2

Nerve Cells, Neural Circuitry, and Behavior

The Nervous System Has Two Classes of Cells Individual nerve cells or neurons are the basic units
Nerve Cells Are the Signaling Units of the of the brain. The human brain contains a huge number
Nervous System of these cells, on the order of 1011 neurons, that can be
Glial Cells Support Nerve Cells classified into at least a thousand different types.
Yet the complexity of human behavior depends less
Each Nerve Cell Is Part of a Circuit That Has One or More
Specific Behavioral Functions on the variety of neurons than on their organization
into anatomical circuits with precise functions. One
Signaling Is Organized in the Same Way in All Nerve Cells
key organizational principle of the brain, therefore, is
The Input Component Produces Graded Local Signals that nerve cells with similar properties can produce
The Trigger Zone Makes the Decision to Generate an different actions because of the way they are
Action Potential interconnected.
The Conductive Component Propagates an All-or-None Because relatively few principles of organization
Action Potential give rise to considerable complexity, it is possible to
The Output Component Releases Neurotransmitter learn a great deal about how the nervous system pro-
duces behavior by focusing on five basic features of
The Transformation of the Neural Signal from Sensory to
Motor Is Illustrated by the Stretch-Reflex Pathway the nervous system:
Nerve Cells Differ Most at the Molecular Level
1. The structural components of individual nerve cells;
Neural Network Models Simulate the Brain’s Parallel 2. The mechanisms by which neurons produce
Processing of Information signals within and between nerve cells;
Neural Connections Can Be Modified by Experience 3. The patterns of connections between nerve cells
and between nerve cells and their targets: muscles
he remarkable range of human behavior and gland effectors;

T depends on a sophisticated array of sensory

receptors connected to a highly flexible neural
organ—the brain—that selects from among the stream
4. The relationship of different patterns of intercon-
nection to different types of behavior; and
5. How neurons and their connections are
modified by experience
of sensory signals those events in the environment that
are important to the individual. In other words, the brain
actively organizes perception, some of which is stored In this chapter we provide an overview of the
in memory for future reference, and some of which is neural control of behavior by introducing these topics
transformed into immediate behavioral responses. All together. We first consider the structure and function of
this is accomplished by interconnected nerve cells. neurons and the glial cells that surround and support
them. We then examine how individual cells
organize and transmit signals and how signaling

1
 Part I / Overall Perspective

between a few interconnected nerve cells produces a

simple behavior, the knee-jerk reflex. Finally, we con-
sider how changes in signaling by specific cells
can modify behavior.

The Nervous System Has Two Classes of Cells

Apical dendrites
There are two main classes of cells in the nervous
system: nerve cells, or neurons, and glial cells, or glia.

Nerve Cells Are the Signaling Units of the

Nervous System
A typical neuron has four morphologically
defined regions: (1) the cell body, (2) dendrites, (3)
axon, and
(4) presynaptic terminals (Figure 2–1).Each region Cell body
has a distinct role in generating signals and
communicating with other nerve cells. Nucleus Basal
dendrites
The cell body or soma is the metabolic center of the
cell.
The cell body usually gives rise to two kinds
Axon
of processes: several short dendrites and one long, tubu- hillock
Axon
lar axon. Dendrites branch out in tree-like fashion and (initial
are the main apparatus for receiving incoming signals segment)

from other nerve cells. The axon typically extends

some distance from the cell body and carries signals Node of Ranvier
to other neurons. An axon can convey electrical signals
over distances ranging from 0.1 mm to 2 m. These elec- Myelin sheath
trical signals, called action potentials, are initiated at a
specialized trigger region near the origin of the axon
Axon

Presynaptic
cell

Figure 2–1 The structure of a neuron. Most neurons in

the vertebrate nervous system have several main features in
common. The cell body contains the nucleus, the storehouse
of genetic information, and gives rise to two types of cell Presynaptic
terminal
processes: axons and dendrites. Axons are the transmitting Synapse
element of neurons; they vary greatly in length. Many axons Postsynaptic
dendrite
are insulated by a sheath of fatty myelin
that is regularly interrupted at gaps called the nodes of Ranvier. Postsynaptic
The action potential, the cell’s conducting signal, is initiated at cells
the initial segment of the axon and propagates to the synapse,
the site at which signals fow from one neuron to another.
Branches of the axon of the presynaptic neuron transmit signals
to the postsynaptic cell. The branches of a single axon may form
synapses with as many as 1,000 postsynaptic neurons. The api-
cal and basal dendrites together with the cell body are the input
elements of the neuron, receiving signals from other neurons.

