Chapter 2

Functional Neuroanatomy

The way structures in the developing brain are Watson, & Ojemann, 2005), to investigate dendritic
related to changes in psychological and cognitive morphology with electron microscopic techniques
development is of interest to child neuropsycholo- (Scheibel, 1990), to measure sequential brain pro-
gists. There are several ways that this relationship cessing during cognitive tasks using visual evoked
can be explored, including: (1) correlating structural potentials (Liotti et al., 2007), and to image the
changes in the developing brain with behavioral brain while a person is completing a task through
changes, (2) investigating behavioral changes and functional magnetic resonance imaging (Pliszka
making inferences about structural maturation of et al. 2006).
the brain, and (3) studying brain dysfunction and Our basic understanding of the brain and its
its relationship to behavioral disorders (Kolb & relationship to complex human behaviors has
Fantie, 1989). been greatly facilitated by technological advances
Although these approaches can yield useful in modern neuroimaging techniques, including
information about the developing brain, they are computed tomography (CT), magnetic resonance
not without shortcomings. For example, because imaging (MRI), regional blood flow (rCBF), and
of the plasticity of the developing brain following positron-emission tomography (PET). Neuroima-
damage, injury in a specific brain region may pro- ging techniques allow researchers to gather direct
duce behavioral losses that vary greatly depending evidence linking cognitive, behavioral, and psy-
on the age of the child. Environmental factors, chosocial disorders to anatomical, physiological,
such as enrichment opportunities and social-cultural and biochemical processes in the brain (Semrud-
experiences, also influence the developing brain Clikeman, 2007). Research findings about the
and the manner in which behaviors are expressed developing brain from these various approaches
(Baron, 2004). Thus, the study of the brain-behavior and methodologies will be used throughout this
relationship is particularly complex in children, and chapter in an effort to explore the biological basis
these factors must enter the equation when drawing of childhood disorders. These techniques will be
conclusions about this relationship. Some have further discussed in Chapter 3. To fully appreciate
criticized neuropsychological approaches because the brain-behavior relationship in children, an
of the level of inferences made when relating overview of the structure and function of the
behavior to brain structure and function, and brain is necessary. This chapter reviews the structures
because of the correlational nature of the research and functions of the neuron and the sub-cortical
(Fletcher & Taylor, 1984). There are now medical and cortical regions from a neurodevelopmental
technologies and new research protocols that perspective. This review serves as a foundation for
avoid some of these shortcomings. These technolo- exploring the complex interaction between anatomi-
gies make it possible to explore the brain during cal development of the brain and the emergence
craniotomies under local anesthesia (McDermott, of childhood behaviors and disorders.

