10

Integrative Functions of
the Nervous System
LEARNING OBJECTIVES of the cortex is highly convoluted and folded into ridges
Upon completion of this chapter, the student should be able to known as gyri. Gyri are separated by grooves called sulci
answer the following questions: (if shallow) or fissures (if deep; see Fig. 4.7). This folding
1. What is the basic layering pattern of the neocortex, greatly increases the surface area of cortex that can be fit
and how do cortical inputs and outputs align with this into the limited and fixed volume within the skull. Indeed,
layering pattern? What is the functional significance of the most of the cortex cannot be seen from the brain surface
variation in the layering pattern between cortical areas? because of this folding.
2. What are the major functions of each of the lobes of the The cerebral cortex can be divided into the left and right
cerebrum? hemispheres and subdivided into a number of lobes (Fig.
3. How does the electroencephalogram (EEG) reflect cortical 10.1; see also Fig. 4.7), including the frontal, parietal,
activity? What are evoked potentials? temporal, and occipital lobes. The frontal and parietal
4. How does cerebral dominance correlate with language
lobes are separated by the central sulcus; both are separated
and hand preference?
5. What is aphasia, and what is compromised in the
from the temporal lobe by the lateral fissure. The occipital
different types of aphasia? and parietal lobes are separated (on the medial surface of
6. How do synaptic and cellular processes support learning the hemisphere) by the parieto-occipital fissure (see Fig.
and memory? How is memory distributed in the brain? 10.1). Buried within the lateral fissure is another lobe, the
7. What role does plasticity play in neural development and insula (see Fig. 4.6A). A group of structures that make up
in response to damage of the nervous system? the limbic lobe is on the medial aspect of the hemisphere,
and its largest part, the hippocampal formation, is folded
into the parahippocampal gyrus of the temporal lobe and
cannot be seen from the surface of the brain.

I
n earlier chapters, the interaction of the nervous system Activity in the two hemispheres of the cerebral cortex
with the body and the outside world was discussed is coordinated by interconnections through the cerebral
in terms of the transduction and analysis of sensory commissures. The bulk of the cortex is connected through
events, the organization of motor function, and relatively the massive corpus callosum (see Figs. 4.9, 10.1), and
simple central processes that link them, such as reflexes parts of the temporal lobes connect through the anterior
(e.g., the stretch reflex and the vestibuloocular reflex). The commissure.
nervous system has other capabilities, so-called integrative There are three types of cerebral cortex: neocortex,
or higher functions, that are less directly tied to specific archicortex, and paleocortex. The neocortex has six corti-
sensory modalities or motor behavior. These functions, in cal layers (Fig. 10.2). In contrast, the archicortex has only
particular, require interactions between different parts of three layers, and the paleocortex has four to five layers.
the cerebral cortex, and, as is being increasingly recognized, In humans, approximately 90% of the cerebral cortex is
between the cerebral cortex and other parts of the brain. The neocortex.
neural basis for some of these higher functions is discussed
in this chapter. Because these functions (as well as sensory
The Neocortex
perception and voluntary motor function) are so highly
dependent on the cerebral cortex, its basic organization is Neuronal Cell Types in the Neocortex
described first. A number of different neuronal cell types in the neocortex
have been described (see Fig. 10.2). Pyramidal cells are
The Cerebral Cortex the most abundant cell type and account for approximately
75% of neocortical neurons. Stellate cells and various
The human cerebral cortex occupies a volume of about other types of nonpyramidal neurons make up the balance.
600 cm3 and has a surface area of 2500cm2. The surface Pyramidal cells have a large triangular cell body, a long

208
 CHAPTER 10 Integrative Functions of the Nervous System 209

Central sulcus Postcentral gyrus

Postcentral sulcus
Precentral gyrus

Superior parietal lobule

Precentral sulcus Intraparietal sulcus
Superior frontal sulcus Inferior parietal lobule:
Superior frontal gyrus Supramarginal gyrus
Angular gyrus
Middle frontal gyrus
Inferior frontal
sulcus
Frontal pole

Inferior frontal gyrus:

Pars triangularis
Pars opercularis
Pars orbitalis
Preoccipital notch

Orbital surface
Middle temporal sulcus
Lateral sulcus
Temporal pole
Superior temporal gyrus Inferior temporal gyrus
Superior temporal sulcus Middle temporal gyrus

Frontal lobe Parietal lobe Temporal lobe Occipital lobe

A
Hippocampal
Central sulcus commissure

Marginal sulcus Superior frontal gyrus

Precuneus Cingulate sulcus

Parieto-occipital fissure Cingulate gyrus

Cuneus
Callosal sulcus
B
Septum G Frontal pole
Calcarine sulcus
Fornix
S R
Subcallosal area
Occipital pole Thalamus
Anterior commissure

Lingual gyrus Lamina terminalis

Isthmus of cingulate gyrus Temporal pole
Optic chiasm
Pineal body Uncus
Parahippocampal gyrus
Collateral sulcus
Posterior commissure

Occipital lobe Parietal lobe Temporal lobe Limbic lobe Frontal lobe
B
Fig. 10.1 Lateral (A) and medial (B) illustrations of the left hemisphere of the human cerebrum with the

major features labeled and the lobes indicated by color. R, G, B, and S indicate, respectively, the rostrum,
genu, body, and splenium of the corpus callosum. (From Haines DE [ed]. Fundamental Neuroscience for
Basic and Clinical Applications. 3rd ed. Philadelphia: Churchill Livingstone; 2006.)
210 S E C T I O N 2 Berne & Levy Physiology

Golgi stain Nissl stain Weigert stain

I. Molecular layer

II. External
granular layer

III.External
pyramidal layer

IV. Internal Outer band

granular layer of Baillarger

V. Internal
pyramidal layer Inner band
of Baillarger

VI.Polymorphic layer

Fig. 10.2 An Area of Neocortex Stained by Three Different Methods. The Nissl stain (center) shows

the cell bodies of all neurons and reveals how different types are distributed among the six layers. The
Golgi stain (left) shows only a sample of the neuronal population but reveals details of their dendrites.
The Weigert stain for myelin (right) demonstrates vertically oriented bundles of axons entering and leaving
the cortex and horizontally coursing fibers that interconnect neurons within a layer. (From Brodmann
K. Vergleichende Lokalisationslehre der Grosshirnrinde in ihren prinzipien Dargestellt auf Grund des
Zellenbaues. Leipzig: JA Barth; 1909.)