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 Chapter 2 / Nerve Cells, Neural Circuitry, and Behavior

0 potential is propagated along the entire length of the

called the initial segment from which they propagate axon to its terminals. In most neurons studied to date
down the axon without failure or distortion at speeds electrical signals in fact travel in one direction. Later in
of 1 to 100 m/s. The amplitude of an action potential this chapter we describe the physiological basis of this
traveling down the axon remains constant at 100 mV principle.
because the action potential is an all-or-none impulse The other principle is connectional specificity,
that is regenerated at regular intervals along the which states that nerve cells do not connect randomly
axon. with one another in the formation of networks. Rather
Action potentials are the signals by which the brain each cell makes specific connections—at particular
receives, analyzes, and conveys information. These contact points—with certain postsynaptic target cells
signals are highly stereotyped throughout the nervous but not with others. The principles of dynamic
system, even though they are initiated by a great vari- polarization and connectional specificity are the
ety of events in the environment that impinge on our basis of the modern connectionist approach to
bodies—from light to mechanical contact, from odor- studying the brain.
ants to pressure waves. The signals that convey infor- Nerve cells are classified into three major
mation about vision are identical to those that carry functional categories: sensory neurons, motor neurons,
information about odors. Here we see a key princi- and interneurons. Sensory neurons carry information
ple of brain function: the information conveyed by from the body’s peripheral sensors into the nervous
an action potential is determined not by the form of system for the purpose of both perception and motor
the signal but by the pathway the signal travels in the coordination. Some primary sensory neurons are
brain. The brain analyzes and interprets patterns of called afferent neurons, and the two terms are used
incoming electrical signals and their pathways, and interchangeably. The term afferent (carried toward the
in turn creates our sensations of sight, touch, smell, central nervous system) applies to all information
and sound. reaching the central nervous system from the
To increase the speed by which action potentials periphery, whether or not this information leads to
are conducted, large axons are wrapped in an insu- sensation. The term sensory should, strictly speaking,
lating sheath of a lipid substance, myelin. The sheath be applied only to afferent inputs that lead to
is interrupted at regular intervals by the nodes of perception. Motor neurons carry commands from the
Ranvier, uninsulated spots on the axon where the brain or spinal cord to muscles and glands (efferent
action potential is regenerated. information). Interneurons are the most numerous and
Near its end the axon divides into fine branches are subdivided into two classes: relay and local. Relay
that contact other neurons at specialized zones of or projection interneurons have long axons and
communication known as synapses. The nerve cell convey signals over consider- able distances, from
trans- mitting a signal is called the presynaptic cell; one brain region to another. Local interneurons have
the cell receiving the signal is the postsynaptic cell. short axons because they form connections with
The presynaptic cell transmits signals from nearby neurons in local circuits.
specialized enlarged regions of its axon’s branches,
called presynaptic terminals or nerve terminals. The Glial Cells Support Nerve Cells.
presynaptic and postsynaptic cells are separated by Glial cells greatly outnumber neurons—there are 2 to
a very narrow space, the synaptic cleft. Most 10 times more glia than neurons in the vertebrate
presynaptic terminals end on the postsynaptic central nervous system. The name for these cells
neuron’s dendrites; but the terminals may also derives
terminate on the cell body or, less often, at the
beginning or end of the axon of the receiving cell
(see Figure 2–1).
By examining the structure of neurons in almost
every region of the nervous system, he could describe
classes of nerve cells and map the precise connections
between many of them.
Two principles of neural organization are
particularly valuable in studying communication in
the nervous system. The first of these has come to
be known as the principle of dynamic polarization. It
states that electrical signals within a nerve cell flow
only in one direction: from the receiving sites of the
neuron, usually the dendrites and cell body, to the
trigger region at the axon. From there the action 3
 Chapter 2 / Nerve Cells, Neural Circuitry, and Behavior