M. Semrud-Clikeman, P.A. Teeter Ellison, Child Neuropsychology, DOI 10.1007/978-0-387-88963-4_2, 25

Springer ScienceBusiness Media, LLC 2009
26 2 Functional Neuroanatomy

Structure and Function of the Neuron or nuclei that have special behavioral functions.
Neurons can be modified through experience, and
The neuron, the basic cellular structure of the they are said to learn, to remember, and to forget as
nervous system, transmits nerve impulses through- a result of experiences (Hinton, 1993). Pathological
out a complex network of interconnecting brain changes in neurons can occur as a result of early
cells. The brain contains approximately 180 billion abnormal experiences. Although these alterations
cells, 50 billion of which transmit and receive are thought to have a profound effect on the mature
sensory-motor signals in the central nervous system organism, the exact nature of these changes is still
(CNS) via 15,000 direct physical connections under investigation. Genetic aberrations also play a
(Carlson, 2007). Investigation of the structure and role in the way neurons develop and function (Cody
function of neurons and their synaptic connections et al. 2005). Damage to or destruction of neurons is
provides insight into basic psychopharmacology at also of concern because neurons typically do not
the molecular level and may provide a method for regenerate (Swaiman, Ashwal, & Ferriero, 2006).
describing how various neuropsychiatric disorders Neurodevelopmental disorders and issues related
emerge and progress (Pliszka, 2003). to recovery of function following brain trauma
The CNS is comprised of two major cell types, will be discussed in detail in later chapters (see
neurons and neuroglia (Carlson, 2007). While neu- Chapter 10).
rons conduct nerve impulses, the neuroglia (nerve
glue) provide structural support and insulate
synapses (the connections between neurons). Glial
cells make up about 50 percent of the total volume Anatomy of the Neuron
of the CNS. Glial cells serve various functions,
including transmission of signals across neurons, The neuron contains four well-defined cellular
structural support for neurons, repair of injured neu- parts, including the cell body, dendrites, axons,
rons, and production of CNS fluid (Carlson, 2007). and axon terminals. The cell body, or soma, is the
Neuroglia infiltrate or invade surrounding tissue in trophic or life center of the neuron (see Fig. 2.1).
both the gray and white matter, and in rare instances Cell bodies vary in size and shape and contain the
these cells replicate uncontrollably during tumor ribonucleic acid (RNA) and deoxyribonucleic acid
activity (Nortz, Hemme-Phillips, & Ris, 2007). (DNA) of the neuron. RNA, the site of protein
Though still relatively infrequent, pediatric brain synthesis, transmits instructions from DNA direct-
tumors are the second most common neoplasm in ing the metabolic functions of the neuron. Biochem-
children under 15 years of age, and as many as ical processes of the neuron, which take place in the
1,0001,500 cases are estimated to occur each year cytoplasm of the cell body, include the energy-pro-
(Sklar, 2002). ducing functions, the self-reproducing functions,
Gray matter is located in the core of the CNS, and the oxidating reactions, whereby energy is
the corpus striata at the base of the right and left made available for the metabolic activities of the
hemispheres, the cortex that covers each hemi- cell (Carlson, 2007). Destruction or damage to the
sphere, and the cerebellum (Carlson, 2007). The cell body can result in the death of the neuron.
cell bodies, the neuroglia, and the blood vessels Dendrites branch off the cell body and receive
that enervate the CNS are gray-brown in color impulses from other neurons (Carlson, 2007).
and constitute the gray matter. White matter covers Dendrites are afferent in nature and conduct
the gray matter and long axons extending out from nerve impulses toward the cell body. Dendritic
the neuron. Axons are generally covered by a myelin spines are the major point of the synapse, the area
sheath, which contains considerable amounts of of transmission from one cell to another. Indivi-
neuroglia and appears white upon inspection. duals with cognitive retardation have fewer spines
White matter has fewer capillaries than gray matter or points of contact across neurons (Klein-
(Carlson, 2007). Tasman, Phillips, & Kelderman, 2007). Dendrites
As the basic functional unit of the CNS, the neu- can transmit neuronal impulses across neurons
ron transmits impulses in aggregated communities through either temporal or graded potentials. In
Structure and Function of the Neuron 27

Soma
(cell body) Dendrites

Terminal
buttons

Axon

Myelin sheath

Messages

Fig. 2.1 Anatomy of the Neuron

Source: From Neil R. Carlson, Physiology of Behavior, 5th edition, p. 21. Copyright # 1994 by Allyn and Bacon. Reprinted by
permission