apical dendrite directed toward the cortical surface, and Cytoarchitecture of Cortical Layers
several basal dendrites. The cells axon emerges from the Each of the six layers of the neocortex has a characteristic
body opposite the apical dendrite, and those from the larger cellular content (see Fig. 10.2). Layer I (molecular layer) has
pyramidal cells project into the subcortical white matter. few neuronal cell bodies and contains mostly axon terminals
The axon may give off collateral branches as it descends synapsing on apical dendrites. Layer II (external granular
through the cortex. The neurotransmitter of pyramidal cells layer) contains mostly stellate cells. Layer III (external pyra-
is an excitatory amino acid (glutamate or aspartate). Stel- midal layer) consists mostly of small pyramidal cells. Layer
late cells, often called granule cells, are interneurons with IV (internal granular layer) contains mostly stellate cells
local connections. They have a small soma and numerous and a dense matrix of axons. Layer V (internal pyramidal
branched dendrites, although many have an apical dendrite layer) is dominated by large pyramidal cells, the main source
and thus look like small pyramidal cells. Some are excitatory of cortical efferents to most subcortical regions. Layer VI
interneurons; these cells are abundant in layer IV of the (multiform layer) contains pyramidal, fusiform, and other
cortex (described in the next section). Their axons remain types of cells.
in the same cortical region, and many ascend into the
upper cortical layers. Some stellate cells are inhibitory inter- Cortical Afferent and Efferent Fibers
neurons whose neurotransmitter is gamma-aminobutyric Most input to the cortex from other regions of the central
acid (GABA). nervous system (CNS) is relayed by neurons in the thalamus,
 CHAPTER 10 Integrative Functions of the Nervous System 211

as described in earlier chapters for sensory and motor markers) are also used. Although several cytoarchitectonic
pathways. The projections from the thalamus to the cortex maps of the cortex have been devised, the one by Korbinian
are a significant component of cortical organization that is Brodmann is most commonly used. In this map, the cortex
observed clearly in the layering pattern. Thalamocortical is divided into 52 discrete areas (Fig. 10.3), numbered in
fibers from thalamic nuclei that have specific (topographi- the order that Brodmann studied them. Areas commonly
cally mapped) cortical projections end chiefly in layer IV referred to include Brodmann areas 3, 1, and 2 (the
but also in layers III and VI. Neurons in other thalamic primary somatosensory cortex located on the postcentral
nuclei (particularly those relaying input from the brainstem gyrus); area 4 (the primary motor cortex located on the
reticular formation) project diffusely and terminate in layers precentral gyrus); area 6 (the premotor and supplementary
I and VI to modulate cortical activity globally, perhaps in motor cortex); areas 41 and 42 (the primary auditory cortex
conjunction with changes in state (e.g., sleep or waking). on the superior temporal gyrus); and area 17 (the primary
In addition to subcortical inputs, every region of the visual cortex, mostly on the medial surface of the occipi-
cortex receives input from other cortical regions. There are tal lobe). Subsequent studies confirmed that Brodmann
some large fiber bundles that connect widely separated cor- areas are distinctive with regard to their cytoarchitecture,
tical regions, and commissural fibers connect corresponding interconnections, and functions, but more recent work has
regions in each hemisphere (these projections terminate in shown that there is some plasticity both in the size of the
layers I and VI), but in relative terms, the largest source of areas and in their internal organization (see the section
synapses in a cortical region is local, either from within the Neural Plasticity).
region itself or from its neighbors. Although cytoarchitectonic maps, like Brodmanns, give
The cortical efferent axons originate from pyramidal cells. the impression of sharp boundaries between contiguous
The smaller pyramidal cells of layers II and III project to areas, the variation between many of the defined cortical
adjacent cortical areas directly and to contralateral regions areas is actually fairly subtle and, rather than sharing a well-
via the corpus callosum. The larger pyramidal cells of layer defined border, most neighboring regions may gradually
V project in many pathways to synaptic targets in the spinal transition into one another. Nevertheless, some areas have
cord, brainstem, striatum, and thalamus. The pyramidal quite distinct cortical characteristics, particularly the primary
cells of layer VI form corticothalamic projections that sensory and motor cortices. For example, the primary and
target the same thalamic nuclei that are their input creating premotor areas are referred to as agranular cortex, because
circuits of reciprocal thalamocortical and corticothalamic no clear layer IV is present in these areas. Moreover, among
interconnections. In addition, intrathalamic connections the motor areas, the primary motor cortex is distinguished
serve to associate activity in different cortical regions. by the presence of large layer V pyramidal cells, the largest
The specific patterns of input from the thalamus have of which are called Betz cells. These enormous cells have
another influence on cortical organization. As discussed in axons that contribute to the corticospinal tracts and whose
the sensory and motor systems, the topographic mapping soma size (diameter > 150m) is necessary for the meta-
of cortical input defines a columnar organization. A bolic maintenance of so much axoplasm. Note that despite
column is a narrow, vertically oriented (from the white being the histological criterion for identifying primary
matter to the cortical surface) region in which the neurons motor cortex, Betz cell axons account for less than 5% of
have correlated activity because of shared input from the all corticospinal fibers.
thalamus. Within a column, there is a great richness in In contrast to the motor areas, the primary sensory corti-
vertical interconnections and fewer lateral interconnections ces (e.g., somatosensory, auditory, and visual) typically have
(to cells in neighboring columns), which suggests that a very prominent layer IV (internal granular layer), which is
columns may be the functional unit of the cortex. Despite dominated by stellate cells (see Fig. 10.2), and therefore they
their relative paucity, however, the lateral interconnections are classified as granular cortices. Indeed, the primary visual
can exert powerful actions, as shown by inhibitory inter- cortex is also known as the striate cortex because of a par-
connection between regions within motor cortex (see Fig. ticularly prominent horizontal sheet of myelinated axons in
9.16). Interestingly, the columnar organization can greatly layer IV known as the stripe of Gennari. In a sense, the terms
influenced by functional interactions, as well as by genetics. granular and agranular are inaccurate because all cortical
(See the section Neural Plasticity.) areas have similar percentages of pyramidal calls (75%) and
nonpyramidal cells (25%). Nevertheless, the key idea is that
Regional Variations in Neocortical Structure the grouping of the cell types into layers varies dramatically
The architecture of the neocortex varies regionally, which between the frontal motor areas, where the nonpyramidal
presumably reflects the functional specialization of cortical neurons do not form a distinct internal granular layer, and
areas. Different aspects of this variation are the bases of the primary sensory cortices, where they do.
several methods for subdividing the cortex into discrete
areas. The most widely used method is cytoarchitectonics, Archicortex and Paleocortex
in which variations in cell density and structure are used;
however, myeloarchitectonics (variations in axon density About 10% of the human cerebral cortex is archicortex and
and size) and chemoarchitectonics (expression of molecular paleocortex. The archicortex has a three-layered structure;
212 S E C T I O N 2 Berne & Levy Physiology

4
6 3 1
25
8 7a
7b
9

19

46 40
10 39 18

45 44 43 41
42
17
47 22
11
21
18
38 37 19

20
A

4
6 312
5
8
7
9
31
24
32 23
33 19
10
26 18
29
30
12 25 27
11 34 35 17

28
38 36 18
19
37

B 20
Fig. 10.3
Brodmanns areas in the human cerebral cortex. (Redrawn from Crosby EC, etal. Correlative