Three types of multipolar cells

Dendrites

Apical
dendrite
Cell body

Cell
body

Basal
dendrite

Dendrites Cell body

Axon Axon
Axon

Motor neuron of spinal cord Pyramidal cell of hippocampus Purkinje cell of cerebellum
roughly triangular cell body; dendrites emerge from both the
Figure 2–3 Multipolar cells have a single axon and many apex (the apical dendrite) and the base (the basal dendrites).
dendrites. They are the most common type of neuron in the Pyramidal cells are found in the hippocampus and throughout
mammalian nervous system. Three examples illustrate the the cerebral cortex. Purkinje cells of the cerebellum are
large diversity of these cells. Spinal motor neurons characterized by a rich and extensive dendritic tree that
innervate skeletal muscle fibers. Pyramidal cells have a accommodates an enormous synaptic input.

from the Greek for glue, but glia do not commonly hold Astrocytes, the third main class of glial cells, owe
nerve cells together. Rather, they surround the cell bod- their name to their irregular, roughly star-shaped cell
ies, axons, and dendrites of neurons. Glia differ from bodies and large numbers of processes (Figure 2–5C).
neurons morphologically; they do not form dendrites Astrocytes have end-feet, dilatations that contact and
and axons. Glia also differ functionally. They do not surround capillaries and arterioles throughout the
have the same membrane properties as neurons; brain (Figure 2–5C). The sheet-like processes of
are not electrically excitable; and are not directly protoplasmic astrocytes envelop nerve cell bodies and
involved in electrical signaling, which is the function synapses, whereas the end-feet of fibrous astrocytes
of nerve cells. contact axons at the nodes of Ranvier.
Glia in the vertebrate nervous system can be divided
The functions of astrocytes are still mysterious. It is
into two major classes: microglia and macroglia.
generally thought that astrocytes are not essential for
Microglia are immune system cells that are mobilized
information processing but support neurons in four
to present antigens and become phagocytes during
ways:
injury, infection, or degenerative diseases. There are
three main types of macroglia: oligodendrocytes,
1. Astrocytes separate cells, thereby insulating neu-
Schwann cells, and astrocytes.
ronal groups and synaptic connections from each
Oligodendrocytes and Schwann cells are small cells
other.
with relatively few processes. Both cells form the myelin
2. Because astrocytes are highly permeable to K+,
sheath that insulates an axon by tightly winding their
they help regulate the K+ concentration in the
membranous processes around the axon in a spiral.
space between neurons. As we shall learn below,
Oligodendrocytes are found in the central nervous K+ flows out of neurons when they fire. Repeti-
sys- tem; each cell envelops from one to 30 axonal tive firing may create excess extracellular K+ that
segments (called internodes), depending on axon could interfere with signaling between cells in the
diameter (Figure 2–5A). Schwann cells occur in the vicinity. Astrocytes can take up the excess K+ and
peripheral nervous system, where each envelops a thus maintain the efficiency of signaling between
single seg- ment of one axon (Figure 2–5B). Upon neurons.
myelination, oligodendrocytes and Schwann cells 3. Astrocytes perform other important housekeeping
influence axons by enhancing signal conduction chores that promote efficient signaling between
and by segregating voltage-sensitive ion channels neurons. For example, as we shall learn later, they
into distinct axonal domains (called node of take up neurotransmitters from synaptic zones after
Ranvier). release.
4. Astrocytes help nourish surrounding neurons by
releasing growth factors. 4
 Chapter 2 / Nerve Cells, Neural Circuitry, and Behavior

A Oligodendrocyte B Schwann cell C Astrocyte

Capillary
End-foot

Layers
of myelin Fibrous
astrocyte

Schwann End-foot
Nodes of cell
Ranvier
Nucleus

Inner
tongue Axon
Neuron
Nodes
of Ranvier

Figure 2–5 The principal types of glial cells are oli- Each cell forms a myelin sheath approximately 1 mm long
godendrocytes and astrocytes in the central nervous sys- between two nodes of Ranvier. The sheath is formed as the
tem and Schwann cells in the peripheral nervous system. inner tongue of the Schwann cell turns around the axon several
B. Oligodendrocytes are small cells with relatively few times, wrapping the axon in layers of membrane. In actuality
processes. In the white matter of the brain, as shown here, the layers of myelin are more compact than what is shown
they provide the myelin sheaths that insulate axons. A single here.
oligodendrocyte can wrap its membranous processes around D. Astrocytes, a major class of glial cells in the central nervous
many axons. In the gray matter perineural oligodendrocytes system, are characterized by their star-like shape and the broad
surround and support the cell bodies of neurons. end-feet on their processes. Because these end-feet put the astro-
C. Schwann cells furnish the myelin sheaths for axons in cyte into contact with both capillaries and neurons, astrocytes are
the peripheral nervous system. During development several thought to have a nutritive function. Astrocytes also play an impor-
Schwann cells are positioned along the length of a single axon. tant role in maintaining the blood-brain barrier (See Appendix D).