this case, as a neuron receives an impulse it can Axons are covered by a myelin sheath made up of
transmit this impulse if the stimulation is close in neurilemma (or Schwann cells), which surround the
time to another impulse or if it is strong enough axon. The myelin sheath gives the axon a white
combined with a previous impulse. appearance and constitutes most of the white matter
The axon is a long projection or axis from the cell in cortical and subcortical areas. Most axons are
body. Most neurons have only one axon, usually myelinated at birth particularly in areas necessary
efferent in nature, that conducts nerve impulses for survival (motor-sucking; tactile sensitivity to
away from the cell body. Axons are typically longer hot, cold, and pain; auditory, and vision). Some
than dendrites and can be as much as one yard in axons continue to myelinate throughout develop-
length. For example, giant pyramidal cells in the ment with myelinization not complete in the frontal
motor cortex send axons to the caudal tip of the lobes until well into the third decade of life. Changes
spinal cord. The axon hillock is a slender process in postnatal brain weight are generally related to
close to the cell body where action potentials arise. increases in dendritic connections and to increases
The axon hillock is highly excitable and is activated in the number of glial cells that form the myelin
through electrochemical processes, thereby turn- sheath along the axon (Shepherd, 2004).
ing on the neuron (Carlson, 2007). The impulse Axons allow the nerve cells to transmit impulses
must be of sufficient strength for the neuron to rapidly, particularly along the Nodes of Ranvier.
fire. Axons follow an all or nothing rule; if the The Nodes of Ranvier are gaps in the myelin and
impulse is not strong enough the neuron will not fire during cell activation, nerve impulses skip from node
and, thus, will not transmit the message to another to node. Myelinated axons permit more rapid trans-
neuron. After the neuron fires, there is a period of mission of signals, and anesthetics seem to be more
time when it will not fire again as the neuron effective at the Nodes of Ranvier. The terminal
recovers. branches of the axon end at the synaptic telodendria.
28 2 Functional Neuroanatomy

The presynaptic and postsynaptic sites are both Types of Neurons

referred to as the synapse. Synapses are specialized
for the release of chemicals known as neurotrans- There are two basic types of neurons: efferent and
mitters. Neurotransmitters are released from synap- afferent. Efferent neurons originate in the motor
tic knobs at the end foot of the neuron in cortex of the CNS, descend through vertical
the presynapse, and they activate neurons at the pathways into subcortical regions, and culminate
postsynapse. Neurotransmitters are released from in the bodys muscles (Gazzaniga, Ivry, &
the presynapse (neuron A), travel across the synap- Mangun, 2002). These large descending tracts
tic cleft, and influence the activity of the adjoining form columns from the motor cortex connecting
neuron (neuron B) (see Fig. 2.2 for a depiction of higher cortical regions through the brain stem and
these activities). There is a collection of vesicles at spinal cord, to the body for the activation of single
the synaptic knob at the end of each synapse, where muscles or muscle groups. Various motor path-
neurotransmitters are stored. Most neurons have ways begin to develop prenatally, while postnatal
thousands of synapses, and each dendritic spine development is marked by changes in primitive
serves as a synapse that is excitatory in nature, reflexes (the Babinski reflex) and automatic
which causes neurons to fire. Synapses are quite reflexes (head and neck righting) (Swaiman et al.,
large for motor neurons and are smaller in the 2006).
cerebellum and other cortical regions. Synapses Afferent neurons, sensory receptors found
usually occur between the axon of one cell and the throughout the body, transmit sensory information
dendrite of another (axondendritic connections). into specific cerebral areas. For example, afferent
Although they can connect onto the soma or cell neurons consist of rods and cones (cells that convey
body of another neuron (axosomatic connection), information about color or black/white) in the visual
synapses rarely occur from axon to axon (axo- system that project into the occipital cortex; hair cells
axonal connections). (convey information about tone) in the auditory

Axons from other neurons

their terminal buttons NEURON B
influence neuron A Synapse

Axons from
other neurons

NEURON A
Messages sent down axon
influence neuron B and
other neurons
To other
neurons

Soma

Dendrite

Fig. 2.2 Anatomy Showing Connections between Neuron A and B with Synaptic Cleft
Source: From Neil R. Carlson, Physiology of Behavior, 5th edition, p. 23. Copyright # 1994 by Allyn and Bacon. Reprinted
with permission
Spinal Cord 29