Anatomy of the Nervous System. New York: Macmillan; 1962.)

the paleocortex has four to five layers. The paleocortex Functions of the Lobes of the Cerebral Cortex
is located at the border between the archicortex and
neocortex. There is no exact correspondence between the folds (lobes
In humans, the hippocampal formation is part of the and gyri) of the cerebral cortex and function; nevertheless,
archicortex. It is folded into the temporal lobe and can be some with the individual lobes of the cerebral hemispheres
viewed only when the brain is dissected. The hippocampal have a general association with function that helps clarify
cortex has three layers: the molecular, pyramidal cell, and cortical organization.
polymorphic layers. They resemble layers I, V, and VI of
the neocortex. The white matter covering the hippocampus Frontal Lobe
is called the alveus, which contains hippocampal afferent One of the main functions of the frontal lobe is motor
and efferent fibers. The efferent axons coalesce to form the behavior. As discussed in Chapter 9, the motor, premotor,
fornix (Fig. 10.4). cingulate motor, and supplementary motor areas are located
 CHAPTER 10 Integrative Functions of the Nervous System 213

Anterior nucleus
Fornix of thalamus
Thalamus
Cingulate gyrus

Basal
forebrain

Prefrontal
cortex

Mammillary Rhinal Hippocampus

body Amygdala cortex
Fig. 10.4 The hippocampus and the amygdala are located on the medial aspect of the temporal lobe.

The fornix, the major output pathway from the hippocampus, projects to the mammillary body, which in
turn connects to the anterior nucleus of the thalamus via the mammillothalamic tract. Also illustrated are
the cingulate gyrus, the basal forebrain area (septal nuclei, bed nucleus of the stria terminalis, nucleus
accumbens), and the prefrontal cortex. (From Purves D. Sleep and wakefulness. In: Purves D, etal [eds].
Neuroscience. 3rd ed. Sunderland, MA: Sinauer; 2004.)

in the frontal lobe, as is the frontal eye field. These areas are information. Connections with the frontal lobe allow
crucial for planning and executing motor behavior. Brocas somatosensory information to aid in voluntary motor activ-
area, essential for the generation of speech, is located in ity. Somatosensory, visual, and auditory information can
the inferior frontal gyrus of the dominant hemisphere for also be transferred to language centers, such as Wernickes
human language (almost always the left hemisphere, as area, as described later. Lesions in the left parietal lobe can
explained later). In addition, the more anterior prefrontal result in Gerstmanns syndrome, which includes a persons
cortex in plays a major role in personality and emotional inability to name his or her fingers (or those of another) and
behavior. a loss of the ability to perform numerical calculations. The
Bilateral lesions of the prefrontal cortex may be produced right parietal lobe is involved in determining spatial context.
either by disease or by a surgical frontal lobotomy. Such Localized lesions can result in the neglect syndrome, in
lesions produce deficits in attention, difficulty in planning which the patient seems unaware of the left side of his or
and problem solving, and inappropriate social behavior. her body and of persons, objects, and events on his or her
Aggressive behavior is also lessened and the motivational- left. (See an In the Clinic box later in this chapter.)
affective component of pain is reduced although pain
sensation remains. Frontal lobotomies are rarely performed Occipital Lobe
today because modern drug therapies provide more focal The major function of the occipital lobe is visual processing
and effective management for mental illness and chronic and perception (see Chapter 8). The primary visual cortex
pain. (Brodmann area 17) lines the calcarine sulcus and is flanked
by secondary (Brodmann area 18) and tertiary (Brodmann
Parietal Lobe area 19) visual cortices. Lesions of these areas in the
The parietal lobe contains the somatosensory cortex (see cuneus gyrus result in blindness in the lower contralateral
Chapter 7) and the adjacent parietal association cortex. visual field; those in the lingual gyrus result in blindness
The parietal association cortex gets information from in the upper contralateral visual field. Connections to the
somatosensory, visual, and auditory cortices and is involved frontal eye fields affect direction of gaze, and projections
in the processing, perception, and integration of sensory to the midbrain assists in the control of convergent eye
214 S E C T I O N 2 Berne & Levy Physiology

Although this syndrome was originally described as follow-

IN THE CLINIC ing large lesions of most or all of the temporal lobe, more
Two areas important for planning and executing motor tasks
recent studies have highlighted the role of the amygdala.
are the parietal cortex and the frontal cortex, the former The amygdala conditions the association of fear with painful
because it integrates sensory information needed to define stimuli and may trigger, via connections to the medial frontal
the context of a task (see Chapter 7) and the latter because cortex and anterior cingulate gyrus, emotional or avoidance
it has neurons that direct all the components for motor responses when these stimuli recur. In addition, the amyg-
execution (see Chapter 9). Mirror neurons have been found in
both the inferior parietal and the inferior frontal cortices
dala projects to the nucleus accumbens, a region of the
of macaques. These cells respond during performance basal ganglia which has been called a reward center. The
of a specific motor task and also during observation of the nucleus accumbens signals pleasurable events in response to
same task performed by another animal. Because these dopaminergic input from the ventral tegmental area of the
mirror cells seem to encode for and respond to very specific brainstem and is also the most common site of the plaques
and particular tasks, it has been speculated that they may
underlie such functions as understanding the intentions of
and tangles associated with Alzheimers disease.
others and empathy, as well as the ability to learn tasks from
observation. In humans, EEG activity consistent with the
behavior of such mirror neurons has been localized to the IN THE CLINIC
inferior frontal and superior parietal lobes. Autism, which
involves an inability to read the intentions and emotions of The functions of the different lobes of the cerebral cortex
others, has been linked to a lack of mirror neurons by EEG have been defined according to both the effects of lesions
evidence. produced by disease or those of surgical interventions to
treat disease in humans and from experiments on animals. In
another approach, the physical manifestations of epileptic
seizures have been correlated with the brain locations that
give rise to seizures (epileptic seizure foci). For example,
epileptic foci in the motor cortex cause movements on
movements, pupillary constriction, and accommodation, all the contralateral side; the exact movements relate to the
of which occur when the eyes adjust for near vision. somatotopic location of the seizure focus. Seizures that
originate in the somatosensory cortex cause an epileptic
Temporal Lobe aura in which a touch sensation is perceived. Similarly,
seizures that start in the visual cortex cause a visual aura
The temporal lobe has many different functions, includ- (scintillations, colors), those in the auditory cortex cause an
ing the processing and perception of sounds and vestibular auditory aura (humming, buzzing, ringing), and those in the
information and higher order visual processing (see Chapter vestibular cortex cause a sensation of spinning. Complex
8). For example, the infratemporal cortex, on its inferior behavior results from seizures that originate in the association
surface, is involved in the recognition of faces. In addition, areas of the temporal lobe; in addition, a malodorous
aura may be perceived if the olfactory cortex is involved
Meyers loop, which forms part of the optic pathway, passes (uncinate fit).
through the temporal lobe. As a result, temporal lobe lesions
can damage vision in upper part of the visual field. Similarly,
a portion of Wernickes area, essential for the understanding
of language, lies in the posterior region of the temporal lobe. The Electrical Activity of the Cortex
The limbic system dominates the medial temporal lobe
(see Fig. 10.4), and it participates in emotional behavior and An electroencephalogram (EEG) is a recording of the
in learning and memory (see the Learning and Memory neuronal electrical activity of the cerebral cortex made by
section). The limbic system helps control emotional electrodes placed on the skull. EEG waves normally reflect
behavior, in part by an influence on the hypothalamus via the summed extracellular currents that result from the gen-
the Papez circuit. This circuit projects from the cingulate eration of synaptic potentials in the pyramidal cells and are
gyrus to the entorhinal cortex and hippocampus, and thus a type of field potential. Because the currents generated
from there, via the fornix, to the mammillary bodies in by a single cell are too small to be detected as discrete events
the hypothalamus. The mammillothalamic tract then con- by an electrode on the skull (to record the activity of a single
nects the hypothalamus with the anterior thalamic nuclei, neuron, a microelectrode must be placed within microns of
which project back to the cingulate gyrus (see Fig. 10.4). In the neuron), the EEG waves reflect the combined activity
addition, the hippocampus and amygdala are connected to of many pyramidal cells. Moreover, for the activity of a
the prefrontal cortex, the basal forebrain, and the anterior group of neurons to generate an event detectable on EEG,
cingulate cortex. they must be oriented so that their individual currents
Bilateral temporal lobe lesions can produce Klver-Bucy summate to produce a detectable field. The arrangement of
syndrome, which is characterized by loss of the ability to pyramidal neurons, with their apical dendrites aligned in
recognize the meaning of objects from visual cues (visual parallel to form a dipole sheet, is particularly favorable for
agnosia); a tendency to examine all objects, even dangerous generating large field potentials. One pole of this sheet is
ones, orally; attention to irrelevant stimuli; hypersexuality; oriented toward the cortical surface and the other toward
a change in dietary habits; and decreased emotionality. the subcortical white matter. Thus the currents generated
 CHAPTER 10 Integrative Functions of the Nervous System 215

