m ay help im prove treatm ents
Brain Research fo r a n x ie t y d i s o r d e r s like p h o b i a s or p o s t - t r a u m a t ic
s t r e s s d i s o r d e r (P T S D ).
Prefrontal Cortex
H ippocam pus
• Prefrontal cortex (PFC)— Seat of the brain’s executive \ A /'

Glossary functions, such as judgm ent, decision making, and problem /
solving. Different parts o f the PFC are involved in using
short-term or “working" mem ory and in retrieving long-term
_ — i. H H a
memories. This area of the brain also helps to control the

J
amygdala during stressful events. Some research shows
Links that people w ho have PTSD or ADHD have reduced activity
in their PFCs. ■
A n te r io r '
C in g u la te C o r te x
A m ygdala
More Information
\ T I 1 I f National Institute of Mental Health
™ i\ Brain, Basics
Glossary
Introduction
action potential— Transmission of signal DNA— The “recipe of life,” containing mutation— A change in the code for
from the cell body to the synaptic terminal inherited genetic information that helps to a gene, which may be harmless or
at the end of the cell’s axon. When the define physical and some behavioral traits. even helpful, but sometimes give rise
The Growing Brain action potential reaches the end of the
epigenetics— The study of how
to disabilities or diseases.
axon the neuron releases chemical
(neurotransmitters) or electrical signals. environmental factors like diet, stress neural circuit—A network of neurons
and post-natal care can change gene and their interconnections.
amygdala— The brain’s “fear hub,” expression (when genes turn on or
Hie Working Brain which helps activate the fight-or-flight off)—without altering DNA sequence. neuron—A nerve cell that is the
basic, working unit of the brain and
response and is also involved in emotions
and memory. gene—A segment of DNA that codes nervous system, which processes
to make proteins and other important and transmits information.
anterior cingulate cortex— Is involved body chemicals.
Brain Basics in Real Life in attention, emotional responses, and neurotransmitter—A chemical produced
many other functions. glutamate—The most common by neurons that carries messages to
neurotransmitter in a person’s body, which other neurons.
axon—The long, fiber-like part of a neuron increases neuronal activity, is involved in
by which the cell sends information to early brain development, and may also nucleus—A structure within a cell that
Brain Research receiving neurons. assist in learning and memory. contains DNA and information the cell
needs for growing, staying alive, and
cell body— Contains the nucleus and hippocampus— A portion of the making new neurons.
cytoplasm of a cell. brain involved in creating and filing
new memories. prefrontal cortex—A highly developed
cell membrane— The boundary area at the front of the brain that, in
separating the inside contents of a cell Hypothaimic-pituitary-adrenal (HPA) humans, plays a role in executive functions
from its surrounding environment. axis— A brain-body circuit which plays such as judgment, decision making and
a critical role in the body’s response problem solving, as well as emotional
cytoplasm— The substance filling a cell, to stress. control and memory.
Links containing all the chemicals and parts
needed for the cell to work properly. impulse—An electrical communication serotonin—A neurotransmitter that
signal sent between neurons by which regulates many functions, including
dendrite— The point of contact for neurons communicate with each other. mood, appetite, and sleep.
More Information receiving impulses on a neuron,
branching off from the cell body. magnetic resonance imaging (MR!)— synapse—The tiny gap between neurons,
An imaging technique that uses where nerve impulses are sent from one
dopamine—A neurotransmitter mainly magnetic fields to take pictures of neuron to another.
involved in controlling movement, the brain’s structure.
managing the release of various
hormones, and aiding the flow of
information to the front of the brain.
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
The Growing Brain NIMH C lin ic a l T ria ls

Clinical trials funded by NIMH
M e d lin e P lu s
Hie Working Brain C onsum er health information provided by the National Library of Medicine
NIH O ffice o f S cien ce E d u ca tio n

Science education resources developed or supported by the National Institutes o f Health (NIH)
Brain Basics in Real Life
NIH R e P O R T E R
Public database o f active grants funded by the National Institutes of Health
Brain Research
Glossary
Links
More Information
< PREVIO U S PAGE NEXT PAGE > 9