Each Nerve Cell Is Part of a Circuit That

Has One or More Specific Behavioral
Functions

Every behavior is mediated by specific sets of to produce the familiar knee jerk. By increasing the
interconnected neurons, and every neuron’s tension of a select group of muscles, the stretch reflex
behavioral function is determined by its Changes the position of the leg, suddenly extending it
connections with other neurons. We can illustrate outward (Figure 2–6).Thus the electrical signals that
this with a simple behavior, the knee-jerk reflex. The produce the stretch reflex carry four kinds of
reflex is initiated when a transient imbalance of the information:
body stretches the quadriceps extensor muscles of 1. Sensory information is conveyed to the
the leg. This stretching elicits sensory information central nervous system (the spinal cord) from
that is conveyed to motor neurons, which in turn muscle
sends commands to the extensor muscles to 2. Motor commands from the central nervous
contract so that balance is restored. This reflex is system are issued to the muscles that carry
useful for clinically, but the underlying mechanism out the knee jerk.
3. Inhibitory commands are issued to motor
is important because it continuously maintains neurons that innervate opposing muscles,
normal tone in the quadriceps and prevents our providing coordination of muscle action.
knees from buckling when we stand or walk. 4. Information about local neuronal activity
The tendon of the quadriceps femoris, an related to the knee jerk is sent to higher
exten- sor muscle that moves the lower leg, is centers of the central nervous system,
attached to the tibia through the tendon of the permitting the brain to coordinate different
kneecap. Tapping this tendon just below the patella behaviors either simultaneously or in series.
stretches the quadriceps femoris. This stretch
initiates reflex contraction of the quadriceps muscle
5
 Part I / Overall Perspective

Stimulus

Sensory
neuron
Muscle Spinal
Quadriceps spindle cord
(extensor)

Hamstring
(flexor)
Inhibitory
Extensor Flexor interneuron
motor motor
neuron neuron
(activated) (inhibited)
neurons that contract the quadriceps, the muscle that was
Figure 2–6 The knee-jerk refex is controlled by a simple
stretched. The sensory neurons act indirectly, through interneu-
circuit of sensory and motor neurons. Tapping the kneecap rons, to inhibit fexor motor neurons that would otherwise
with a refex hammer pulls on the tendon of the quadriceps contract the opposing muscle, the hamstring. These actions
femoris, a muscle that extends the lower leg. When the muscle combine to produce the refex behavior. In the drawing each
stretches in response to the pull of the tendon, information extensor and fexor motor neuron represents a population of
regarding this change in the muscle is conveyed to the central many cells.
nervous system by sensory neurons. In the spinal cord the
sensory neurons form excitatory synapses with extensor motor

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 Chapter 2 / Nerve Cells, Neural Circuitry, and Behavior

Model Sensory Motor Local Projection Neuroendocrin

neuron neuron neuron interneuron interneuron e cell

Region:

Input

Integrative

Conductive

Central
neuron
Output

Capillary
Muscle
Central
Central neuron
neuron

Figure 2–9 Most neurons, regardless of type, have four to the cell, whereas the output signal is a chemical substance
functional regions in which different types of signals are ejected by the cell into the synaptic cleft. Not all neurons share
generated. Thus the functional organization of most neurons all these features; for example, local interneurons often lack a
can be represented schematically by a model neuron. The input, conductive component.
integrative, and conductive signals are all electrical and integral