Drowsy (8 to 12 Hz) alpha waves

50 V
Stage 1 (3 to 7 Hz) theta waves
Theta waves
1 sec

Stage 2 (12 to 14 Hz) sleep spindles and K complexes

Sleep spindle K complex

Stage 4 ( to 2 Hz) delta waves

REM sleep

Fig. 10.5 Electroencephalographic tracings during drowsiness; stages 1, 2, and 4 of slow-wave

(nonrapid eye movement [non-REM]) sleep; and REM sleep. (Modified from Shepherd GM. Neurobiology.
London: Oxford University Press; 1983.)

by cortical pyramidal cells have a similar orientation and frequencies of the EEG recorded over the parietal and
can therefore summate to produce a field potential that occipital lobes are about 8 to 12Hz, the alpha rhythm.
can be detected. The need for summation also explains why If the subject is asked to open the eyes, the wave becomes
EEG signals reflect primarily synaptic potentials rather than less synchronized, and the dominant frequency increases to
action potentials: electrical events must overlap in time in 13 to 30Hz, which is called the beta rhythm. The delta
order to sum, and synaptic potentials have much longer (0.5 to 2Hz) and theta (3 to 7Hz) rhythms are observed
durations than do action potentials. during sleep (see the following discussion; Fig. 10.5). Also,
The sign of an EEG wave can be positive or negative, brief EEG waves do exist and, because of their shape, are
but its direction alone does not indicate whether pyramidal sometimes referred to as spikes, but this does not imply
cells are being excited or inhibited. For instance, a negative that they are associated with action potentials.
EEG potential may be generated at the surface of the skull
(or cortex) by excitation of apical dendrites or by inhibition Evoked Potentials
near the somas. Conversely, a positive EEG wave can be
produced by inhibition of apical dendrites or by excitation An EEG change that can be elicited by a stimulus is called
near the somas. a cortical evoked potential. A cortical evoked potential is
A normal EEG tracing consists of waves of various best recorded from the part of the skull located over the
frequencies. The dominant frequencies depend on several cortical area being activated. For example, a visual stimulus
factors, including the state of wakefulness, the age of the results in an evoked potential that can be recorded best
subject, the location of the recording electrodes, and the over the occipital bone, whereas a somatosensory evoked
absence or presence of drugs or disease. When a normal potential is recorded most effectively near the junction of
awake adult is relaxed with the eyes closed, the dominant the frontal and parietal bones. Evoked potentials reflect the
216 S E C T I O N 2 Berne & Levy Physiology