\ T I 1 I f National Institute of Mental Health
™ i\ Brain, Basics
Brain Research
Introduction
Modern research tools and techniques are giving scientists G e n e S tu d ie s
a more detailed understanding o f the brain than ever before.
Advanced technologies are also making it faster, easier, and
The Growing Brain B r a in Im a g in g m ore affordable to study genes. Scientists have found many
different genes and groups o f genes that appear to increase
Using brain imaging technologies such as m agnetic resonance risk or provide protection from various mental disorders. Other
imaging (MRI), which uses m agnetic fields to take pictures of genes may change the w ay a person responds to a certain
Hie Working Brain the brain’s structure, studies show that brain growth in children medication. This inform ation may som eday make it possible
with a u tis m appears to peak early. And as they grow there to predict w ho will develop a mental disorder and to tailor the
are differences in brain developm ent in children w ho develop treatm ent for a person’s specific conditions.
bipolar disorder than children w ho do not.
Brain Basics in Real Life Such brain research help increase the understanding of
Studies comparing such children to those with normal brain how the brain grows and w orks and the effects o f genes and
developm ent may help scientists to pinpoint when and where environm ent on mental health. This know ledge is allowing
1 mental disorders begin and perhaps how to slow or stop them scientists to make im portant discoveries that could change the
from progressing. Functional m agnetic resonance imaging w ay we think about and treat mental illnesses.
B rain R esearch (fM RI) is another im portant research tool in understanding how
the brain functions. The National Institute o f Mental Health supports m any studies
on mental health and the brain. You can read about som e of
Another type of brain scan called magnetoencephalography, these studies online at w w w .n im h .n ih .g o v .
Glossary o r MEG, can capture split-second changes in the brain. Using
MEG, som e scientists have found a specific pattern o f brain
activity that may help predict w ho is most likely to respond to
fast-acting antidepressant medications. C urrently available
Links antidepressants usually take four to six w eeks to reach th e ir full
effect, which can be a difficult w ait fo r some people struggling
with depression. However, recent research points to a possible
new class o f antidepressants that can relieve sym ptom s o f the
illness in just a few hours. Knowing who m ight respond to such
More Information m edications could reduce the am ount of trial and error and
frustration that many people with depression experience when
starting treatment.
Mil National Institute of Mental Health
Brain Basics
More Information
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
Phone: 301 - 443-4513 or

Tire Working Brain 1 - 866-61 5 - n im h (6464) toll-free
& TTY: 301 - 443-8431 or
-------------------------------------------------------------------------------------- 1 - 866 - 415-8051 toll-free
FAX: 301 - 443-4279
Brain Basics in Real life
Email: n im h in fo @ n ih .g o v
W eb site: h ttp ://w w w .n im h .n ih .g o v
Brain Research
Glossary
Links
More Information
< PREVIO U S PAGE 10

Neurotransmitters Overview
Neurotransmitter molecules include amino acids, amines, and peptides. Major examples
are these. Amino Acids: GABA, Glutamate, Glycine, and Aspartate. Amines:
Achetylcholine, (ACh), Dopamine (DA), Norepinephrine (NE), Epinephrine, Histamine,
and Serotonin (5-HT). Peptides: Hypothalamic Neuropeptides Oxytocin and
Vasopressin, Substance P, and Beta-Endorphin.
Neurotransmitters are synthesized in nerve cell bodies or presynaptic terminals, stored

in presynaptic terminals, released from these terminals upon stimulation, and have
specific receptors on the postsynaptic cells.
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).
Glutamate. Glutamate is the major excitatory amino acid neurotransmitter in the

central nervous system. It binds to several types of postsynaptic receptors, which allow
Na+ and sometimes Ca++ to enter the cell and cause a depolarization that excites the
neuron.
GABA. Gamma-amino butyric acid, or GABA, is the major inhibitory

neurotransmitter in the CNS. Like glutamate, it is an amino acid neurotransmitter, and
has different types of postsynaptic receptors. The receptors change gated chloride (Cl-)
channels, or potassium (K+) and calcium (Ca2+) channels. These synaptic receptors are
sites where several drugs act to control neural excitability, anxiety, and sleep.
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.
Norepinephrine. In the brain, norepinephrine is a neurotransmitter that affects

alertness and mood, and responds during anxiety or panic. The locus coeruleus in the
brain stem sends nerve fibers that branch out to secrete norepinephine in widespread
areas of brain. Peripherally, in the sympathetic nervous system, it is released by
postganglionic nerve fibers to act on end organs like the heart and blood vessels. For
example, it can increase the heart rate and constrict blood vessels. Norepinephrine is
also called noradrenaline.
Epinephrine and norepinephrine are closely related and both can be secreted
during emergencies or anxiety episodes. Epinephrine can also be used as a drug to
control severe allergic reactions or other conditions. Epinephrine is also called
adrenaline.
Dopamine. Related chemically to norepinephrine and epinephrine, dopamine

helps control movement and posture. The loss of dopamine in the basal ganglia causes
the muscle rigidity typical of Parkinson’s Disease. The dopamine precursor L-DOPA can
be used in treatment, Dopamine also modulates mood, plays a role in positive
reinforcement and reward, and contributes to the dependency found with addictive
drugs. Antipsychotic drugs act typically as dopamine antagonists.
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.
Hypothalamic Neuropeptides. The hypothalamus, centrally located in