ranges from –40 to –80 mV; in muscle cells it is greater

Signaling Is Organized in the Same
still, about –90 mV. As we shall see in Chapter 6, the
Way in All Nerve Cells resting membrane potential results from two factors:
the unequal distribution of electrically charged ions, in
particular the positively charged Na+ and K+ ions, and
To produce a behavior, a stretch reflex for the selective permeability of the membrane.
example, each participating sensory and motor The unequal distribution of positively charged ions
nerve cell must generate four different signals in on either side of the cell membrane is maintained
sequence, each at different site within the cell: an
input signal, a trigger signal, a conducting signal,
and an output signal. Regardless of cell size and
shape, transmitter biochemistry, or behavioral
function, almost all neurons can be described by a
model neuron that has four functional components
that generate the four types of signals: a receptive
component, a summing or integrative component, a
long-range signaling component, and a secretory
component (Figure 2–9).
The different types of signals generated in a neu-
ron are determined in part by the electrical properties
of the cell membrane. Every cell, including a neuron,
maintains a certain difference in the electrical potential
on either side of the plasma membrane when the cell is
at rest. This is called the resting membrane potential. In a
typical resting neuron the voltage of the inside of the cell
is about 65 mV more negative than the voltage outside
the cell. Because the voltage outside the membrane is
defined as zero, we say the resting membrane potential
is –65 mV. The resting potential in different nerve cells 7
 Part I / Overall Perspective

by two main mechanisms. Intracellular Na+ and K+

concentrations are largely controlled by a
membrane protein that actively pumps Na+ out of
the cell and K+ back into it. This Na+-K+ pump, about
which we shall learn more in Chapter 6, keeps the
Na+ concentration in the cell low (about one-tenth the
concentration outside the cell) and the K+
concentration high (about 20 times the concentration
outside). The extracellular concentrations of Na+ and
K+ are maintained by the kidneys.
The cell membrane is selectively permeable to K+
because the otherwise impermeable membrane
contains proteins that form pores called ion
channels. The channels that are active when the
cell is at rest are highly permeable to K+ but
considerably less permeable to Na+. The K+ ions
tend to leak out of these open channels, down the
ion’s concentration gradient. As K+ ions exit the cell,
they leave behind a cloud of unneutralized negative
charge on the inner surface of the membrane, so
that the net charge inside the membrane is more
negative than that outside.
A cell, such as nerve and muscle, is said to be
excitable when its membrane potential can be
quickly and significantly altered. This change serves
as a signaling mechanism. In some neurons
reducing the membrane potential by 10 mV (from –
65 to –55 mV) makes the

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 Chapter 2 / Nerve Cells, Neural Circuitry, and Behavior