activity in large numbers of cortical neurons. They may also that seen in the EEG from an aroused subject (see Fig.
reflect activity in subcortical structures. 10.5, bottom trace). Because of the similarity of the EEG
Evoked potentials are small in comparison with the size of to that of an awake individual and the difficulty awaking
the EEG waves. However, their appearance can be enhanced the person, the term paradoxical sleep characterizes this
by a process called signal averaging. In this process, the type of sleep. Muscle tone is completely lost, but phasic
stimulation is repeated, and the EEGs recorded during each contractions occur in a number of muscles, most notably
trial are electronically averaged. With each repetition of the the eye muscles. The resulting rapid eye movements are
stimulus, the evoked potential occurs at a fixed time after basis of the name for this type of sleep. Many autonomic
the stimulus. When the records are averaged, the compo- changes also take place. Temperature regulation is lost, and
nents of the EEG that have a random temporal association meiosis occurs. Penile erection may occur during this type
with the stimulus cancel each other, whereas the evoked of sleep. Heart rate, blood pressure, and respiration change
potentials sum. intermittently. Several episodes of REM sleep occur each
night. Although it is difficult to arouse a person from REM
sleep, internal arousal is common. Most dreaming occurs
IN THE CLINIC during REM sleep.
Evoked potentials are used clinically to assess the integrity
of a sensory pathway, at least to the level of the primary
sensory receiving area. These potentials can be recorded IN THE CLINIC
in comatose individuals, as well as in infants too young
to undergo a sensory examination. The initial parts of the The sleep-wake cycle has an endogenous periodicity of
auditory evoked potential actually reflect activity in the about 25 hours, but it normally becomes entrained to the
brainstem; therefore, this evoked potential can be used to day-night cycle. The source of circadian periodicity appears
assess the function of brainstem structures. to be the suprachiasmatic nucleus of the hypothalamus. This
nucleus receives projections from the retina, and its neurons
seem to form a biological clock that adapts to the light-dark
cycle. However, the entrainment can be disrupted when the
subject is isolated from the environment or changes time
zones (jet lag). Destruction of the suprachiasmatic nucleus
Sleep-Wake Cycle disrupts a number of biological rhythms, including the
sleep-wake cycle.
Sleep and wakefulness are among the many functions of
the body that show circadian (about 1-day) periodicity.
Characteristic changes in the EEG can be correlated with
changes in the behavioral state during the sleep-wake The proportion of slow-wave (non-REM) sleep to REM
cycle. Beta wave activity dominates in an awake, aroused sleep varies with age. Newborns spend about half of their
individual. The EEG is said to be desynchronized; it sleep time in REM sleep, whereas elderly people have little
displays low-voltage, high-frequency activity. In relaxed REM sleep. About 20% to 25% of the sleep of young adults
individuals with their eyes closed, the EEG is dominated is REM sleep.
by alpha waves (see Fig. 10.5). A person falling asleep The mechanism of sleep is incompletely understood.
passes sequentially through four stages of slow-wave sleep Stimulation in the brainstem in a large region known as the
(called stages 1 through 4) over a period of 30 to 45 minutes reticular activating system causes arousal and low-voltage,
(see Fig. 10.5). In stage 1, alpha waves are interspersed fast EEG activity. Sleep was once thought to be caused by
with lower frequency waves called theta waves. In stage a reduced level of activity in the reticular activating system.
2, the waves slow further, but the slow-wave activity is However, substantial data, including the observations that
interrupted by sleep spindles, which are bursts of activity anesthesia of the lower brainstem results in arousal and that
at 12 to 14Hz, and by large K complexes (large, slow stimulation in the medulla near the nucleus of the solitary
potentials). Stage 3 sleep is associated with delta waves and tract can induce sleep, suggest that sleep is an active process.
with occasional sleep spindles. Stage 4 is characterized by Investigators have tried to find a relationship between sleep
delta waves without spindles. mechanisms and brainstem networks in which particular
During slow-wave sleep, the muscles of the body relax, neurotransmitters, including serotonin, norepinephrine,
but the posture is adjusted intermittently. The heart rate and acetylcholine, are used; manipulations of the levels of
and blood pressure decrease, and gastrointestinal motility these transmitters in the brain can affect the sleep-wake
increases. The ease with which individuals can be awakened cycle. However, a detailed neurochemical explanation of the
decreases progressively as they pass through these sleep neural mechanisms of sleep is not yet available.
stages. As individuals awaken, they pass through the sleep Similarly, the purpose of sleep is still unclear. However,
stages in reverse order. it must have a high value because so much of life is
About every 90 minutes, slow-wave sleep changes to a spent in sleep and because lack of sleep can be debilitat-
different form of sleep, called rapid eye movement (REM) ing. Medically important disorders of the sleep-wake cycle
sleep. In REM sleep, the EEG again becomes desynchro- include insomnia, bed-wetting, sleepwalking, sleep apnea,
nized. The low-voltage, fast activity of REM sleep resembles and narcolepsy.
 CHAPTER 10 Integrative Functions of the Nervous System 217

1 sec
A

50 V
B

100 V
C

100 V
D
Fig. 10.6 Electroencephalographic (EEG) Abnormalities in Several Forms of Epilepsy. A, EEG

tracings during the tonic (left) and clonic (right) phases of a tonic-clonic (grand mal) seizure. B, Spike
and wave components of an absence (petit mal) seizure. C, EEG tracing in a person with temporal lobe
epilepsy. D, EEG tracing of a focal seizure. (Redrawn from Eyzaguirre C, Fidone SJ. Physiology of the
Nervous System. 2nd ed. St. Louis: Mosby; 1975.)

IN THE CLINIC
The EEG becomes abnormal in a variety of pathological grand mal seizures. In petit mal epilepsy, consciousness is
circumstances. For example, during coma, the EEG is lost transiently (typically for less than 15 seconds), and the
dominated by delta activity. Brain death is defined by a EEG displays spike and wave activity (see Fig. 10.6B). In
maintained flat EEG wave. grand mal seizures, consciousness is lost for a longer period,
Epilepsy commonly causes, and can be diagnosed by, and the affected individual may fall if standing when the
specific EEG abnormalities. There are many forms of epilepsy, seizure starts. The seizure begins with a generalized increase
and examples of EEG patterns from some of these types of in muscle tone (tonic phase), followed by a series of jerky
epilepsy are shown in Figure 10.6. Epileptic seizures can be movements (clonic phase). The bowel and bladder may be
either partial or generalized. evacuated. The EEG shows widely distributed seizure activity
One form of partial seizures originates in the motor cortex (see Fig. 10.6A).
and results in localized contractions of contralateral muscles. EEG spikes that occur between full-blown seizures are
The contractions may then spread to other muscles; such called interictal spikes. Similar events can be studied
spread follows the somatotopic sequence of the motor cortex experimentally. These spikes arise from abrupt, long-lasting
(see Chapter 9). This stereotypical progression is called a depolarizations, called depolarization shifts, that trigger
Jacksonian march. Complex partial seizures (which may repetitive action potentials in cortical neurons. These
occur in psychomotor epilepsy) originate in the limbic depolarization shifts may reflect several changes in epileptic
structures of the temporal lobe and result in illusions and foci. Such changes include regenerative Ca++-mediated
semipurposeful motor activity. During and between focal dendritic action potentials in cortical neurons and a reduction
seizures, scalp recordings may reveal EEG spikes (see Fig. in inhibitory interactions in cortical circuits. Electrical field
10.6C and D). potentials and the release of K+ and excitatory amino acids
Generalized seizures involve wide areas of the brain and from hyperactive neurons may also contribute to the increased
loss of consciousness. Two major types are petit mal and cortical excitability.

Cerebral Dominance and Language has been demonstrated (1) by the effects of lesions of the
left hemisphere that produce deficits in language function
Although right-handedness represents a sensorimotor (aphasia) and (2) by the transient aphasia (inability to
dominance of the left hemisphere and left-handedness speak or write) that results when a short-acting anesthetic
represents a sensorimotor dominance of the right hemi- is introduced into the left carotid artery. Lesions of the
sphere, cerebral dominance is assigned to the hemisphere nondominant hemisphere and injection of anesthetic into
in which language is to communicate; in humans, the left it do not usually affect language substantially.
hemisphere is the dominant hemisphere in more than Several areas in the left hemisphere are involved in lan-
90% of both right- and left-handed people. This dominance guage. Wernickes area is a large area in the posterior part
218 S E C T I O N 2 Berne & Levy Physiology