system that project into the temporal cortex, and Astrocytomas in childhood most frequently
pain, touch, temperature, and pressure sensors in occur in the cerebellum and the brain stem. These
the skin that project into the parietal cortex. Somes- tumors are found equally in males and females.
thetic senses are the first to become functional in the Although astrocytomas can occur at any age, the
fetus, as early as 78 weeks gestation, while auditory most frequent incidence is between five and nine
and visual neural maturation occurs later in embryo- years of age (Hunter et al., 2005). Oligodendroglia
nic development (Gazzaniga et al., 2002). Other cells cells form and maintain the myelin sheath and,
in the corpus callosum (a large bundle of fibers con- when injured, swell in size. Tumors rarely occur in
necting the two hemispheres) and the frontal lobes do oligodendroglia cells; when they do they grow
not become fully functional until the teenage years slowly and are found primarily in the cortex and
through the 20s. white matter. While about 4060 percent of these
tumors can be detected by skull X-rays after they
calcify (Cohen & Duggner, 1994), radionuclide
brain scans, angiography, and computed tomogra-
Types of Neuroglia phy scans have been helpful in the diagnostic phase
of tumor processes. Finally, microglia cells are
predominantly found in the gray matter (Carlson,
The neuroglia cells serve a number of important 2007). Following disease or injury, microglia prolif-
functions in the CNS: (1) providing structural erate, move to the site of injury, and perform a
support to neurons; (2) aiding in the regeneration phagocytic function by cleaning up damaged tissue.
of injured nerve fibers; (3) occupying injured sites by Tumors rarely occur in microglia cells.
producing scar tissue, and (4) transporting gas, These cells develop at different rates depending
water, and metabolites from blood, and removing on location in the brain, experience of the baby,
wastes from nerve cells (Carlson, 2007). The three and genetic programming. In order to understand
major types of neuroglia (astrocytes, oligodendro- difficulties children have in development it is first
glia, and microglia) have distinct functions and important to understand how a typical brain devel-
serve multiple purposes in the CNS. Astrocytes ops. The following section provides a brief overview
have three primary functions: (1) forming the of the course of neuronal development.
blood-brain barrier; (2) supporting the cellular
structure of the brain, and (3) directing the migration
of neurons during early development. Astrocytes are
the largest in size and the most abundant type of Spinal Cord
neuroglia (Carlson, 2007). These star-shaped glial
cells attach to capillary blood vessels and cover The spinal cord serves two major functions: connecting
approximately 80 percent of each capillary. Astrocytes, the brain and the body via large sensory and motor
found primarily in the pia matter (fine membrane on neurons. The spinal cord comprises gray matter
the surface of the brain), cover large blood vessels. and white matter. Gray matter is the central, inter-
When injury occurs to the spinal cord or to the brain, ior region of the spinal cord and is shaped like a
through either disease or trauma, astrocytes go into butterfly. It appears gray on inspection and is made
hypertrophy (Morris, Krawiecki, Kullgren, Ingram, & up of cell bodies. Neurons leave the spinal cord in
Kurczynski, 2000). These cells multiply quickly, form- segments called dermatomes and enter into muscles
ing a glial scar that fills in gaps in the cellular structure and organs. Motor commands from higher cortical
caused by injury. Astrocytes may also serve a phago- centers are conducted at these sites. Sensory recep-
cytic function by removing destroyed tissue and clean- tors connect with motor neurons in the gray matter
ing up the site of injury. Astrocytoma, a type of of the spinal cord, via interneurons. Interneurons
primary neoplasm that frequently reoccurs after remain in the spinal cord and mediate motor activity
surgery, is the second most common brain tumor in with sensory stimuli. Interneurons also provide for
adults (Hunter et al., 2005); though rare, astrocytomas cooperation among different spinal segments, which
do occur in children as well. control distant muscle groups. For example,
30 2 Functional Neuroanatomy