membrane much more permeable to Na+ than to K+. Thus, unlike the action potential, which is all or none,
The resultant influx of positively charged Na+ neutral- receptor potentials are graded. Most receptor poten-
izes the negative charge inside the cell and causes a tials are depolarizing (excitatory). However, hyperpo-
brief and explosive change in membrane potential to larizing (inhibitory) receptor potentials are found in
+40 mV. This action potential is conducted down the the retina.
cell’s axon to the axon’s terminal, where it initiates an The receptor potential is the first representation
elaborate chemical communication with other neu- of stretch to be coded in the nervous system. This
rons or muscle cells. The action potential is actively signal spreads passively, however, and therefore
propagated along the axon so that its amplitude does does not travel much farther than 1 to 2 mm. In fact, 1
not diminish by the time it reaches the axon terminal. mm down the axon the amplitude of the signal is only
An action potential typically lasts approximately 1 ms, about one- third what it was at the site of generation.
after which the membrane returns to its resting state, To be carried successfully to the central nervous
with its normal separation of charges and higher per- system, the local signal must be amplified—it must
meability to K+ than to Na+. generate an action potential. In the knee-jerk reflex
In addition to the long distance signals the receptor potential in the sensory neuron must
represented by the action potential, nerve cells also pro- reach the first node of Ranvier in the axon. If it is
duce local signals—receptor potentials and synaptic large enough, the signal triggers an action potential
potentials—that are not actively propagated and that that then propagates without failure to the axon
typically decay within just a few millimeters. terminals in the spinal cord (Figure 2–10C). At the
The change in membrane potential that generates synapse between the sensory neuron and a motor
long-range and local signals can be either a decrease neuron, the action potential produces a chain of
or an increase from the resting potential. The resting events that results in an input signal to the motor
membrane potential therefore provides the baseline on neuron.
which all signaling occurs. A reduction in membrane In the knee-jerk reflex the action potential in the
potential is called depolarization. Because depolarization presynaptic terminal of the sensory neuron initiates
enhances a cell’s ability to generate an action potential, the release of a chemical substance, or neurotransmit-
it is excitatory. In contrast, an increase in membrane ter, into the synaptic cleft (Figure 2–10D). After diffus-
potential is called hyperpolarization. Hyperpolarization ing across the cleft, the transmitter binds to receptor
makes a cell less likely to generate an action potential proteins in the postsynaptic membrane of the motor
and is therefore inhibitory. neuron, thereby directly or indirectly opening ion
channels. The ensuing flow of current alters the mem-
The Input Component Produces Graded brane potential of the motor cell, a change called the
Local Signals synaptic potential.
Like the receptor potential, the synaptic poten-
In most neurons at rest no current flows from one part tial is graded; its amplitude depends on how much
of the cell to another, so the resting potential is the same transmitter is released. In the same cell the synaptic
throughout. In sensory neurons current flow is typi- potential can be either depolarizing or hyperpolariz-
cally initiated by a physical stimulus, which activates ing depending on the type of receptor molecule that is
specialized receptor proteins at the neuron’s recep- activated. Synaptic potentials, like receptor potentials,
tive surface. In our example of the knee-jerk reflex, spread passively and thus are local changes in poten-
stretching of the muscle activates specific ion tial unless the signal reaches beyond the axon’s initial
channels that open in response to stretch of the segment and thus can give rise to an action potential.
sensory neuron membrane. The opening of these The features of receptor and synaptic potentials are
channels when the cell is stretched permits the rapid summarized in Table 2–1.
influx of Na+ ions into the sensory cell. This ionic
current changes the membrane potential, producing a
The Trigger Zone Makes the Decision to Generate
local signal called the receptor potential.
an Action Potential
The amplitude and duration of a receptor poten-
tial depend on the intensity of the muscle stretch: The Sherrington first pointed out that the function of
larger or longer-lasting the stretch, the larger or longer- the nervous system is to weigh the consequences of
lasting the resulting receptor potential (Figure 2–10A). different types of information and then decide on
appropriate responses. This integrative function of the
nervous system is clearly seen in the actions of the
trigger zone of the neuron, the initial segment of the
axon.
9
 Part I / Overall Perspective

Muscle Trigger Sensory neuron

spindle zone cell body
Myelinated
axon
Synaptic
Stretch terminal

A Receptor potential B Trigger action C Action potential D Output signal

(transmitter release)
20
0
Stimulus (stretch) –20
–40
Spike threshold
Amplitude –60
–80
Duration
Membrane potential (mV)

20
0
–20
–40
Spike threshold
–60
–80

20
0
–20
–40
–60
–80

0 5 10 0 5 10 0 5 10
Time (s)

Figure 2–10 Each of the neuron’s four signaling compo- receptor potential determines the duration of the train of action
nents produces a characteristic signal. The fgure shows potentials. Thus the graded amplitude and duration of the
a sensory neuron activated by stretching of a muscle, which receptor potential is translated into a frequency code in the
the neuron senses through a specialized receptor, the muscle action potentials generated at the trigger zone. All action poten-
spindle. tials produced are propagated faithfully along the axon.
A. The input signal, called a receptor potential, is graded in C. Action potentials are all-or-none. Because all action poten-
amplitude and duration, proportional to the amplitude and dura- tials have a similar amplitude and duration, the frequency
tion of the stimulus. and duration of fring represents the information carried by
B. The trigger zone sums the depolarization generated by the the signal.
receptor potential. An action potential is generated only if the D. When the action potential reaches the synaptic terminal,
receptor potential exceeds a certain voltage threshold. Once it initiates the release of a neurotransmitter, the chemical sub-
this threshold is surpassed, any further increase in amplitude stance that serves as the output signal. The frequency of action
of the receptor potential can only increase the frequency with potentials determines exactly how much neurotransmitter is
which the action potentials are generated, because action released by the cell.
potentials have a constant amplitude. The duration of the