of the superior temporal gyrus, extending from behind the is presumably transferred between the two hemispheres
auditory cortex into the parietal lobe. Another important through the corpus callosum. This finding can be confirmed
language area, Brocas area, is in the posterior part of by cutting both the optic chiasm and the corpus callosum
the inferior frontal gyrus, close to the face representation before training (see Fig. 10.7C). Then the information is
of the motor cortex. Damage to Wernickes area results not transferred, and each hemisphere must learn the task
in receptive aphasia, in which the person has difficulty independently.
comprehending spoken and written language; however, A similar experiment was conducted in human patients
speech production remains fluent, if meaningless. Con- who had undergone surgical transection of the corpus cal-
versely, a lesion in Brocas area causes expressive aphasia, losum to prevent the interhemispheric spread of epilepsy
in which individuals have difficulty in generating speech (Fig. 10.8). The optic chiasm remained intact, but visual
and writing, although they can understand language information was directed to one or the other hemisphere by
relatively well. the patients fixing vision on the central point of the screen.
The terms sensory aphasia and motor aphasia are often A picture or name of an object was then flashed to one
interchanged with receptive aphasia and expressive aphasia, side of the fixation point, so that visual information about
respectively. The former terms, however, are misleading: A the picture reached only the contralateral hemisphere. An
person with receptive aphasia may not have auditory or opening beneath the screen allowed the patient to manipu-
visual impairment, and one with expressive aphasia may late objects that could not be seen. The objects included
have normal motor control of the muscles responsible for those shown in the projected pictures. Normal individuals
speech or writing. Aphasia does not depend on a deficit of would be able to locate the correct object with either hand.
sensation or of motor skill; rather, it is an inability to decode However, patients with a transected corpus callosum could
language-encoded sensory information into concepts or to locate the correct object only with the hand ipsilateral to
encode concepts into language. However, lesions in the the projected image (contralateral to the hemisphere that
dominant hemisphere may be large enough to result in received the visual information). For the hand to explore
mixed forms of aphasia, as well as sensory changes or and recognize the correct object, the visual information
paralysis of some of the muscles used to express language. must have access to the somatosensory and motor areas of
For example, the latter situation could occur with a lesion the cortex. With the corpus callosum cut, the visual and
of the face representation portion of the motor cortex that motor areas are interconnected only on the same side of
results in an inability to manipulate the motor apparatus the brain.
needed for speaking (vocal cords, jaws, tongue, lips) and Another test was to ask the patient to verbally identify
would be manifest as unclear speech because of dysarthria, what object was seen in the picture. The patient would make
a mechanical deficit. An affected individual would, however, a correct verbal response to a picture that was projected to
be able to write if the motor cortex serving the upper limb the right of the fixation point because the visual informa-
were unaffected. tion reached only the left (language-dominant) hemisphere.
However, the patient could not verbally identify a picture
Interhemispheric Communication and that was presented to the left hemifield because visual
the Corpus Callosum information reached only the right hemisphere.
Similar observations can be made in patients with a
The two cerebral hemispheres can function somewhat transected corpus callosum when different forms of stimuli
independently, as in the control of one hand. However, are used. For example, when such patients are given a verbal
information must be transferred between the hemispheres command to raise the right arm, they do so without dif-
to coordinate activity on the two sides of the body. Much ficulty. The language centers in the left hemisphere send
of that information is transmitted through the corpus cal- signals to the ipsilateral motor areas, and these signals
losum, although some is transmitted through other com- produce the movement of the right arm. However, these
missures (e.g., the anterior commissure or the hippocampal patients cannot respond to a command to raise the left
commissure). arm. The language areas on the left side cannot influence
The importance of the corpus callosum for interhemi- the motor areas on the right unless the corpus callosum is
spheric transfer of information is illustrated in Figure 10.7A. intact. Somatosensory stimuli applied to the right side of
An animal with an intact optic chiasm and corpus callosum the body can be described by patients with a transected
and with the left eye closed learns a visual discrimination corpus callosum, but these patients cannot describe the
task (see Fig. 10.7A). The information is transmitted to same stimuli applied to the left side of the body. Informa-
both hemispheres through bilateral connections made by tion that reaches the right somatosensory areas of the cortex
the optic chiasm or through the corpus callosum, or both. cannot reach the language centers if the corpus callosum
When the animal is tested with the left eye open and the has been cut.
right eye closed (see Fig. 10.7A, center), the task can still In addition to language, other differences in the
be performed because both hemispheres have learned the functional capabilities of the two hemispheres can be
task. If the optic chiasm is transected before the animal is compared by exploring the performance of individuals
trained, the result is the same (see Fig. 10.7B). Information with a transected corpus callosum. Such patients solve
 CHAPTER 10 Integrative Functions of the Nervous System 219

LEARNING TESTING CONCLUSION

Both hemispheres involved Since both hemispheres involved, Interocular transfer due to intact
interocular transfer complete optic chiasm and/or
corpus callosum
A

Optic
chiasm split

LEARNING TESTING CONCLUSION

Right hemisphere trained Left hemisphere knows Learning transferred
problem via corpus callosum
B

Optic
chiasm split

Corpus callosum
section

LEARNING TESTING CONCLUSION

Right hemisphere trained Left hemisphere does Transfer pathway
not know problem was blocked
C
Fig. 10.7 Role of the Corpus Callosum in the Interhemispheric Transfer of Visual Information When
Learning Involves One Eye. A, Discrimination depends on distinguishing between a cross and a circle.
B, Discrimination is between triangles oriented with the apex up or down. C, Discrimination is between
vertical and horizontal bars.
220 S E C T I O N 2 Berne & Levy Physiology

three-dimensional puzzles better with the right than with the

left hemisphere, which suggests that the right hemisphere
has specialized functions for spatial tasks. Other functions
that seem to be more associated with the right than the
Fixation point left hemisphere are facial expression, body language, and
speech intonation (Fig. 10.9). Patients with a transected
corpus callosum lack normal interhemispheric coordina-
tion. When they are dressing, for example, one hand may
button a shirt while the other tries to unbutton it. Observa-
tion of these patients indicates that the two hemispheres
can operate quite independently when they are no longer
interconnected. However, one hemisphere can express
itself with language, whereas the other communicates only
nonverbally.

IN THE CLINIC
One of the more striking examples of interhemispheric
differences is the phenomenon of cortical neglect,
which is a consequence of a lesion in the parietal cortex
of the nondominant (usually right) hemisphere. In such
cases, the patient ignores objects and individuals in the
A left visual field, draws objects that are incomplete on the
left, denies the existence of his or her left arm and leg, and
fails to dress the left side of his or her body. The patient
also denies having any such difficulties (anosognosia).
Although the patient may respond to touch and pinprick
Fixation point
on the left side of the body, he or she cannot identify
objects placed in the left hand. The lesion is adjacent to
the first somatosensory (SI) cortex, as well as the visual
association cortex, and it suggests that this region plays a
special role in the perception of body image and immediate
extrapersonal space. Similar lesions on the dominant side
result only in loss of some higher order somesthesias, such
Key Ring as agraphesthesia (inability to identify characters drawn on
the palm) and astereognosis (inability to identify an object
only by touch).