interneurons connect cervical and lumbar regions Table 2.1 Cranial nerves
of the spinal cord to coordinate forelimbs and legs Number Name Function
for walking. White matter surrounds the gray matter I Olfactory Smell
and consists of the myelin sheath (Brodal, 2004). II Optic Vision
The spinal cord conducts signals to and from III Oculomotor Eye movement
IV Trochlear Eye movement
higher cortical regions, including the brain stem,
V Trigeminal Masticatory movement
the cerebellum, and the cortex. The posterior root VI Abducens Eye movement
of the spinal cord is afferent in nature, where sen- VII Facial Face movement
sory fibers enter into the gray matter, synapse with VIII Auditory Hearing
other neurons, and ascend into higher cortical areas IX Glossopharyngeal Tongue and pharynx
in pathways. Conversely, the anterior root is effer- movement
ent in nature and is made up of motor fibers that X Vagus Heart, blood vessels,
viscera, larynx, and
receive motor signals from higher cortical areas and pharynx movement
communicate to muscle groups for movement. XI Spinal Accessory Strength of neck and
Nerve fibers enter and leave the spinal cord at reg- shoulder muscles
ular intervals (dermatomes) and provide sensory XII Hypoglossal Tongue muscles
and motor innervation to specific body segments.
There are a total of 30 segments innervating the
spinal cord: eight cervical, 12 thoracic, five lumbar, Role and Function of the Meninges
and five sacral (Brodal, 2004). Damage to the spinal
cord at specific sites produces localized sensory and Both the spinal cord and the brain are surrounded
motor dysfunction in the body. by a protective layer of tissue called the meninges.
Unlike the brain, the spinal cord has little diver- The meninges comprise three layers: the dura mater,
sification or specialization, but it does carry out the arachnoid, and the pia mater. The dura mater is
sensory, motor, and integrative functions. Four the tough outer layer of the spinal cord and the
such functions are carried out in the spinal cord: brain, and has the consistency of a thin rubber
(1) reflex activity, whereby a stimulus is followed glove. The dura mater attaches to the bones cover-
by a coordinated motor response; (2) reciprocal ing the cranium and receives blood vessels that
activity, whereby one activity starts or stops another innervate the brain (Brodal, 2004). Head injury
(i.e., excitatory or inhibitory); (3) monitoring activity, may form an epidural hematoma, causing blood to
whereby incoming messages are controlled, coded, accumulate in the region between the skull and the
and transmitted, and (4) transmission activity, whereby dura mater. The dura mater is supplied with blood
messages are transmitted to and from the brain by tiny vessels on its outermost layer near the skull.
through the white matter (Kolb & Whishaw, 2003). The subdural space, a fluid-filled layer, separates
In summary, the spinal cord is one of two major divi- the dura mater from the arachnoid space. Accumu-
sions of the CNS; the second is the brain. lation of blood in the subdural area following injury
can put enormous pressure on the brain (Swaiman
et al., 2006). The arachnoid, a spiderlike web, is a
Structure and Function of the Brain delicate network of tissue under the dura mater.
Blood accumulation between the dura mater and
The nervous system is divided into two basic the arachnoid following injury is referred to as a
systems: the peripheral (PNS) and the central subdural hematoma. Finally, the pia mater is the
nervous system (CNS). The PNS consists of the fragile, innermost layer of the meninges and con-
spinal, cranial, and peripheral nerves that connect tains small blood vessels. The pia mater surrounds
the CNS to the rest of the body. Table 2.1 lists the the arteries and veins that supply blood to the brain;
cranial nerves and their functions. The CNS is it serves as a barrier keeping out harmful substances
completely encased in bone, is surrounded by pro- that might invade the brain.
tective coverings (meninges), and consists of two Bilateral infections that attack the meninges,
major structures: (1) the spinal cord in the vertebral referred to as meningitis, can have serious conse-
column, and (2) the brain within the skull. quences for the developing brain (Swaiman et al.,
Role and Function of the Meninges 31