Action potentials are generated by a sudden influx Because the initial segment of the axon has the
of Na+ through channels in the cell membrane that highest density of voltage-sensitive Na+ channels and
open and close in response to changes in membrane therefore the lowest threshold for generating an action
potential. When an input signal (a receptor potential or potential, an input signal spreading passively along the
synaptic potential) depolarizes an area of membrane, cell membrane is more likely to give rise to an action
the local change in membrane potential opens local potential at the initial segment than at other sites in
Na+ channels that allow Na+ to flow down its concen- the cell. This part of the axon is therefore known as the
tration gradient, from outside the cell where the Na+ trigger zone. It is here that the activity of all receptor
concentration is high to inside where it is low. (or synaptic) potentials is summed and where, if the

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 Chapter 2 / Nerve Cells, Neural Circuitry, and Behavior

sum of the input signals reaches threshold, the neuron

generates an action potential.
If signals are stereotyped and reflect only the most
elementary properties of the stimulus, how can they
The Conductive Component Propagates an
carry the rich variety of information needed for com-
All-or-None Action Potential
plex behavior? How is a message that carries visual
The action potential is all-or-none: Stimuli below the information about a bee distinguished from one that
threshold do not produce a signal, but stimuli above carries pain information about the bee’s sting, and how
the threshold all produce the signals of the same are these sensory signals distinguished from motor sig-
amplitude. However much the stimuli vary in inten- nals for voluntary movement? The answer is simple
sity or duration, the amplitude and duration of each and yet is one of the most important organizational
action potential are pretty much the same. In addition, principles of the nervous system: Pathways of con-
unlike receptor and synaptic potentials, which spread nected neurons, not individual neurons, convey infor-
passively and decrease in amplitude, the action poten- mation. Interconnected neurons form anatomically and
tial does not decay as it travels along the axon to functionally distinct pathways. The neural pathways
its target—a distance that can be as great as 2 m— activated by receptor cells in the retina that respond to
because it is periodically regenerated. This light are completely distinct from the pathways acti-
conducting signal can travel at rates as fast as 100 vated by sensory cells in the skin that respond to touch.
m/s.
Only two features of the conducting signal The Output Component Releases Neurotransmitter
con- vey information: the number of action
potentials and the time intervals between them When an action potential reaches a neuron’s terminal it
(Figure 2–10C). As Adrian put it in 1928, stimulates the release of chemical substances from the
summarizing his work on sensory fibers: “all cell. These substances, called neurotransmitters, can be
impulses are very much alike, whether the message small organic molecules, such as *-glutamate and
is destined to arouse the sensation of light, of touch, or acetylcholine, or peptides like substance P or LHRH
of pain; if they are crowded together the sensation (luteinizing hormone releasing hormone).
is intense, if they are separated by long inter- vals the Neurotransmitter molecules are held in subcellular
sensation is correspondingly feeble.” Thus, what organelles called synaptic vesicles, which accumulate at
determines the intensity of sensation or speed of specialized release sites in the terminals of the axon called
movement is the frequency of the action potentials. active zones. To eject their transmitter substance into the
Likewise, the duration of a sensation or movement is synaptic cleft, the vesicles move up to and fuse with the
determined by the period over which action neuron’s plasma membrane, then burst open, a process
potentials are generated. known as exocytosis. The molecular machinery of neuro-
In addition to the frequency of the action transmitter release is described in Chapters 11 and 12.
potentials, the pattern of action potentials also Once released, the neurotransmitter is the neu-
conveys important information. For example, some ron’s output signal. Like the input signal, it is graded.
neurons are not silent in the absence of stimulation but The amount of transmitter released is determined by
are spontaneously active. Some spontaneously active the number and frequency of the action potentials that
nerve cells (beating neurons) reach the presynaptic terminals (Figure 2–10C,D). After
fire action potentials regularly; other neurons release the transmitter diffuses across the synaptic cleft
(bursting neurons) fire in brief bursts of action and binds to receptors on the postsynaptic neuron. This
potentials. These diverse cells respond differently to binding causes the postsynaptic cell to generate a syn-
the same excitatory synaptic input. An excitatory aptic potential. Whether the synaptic potential has an
synaptic potential may initiate one or more action excitatory or inhibitory effect depends on the type of
potentials in a cell that does not have a receptor in the postsynaptic cell, not on the particular
spontaneous activity, but in spontaneously active chemical neurotransmitter. The same transmitter sub-
cells that same input will modulate the rhythm by stance can have different effects at different receptors.
increasing the rate of firing of action potentials.
The Transformation of the Neural Signal from
Sensory to Motor Is Illustrated by the Stretch-Reflex
Pathway
We have seen that the properties of a signal are trans-
formed as the signal moves from one component of