Speech
Right Left Learning and Memory
hand hand
Major functions of the higher levels of the nervous system
are learning and memory. Learning is a neural mechanism
Ring Key by which the organisms behavior changes as a result of
experience. Memory is the storage mechanism for what is
B learned.
Fig. 10.8 Illustration of Tests in a Patient With a Transected The neural circuitry involved in memory and learning
Corpus Callosum. A, The patient fixes on a point on a rear projec- in mammals is complex and difficult to study. Alternative
tion screen, and pictures are projected to either side of the fixation
point. The hand can palpate objects that correspond to the projected
approaches are animal studies (especially in the simpler
pictures, but these objects cannot be seen. B, Response by the left nervous systems of invertebrates), analysis of the functional
hand to a picture of a key in the left field of view. However, the verbal consequences of lesions, and anatomical/physiological
response is that the patient sees a picture of a ring. (Redrawn from studies at the cellular and pathway level. For example, in
Sperry RW. In: Schmitt FO, Worden FG [eds]. The Neurosciences: the marine mollusk Aplysia, it has been possible to isolate
Third Study Program. Cambridge, MA: MIT Press; 1974.)
a connection between a single sensory neuron and a motor
neuron, which shows aspects of habituation (learning not
to respond to repetitions of an insignificant stimulus),
sensitization (increased responsiveness to innocuous
stimuli that follow the presentation of a strong or noxious
stimulus), and even associative conditioning (learning to
respond to a previously insignificant event after it has been
 CHAPTER 10 Integrative Functions of the Nervous System 221

Visual field

Left Right

R L R L

Left-side olfaction Right-side olfaction

Verbal memory Memory for shapes

Motor speech Stereognosis, left side

Stereognosis, right side Hearing

Right-hand skills (left-ear preference)
(e.g., writing)

Hearing 372 Musical ability

(right-ear preference) x 49
18228
Understanding language Recognition of forms,
faces, and body image
Mathematical ability

Right visual field Left visual field

Left hemisphere Right hemisphere

Fig. 10.9 Schematic illustration of the functional specializations of the left and right hemispheres, as

determined in patients after section of the corpus callosum. (Modified from Siegel A, Sapru HN. Essential
Neuroscience. 5th ed. Philadelphia: Lippincott Williams & Wilkins; 2005.)

paired with a significant one). In the case of habituation, most extensively in the cerebellum (see Chapter 9), but it
the amount of transmitter released in successive responses also occurs in the hippocampus and in other regions of
gradually diminishes. The change involves an alteration in the CNS.
the Ca++ current that triggers release of neurotransmitter. LTP has been studied most intensively in slices of the
The cause of this change is inactivation of presynaptic Ca++ hippocampus in vitro, but it has also been studied in the
channels by repeated action potentials. Long-term habitu- neocortex, cerebellum, and other parts of the CNS. Repeti-
ation can also be produced. In this case, the numbers of tive activation of an afferent pathway to the hippocampus
synaptic endings and active zones in the remaining terminals or repetitive activation of one of the intrinsic connections
decreases. increases the responses of pyramidal cells. The increased
responses (the LTP) last for hours in vitro (and even days
Long-Term Potentiation to weeks in vivo). The forms of LTP differ, depending
Additional models of learning, provided by synaptic on the particular synaptic system. The mechanism of the
phenomena, are called long-term potentiation (LTP) enhanced synaptic efficacy seems to involve both pre-
and long-term depression (LTD). LTD has been studied synaptic and postsynaptic events. The neurotransmitters
222 S E C T I O N 2 Berne & Levy Physiology

involved in LTP include excitatory amino acids that act on

N-methyl-D-aspartate (NMDA) receptors, the responses of AT THE CELLULAR LEVEL
which are associated with an influx of Ca++ into the post- Cellular studies of the hippocampus and the entorhinal cortex
synaptic neuron. Second messenger pathways (including (which is adjacent and parallel to the hippocampus) have
G proteins, Ca++/calmodulin-dependent kinase II, protein demonstrated the existence of place cells that fire when
kinase G, and protein kinase C) are also involved, and these the subject enters a specific place in a test environment.
kinases cause protein phosphorylation and changes in the Although there are many place cells, they are not distributed
in an orderly manner that would resemble a topographic
responsiveness of neurotransmitter receptors. A retrograde map. Place cells appear in very young animals as soon as
messenger, perhaps nitric oxide (or carbon monoxide), may they are able to explore. More recent studies reveal the
be released from postsynaptic neurons to act on presynaptic presence of grid cells, which also respond to specific
endings in such a way that transmitter release is enhanced. sites but are distributed in hexagonal arrays that resembles
Immediate-early genes are also activated during LTP. Hence, an orderly map of the environmental space in the posterior
entorhinal cortex. Although this cognitive map is fixed for
changes in gene expression may also be involved. any context, changes to the environment or removal of the
person to a new test environment causes the generation of a
Memory new and appropriate map of grid cells.
With regard to the stages of memory storage, a distinction Because the entorhinal cortex is a major source of
between short-term memory and long-term memory is input to the hippocampus, it is interesting that studies of
Londons taxi driverswho must demonstrate an extremely
useful. Recent events appear to be stored in short-term detailed knowledge of the citys streets and most efficient
memory by ongoing neural activity because short-term routes before being licensedindicate that the posterior
memory persists for only minutes. Short-term memory is hippocampus in trained and experienced drivers is larger
used, for instance, to remember page numbers in a book than that in beginners or the general population. Getting
after looking them up in the index. Long-term memory lost, a common complaint associated with amnesia, may be
due to the loss of spatial memory skills.
can be subdivided into an intermediate form, which can
be disrupted, and a long-lasting form, which is difficult to
disrupt. Memory loss, or amnesia, can be caused by a loss
of memory information per se, or it can result from interfer-
ence with the mechanism for accessing the information.
Long-term memory probably involves structural changes
because it can remain intact even after events that disrupt some degree of plasticity remains in the adult brain, as
short-term memory. evidenced by responses to certain manipulations, such as
The temporal lobes appear to be particularly important lesions of the brain, sensory deprivation, or even experience.
for memory because bilateral removal of the hippocampal The capability for developmental plasticity may be
formation can severely and permanently disrupt recent maximal for some neural systems at a time referred to
memory. Short-term and long-term memories are unaf- as the critical period. For example, it is possible to alter
fected, but new long-term memories can no longer be some connections formed in the visual pathways during
established. Thus patients with such amnesia remember their development by preventing one eye from providing
events before their surgery but fail to recall new events, input, but only during a specific critical period early in
even with multiple exposures, and must be reintroduced development. In such visually deprived animals, the visual
repeatedly to people they meet after the surgery. This loss connections become abnormal (Fig. 10.10), and restora-
of declarative memory involves the conscious recall of tion of normal visual input after the critical period does
personal events, places, and general history. Such patients, not undo the abnormality, nor does it restore functional
however, can still learn some tasks because they retain pro- vision from the deprived eye. In contrast, similar visual
cedural memory, which involves associational and motor deprivation later in life does not result in abnormal con-
skills. If such patients are given a complex task to perform nections. The plastic changes seen in such experiments may
(e.g., mirror writing), they not only improve during the first reflect a competition for synaptic connections, whereby the
training session but also perform better on subsequent days less functional connections are pruned away. Research has
despite their denial of having any earlier experience with shown a correlation between the form of a gene that modu-
that task. The cerebral structures involved in procedural lates the efficacy of synaptic pruning and the probability of
memory are not yet defined. schizophrenia.
Plastic changes can also occur after injury to the brain in
Neural Plasticity adults. Sprouting of new axons does occur in the damaged
Plasticity most commonly refers to the ability of the CNS to CNS; however, the sprouts do not necessarily restore normal
change its connectivity. Such changes can occur in various function, and many neural pathways do not appear to
contexts, including learning and memory (see previous dis- produce sprouts. Additional knowledge concerning neural
cussion), damage, and development. Damage to the CNS plasticity in the adult CNS is vital if medical therapy is to
can induce remodeling of neural pathways and thereby alter be improved for many diseases of the CNS and after neural
behavior. Plasticity is greatest in the developing brain, but trauma. Research is currently being conducted to explore
 CHAPTER 10 Integrative Functions of the Nervous System 223