the brain, controls the secretion of neuropeptides from the pituitary gland, an
outcropping extending below the hypothalamus. Oxytocin, composed of amino acids,
controls milk ejection and plays a role in bonding between mother and infant and
possibly adults as well. Vasopressin, also composed of amino acids, is released from
the posterior pituitary to control kidney function and regulate blood volume and
pressure. Both neuropeptides are thought to affect neurons in the brain as well as their
peripheral actions.
Supstance P. This large peptide containing 12 amino acids is released by nerve

fibers in the spinal cord when they are triggered by painful stimuli acting on
nociceptors. This release is combined with that of the excitatory neurotransmitter
glutamate, activating the spinothalamic pain pathway. Opiate drugs are thoughts to
have an analgesic effect by reducing this activation. Substance P has effects in the brain
and other tissues as well.
Endorphin. A peptide involved in reducing pain and responding to pleasure, it is

structurally similar to opium and the opiate drugs (morphine, heroin, etc.), which can
attach to the endorphin receptor site. The name is a combination of “endogenous”
(produced by the body) and morphine. There are several types, and a well –known one
is beta-endorphin.
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Neuroscience ABCs

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Neuroscience ABCs

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Neuroscience

Robert Lavine, Ph.D. Robert Lavine, Ph.D.

Neuro Cognitive Publications

The book can be read alone or along

The goal is also to provide a foundation

Since neuroscience is complex, you may

Research findings and clinical exam

1. Structure and Function

2. The Nerve Impulse How nerve cells become excited and 25

3. Sensory Reception How receptors convert energy from the 47

4. Synaptic Transmission How signals pass from neuron to neuron 59

5. Somatic Sensation How the nervous system converts touch, 85

6. Auditory and How neural mechanisms function in 105

8. Chemical Senses How the nervous system responds to 157

10. Autonomic Function How sympathetic and parasympathetic 197

11. Cerebral Physiology and How cerebral activity relates to 207

How the nervous system is designed to communicate

put). There are then four types of peripheral

Ventral horn (Gray matter)

Table 1-1. Cranial Nerves and Their Major Functions

VIII Auditory Hearing

at the midline into a left and right half and

lateral ascending ramus

frontal eye som esthetic

This fMRI image show a side view of the inside

Adapted from: Cantlon JF, Brannon EM, Carter EJ,

Chemical Transmitter Release Velocity

Giant Axons from the

The giant squid can reach 40 feet long

The membrane has two layers of lipid or

Adapted from Depolarizing Pre-Pulse,

and vibrations set up by the predator at a

Conduction Some receptors in the somatic sensory system

EXTERNAL RECEPTOR SENSORY

1_3)‘ The electrical potential change within muscle

You willprobablyfind that while most adults

2. Describe the role of receptors and effectors.

4. Compare the white and gray areas of the CNS.

5. List in outline form the major structures within the brain.

8. Define transduction in a sensory receptor.

9. Give an example o f how a spinal motoneuron might integrate information from

How nerve cells become excited and conduct impulses

Figure 2-2. An unmyelinated axon (left) and class­

If the membrane were permeable only to

Table 2-1. Common Electrical Symbols

Figure 2-8. Membrane potential measured in a large The Action Potential

Figure 2-10. An action potential, originating with a Na+( 15 mM

10-12 mole/cm2 of membrane). Although

potassium conductance brings the mem­ msec

brane closer to Er (-9 0 mV) than during

10-12 mole/cm2 of membrane). Although

potassium conductance brings the mem­ msec

brane closer to Er (-9 0 mV) than during

few milliseconds have passed (the sodium con­

Conduction o f the Action Potential

At some segment of axon membrane, depolari­

Figure 2-15. Conduction velocity of myelinated ax­

Figure 2-16. Types of action potential recordings.

Review Giant Axons to

The giant squid can reach 40 feet long

The membrane has two layers of lipid or

Adapted from Depolarizing Pre-Pulse,

No, the sodium, channels are not just little

The side-view above shows part of a

Figure 2-2. An unmyelinated axon (left) and class

potassium conductance brings the mem msec

potassium conductance brings the mem msec

few milliseconds have passed (the sodium con

At some segment of axon membrane, depolari

Figure 2-15. Conduction velocity of myelinated ax

anism is not known definitely, the mechan

movement of its protein components, open

transmitters are acetylcholine and norepineph

direct change of postsynaptic membrane con Acetylcholine (ACh)

Figure 4-15. Major chemical transmitters and pre

1. Diagram a neuromuscular junction, labeling the presynaptic terminal, synaptic vesi