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 Part I / Overall Perspective

a neuron to another or between neurons. This short one that regeneration of the signal is not
trans- formative chain of events can be seen in the required. In these neurons the input signals are
relay of signals for the stretch reflex. summed and spread passively to the pre-synaptic
When a muscle is stretched, the amplitude terminal region nearby where transmitter is
and duration of the stimulus are reflected in the released. Neurons that are spontaneously active do not
amplitude and duration of the receptor potential require sensory or synaptic inputs to fire action
generated in the sensory neuron (Figure 2–12A). If potentials because they have a special class of ion
the receptor potential exceeds the threshold for an channels that permit Na+ current flow even in the
action potential in that cell, the graded signal is absence of excitatory synaptic input.
transformed at the trigger zone into an action Even cells that are similar morphologically can
potential, an all-or-none signal. The more the differ importantly in molecular details. For example,
receptor potential exceeds threshold, the greater they can have different combinations of ion channels.
the depolarization and consequently the greater As we shall learn in Chapter 7, different ion channels
the frequency of action potentials in the axon. The provide neurons with various thresholds, excitability
duration of the input signal also determines the properties, and firing patterns (Figure 2–11).
duration of the train of action potentials. Neurons with different ion channels can therefore
The information encoded by the frequency encode synaptic potentials into different firing
and duration of firing is faithfully conveyed along patterns and thereby convey different information.
the axon to its terminals, where the firing of Neurons also differ in the chemical substances they
action potentials determines the amount of use as transmitters and in the receptors that receive
transmitter released. These stages of signaling have transmitter substances from other neurons. Indeed,
their counterparts in the motor neuron (Figure 2–12B) many drugs that act on the brain do so by modifying the
and in the muscle (Figure 2–12C). actions of specific chemical transmitters or
Nerve Cells Differ Most at the Molecular receptors.
Level Despite the differences among nerve cells, the
The model of neuronal signaling we have outlined is basic mechanisms of electrical signaling are
a simplification that applies to most neurons but there surprisingly similar. This simplicity is fortunate, for
are some important variations. For example, some understanding the molecular mechanisms of
neurons do not generate action potentials. These signaling in one kind of nerve cell aids the
are typically local interneurons without a understanding of these mechanisms in many other
conductive component; they have no axon or such a nerve cells.

12
 Part I / Overall Perspective

A A

A Sensory signals B Motor signals C Muscle signals

Stimulus Input Integration Conduction Output Input Integration Conduction Output Input Integration Conduction Output
(transmitter (transmitter (behavior)
release) release)
Action
Graded potential Action Action Graded Action Action Action Graded Action Action
receptor potential potential synaptic potential potential potential synaptic potential potential
potential Receptor potential Synaptic potential Synaptic
potential potential potential

Stretch Contraction

Muscle spindle Sensory neuron Motor neuron Muscle

Figure 2–12. The sequence of signals that produces a that innervates the stretched muscle; it then binds to receptor
refex action. molecules on the external membrane of the motor neuron.
A. The stretching of a muscle produces a receptor potential in B. This interaction initiates a synaptic potential that spreads
the specialized receptor (the muscle spindle). The amplitude passively to the trigger zone of the motor neuron’s axon,
of the receptor potential is proportional to the intensity of the where it initiates an action potential that propagates actively
stretch. This potential spreads passively to the integrative or to the terminal of the motor neuron’s axon. The action potential
trigger zone at the frst node of Ranvier. If the receptor poten- releases a neuro transmitter where the axon terminal meets
tial is suffciently large, it triggers an action potential that then a muscle fber.
propagates actively and without change along the axon to the C. The neurotransmitter binds receptors on the muscle fber,
axon terminal. At specialized sites in the terminal the action triggering a synaptic potential in the muscle. If suffciently
potential leads to an output signal, the release of a chemical large, or if combined with signals from other motor neurons,
neurotransmitter. The transmitter diffuses across the synaptic the synaptic potential will generate an action potential in the
cleft between the axon terminal and a target motor neuron muscle, causing contraction of the muscle fber.

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