1mm

157 L

1mm

B
Fig. 10.10 Plasticity in the Visual Pathway as a Result of Sensory Deprivation During Development.

The ocular dominance columns are demonstrated by autoradiography after injection of a radioactive tracer
into one eye. The tracer is transported to the lateral geniculate nucleus and then transneurally transported
to the striate cortex. The cortex is labeled in bands that alternate with unlabeled bands whose input is
from the uninjected eye. A, Normal pattern. B, Changed pattern in an animal raised with monocular
visual deprivation. The injection was made into the nondeprived eye, and the ocular dominance columns
for this eye were clearly expanded. Other experiments showed that the ocular dominance columns for
the deprived eye contracted. (A, From Hubel DH, Wiesel TN. Proc R Soc Lond B. 1977;198:1. B, From
LeVay S, etal. J Comp Neurol. 1980;191:1.)
224 S E C T I O N 2 Berne & Levy Physiology

the potential of human embryonic stem cells for restoring 5

CNS function. 4
Phantom limb sensation is an example of neural plasticity
in adults. A patient whose limb has been amputated often 3
Right hand
perceives sensations on the missing limb when stimulated
2
elsewhere on the body. Functional imaging studies suggest
that this is a result of the spread of connections from the 1
surrounding cortical territories into the cortical region that
had served the amputated limb.
Central sulcus

IN THE CLINIC Superior

frontal gyrus
It was traditional policy to delay corrective surgery for a child Postcentral
born with a congenital cataract until the child was older and Precentral gyrus
gyrus 5
more able to cope with the stress of surgery. However, if
3
the correction is deferred until after the critical period, full
Precentral 1
recovery of function is unlikely. Similarly, children born with
amblyopia, a condition characterized by strabismus (cross- sulcus
eye) because of relative weakness of one of the extraocular
muscles, tend to use the unaffected eye in preference. In Inferior
both cases, early surgery is now common practice so that frontal gyrus
A Left cortex
the cortical circuitry can be correctly sculpted by balanced
input from the two eyes.

5
4
Such remapping can also occur after surgical amputa-
tion of the second and third digits of the hand. Before Right hand
surgery, each of the digits was represented in discrete and
somatotopically organized areas of the postcentral gyrus (SI
cortex). After surgery, the area that represented the ampu- 1
tated digits is now mapped with an enlarged representation
of the adjacent digits (Fig. 10.11). Conversely, individuals Central sulcus
born with syndactyly (fusion of two or more digits of the
hand) have a single or mostly overlapping representation Postcentral
of these digits in the SI cortex. After corrective surgery, the Precentral gyrus
gyrus
independent digits come to have distinctive representations.
Postcentral
Even more remarkable is that monkeys that were trained on 5 sulcus
a sensory discrimination task requiring repeated daily use 4
of their fingertips showed cortical differences after training. 1 Supramarginal
Not only were the SI cortical territories of their fingertips gyrus
larger than before training but also the number of corti-
Lateral sulcus
cally recorded receptive fields on the fingertips was likewise
increased. B Left cortex
Fig. 10.11 Representation of the digit region of the left first

somatosensory (SI) cortex (A) and reorganization of this representation

(B) after amputation of the second and third digits. (From Haines DE
[ed]. Fundamental Neuroscience for Basic and Clinical Applications.
3rd ed. Philadelphia: Churchill Livingstone; 2006.)

Key Points
1. The cerebral cortex can be divided into lobes on understanding of language, and Brocas area (in
the basis of the pattern of gyri and sulci. Each lobe the inferior frontal lobe) is responsible for its
has distinctive functions, as shown by the effects of expression.
lesions. The left cerebral hemisphere is dominant 2. The neocortex contains pyramidal cells and several
for language in most individuals. Wernickes area (in kinds of interneurons. Specific thalamocortical
the posterior temporal lobe) is responsible for the afferent fibers terminate mainly in layer IV of the
 CHAPTER 10 Integrative Functions of the Nervous System 225

neocortex; diffuse thalamocortical afferent fibers and its circadian rhythmicity is controlled by the
synapse in layers I and VI. Axons from pyramidal suprachiasmatic nucleus.
cells in layer V are the major source of output 6. Information is transferred between the two
to subcortical targets, including the spinal cord, hemispheres primarily through the corpus callosum.
brainstem, striatum, and thalamus. The right hemisphere is more capable than the left
3. The cortical structure varies in different regions. in spatial tasks, facial expression, body language, and
Brodmanns designations reflect these variations in speech intonation. The left hemisphere is specialized
cortical structure and correspond to functionally for the understanding and generation of language, for
discrete areas. logic, and for mathematical computation.
4. The EEG reflects electrical fields generated by the 7. Learning and memory can be studied on the cellular
activity of pyramidal and varies with the state of the level, in invertebrates, and in higher animals.
sleep-wake cycle, disease, and other factors. Cortical Memory includes short-term (lasting minutes),
evoked potentials are stimulus-triggered changes in recent, and long-term storage processes and a retrieval
the EEG and are useful clinical data about sensory mechanism. The hippocampal formation is important
transmission. for storing declarative and spatial memory.
5. EEG patterns during sleep are divided into slow-wave 8. Lesion studies and behavioral studies indicate
and REM forms. Slow-wave sleep progresses through that plasticity occurs in the brain throughout life.
stages 1 through 4, each with a characteristic EEG However, there appears to be more plasticity early in
pattern. Most dreams occur in REM sleep. Sleep life, and synaptic competition in critical periods is
is produced actively by a brainstem mechanism, important for the establishment of neural circuitry.

Additional Reading
Squire L, Berg D. Fundamental Neuroscience. 4th ed. New York:
Academic Press; 2012.