2006). The first year of life is the time of greatest risk have a devastating affect on the developing brain
for meningitis. The earlier the infection occurs, the and may cause cognitive delays, particularly for non-
higher the mortality rate. Some of the long-term verbal information; emotional, psychiatric, or beha-
consequences of meningitis are mental retardation, vioral disturbances, and slow motor development
hydrocephalus, seizures, deafness, and hyperactivity (Fletcher, Dennis, & Northrup, 2000). Surgical
(Swaiman et al., 2006). Cerebrospinal fluid (CSF), a shunting drains CSF outside the skull. Recent
clear, colorless fluid, fills the ventricles and the sub- advances in microsurgery in utero have produced
arachnoid space (Wilkinson, 1986). CSF contains successful results by reducing some of the more
concentrations of sodium, chloride, and magnesium, severe long-term negative effects of brain dysfunc-
as well as levels of neurotransmitters and other tion or damage that can occur when hydrocephalus
agents. An assay of the composition of these chemi- is untreated. Residual effects of hydrocephaly, ran-
cals can be important for diagnosing disease pro- ging from mild to severe, depend on individual
cesses. CSF reproduces at such a rate that total variables including the age of the child at the time
replacement occurs several times a day. The choroid of shunting and the presence of other neurological
plexus, located in the floor of the ventricles, produces or medical complications that often accompany
the CSF, while the lateral ventricles contain the high- this disorder (Fletcher et al., 2000).
est amounts of CSF. Infectious and metabolic dis-
orders, such as meningitis, encephalitis, and tumors,
as well as traumatic injury, can cause discernible
changes in the CSF. Ventricles
Cerebrospinal fluid has three major functions.
Specifically, it (1) protects against injury to the brain The ventricles, large cavities filled with cerebrospinal
and spinal cord; (2) diffuses materials into and away fluid (CSF), reside in various regions of the brain.
from the brain, and (3) maintains a special environ- The fourth ventricle, also referred to as the aqueduct
ment for brain tissues. Interference in the circulation of Sylvius, resides in the brain stem at the level of the
and drainage of CSF can result in hydrocephalus, pons and the medulla. The third ventricle is located
which causes cranial pressure. Hydrocephalus can in the diencephalon, and the lateral ventricles

THALAMUS

CORPUS CALLOSUM CENTRAL SULCUS

TAL
ON
FR
R
IO
R
PE

GYRUS
GULATE
CIN
SU

SP
LE
NIU
GE M
NU

L
ORBITA
GYRUS

HYPOTHALAMUS
CEREBELLUM
MIDBRAIN
PONS MEDULLA

Fig. 2.3 Sagittal Section of the Brain Showing Brain Stem, in D. Saklofske (Ed.), International Handbook of Personality
Midbrain, and Forebrain Structure and Intelligence in Clinical Psychology and Neuropsychology,
Source: Adapted from M. Semrud-Clikeman and P. A. copyright # 1995 by Plenum Press, New York
Teeter, Personality, Intelligence, and Neuropsychology,
32 2 Functional Neuroanatomy

are found in the forebrain region (see Fig. 2.3). Ven- Structure and Function of the Brain Stem
tricles provide equilibrium as well as the CSF
transporting nutrients and wastes throughout the The brain stem comprises five areas, including the
brain. When these ventricles appear enlarged, a fourth ventricle, the medulla oblongata, the pons
diagnosis of a tumor or disease processes, includ- (bridge), the midbrain (mesencephalon), and the
ing hydrocephalus, encephalitis, and meningitis, diencephalon. Figure 2.4 shows a schematic of
may be made. these structures and Fig. 2.5 shows a magnetic

Fig. 2.4 MRI Sagittal Section of CNS Analogous to Brain Areas Depicted in Figure 2.3

CORPUS CALLOSUIM
LATERAL VENTRICLES

CAUDATE NUCLEUS
BASAL GANGLIA
BASAL GANGLIA
INSULA
INTERNAL CAPSULE
SUBSTANTIA NIGRA

PUTAMEN RED NUCLEUS

GLOBUS PALLIDUS
MESENCEPHALON
THALAMUS
SUPERIOR COLLICULUS
LATERAL VENTRICLE

LATERAL VENTRICLE

THIRD
VENTRICLE

Fig. 2.5 Coronal Section Showing Structures of the Right and Left Hemisphere with Ventricular Systems
Structure and Function of the Brain Stem 33

resonance image of these same structures. The are necessary for adequate direction and maintenance
major regions of the brain stem are discussed in of attention. When enough information reaches the
detail in the following sections. RAS, it signals the cortex and produces cortical arou-
sal and wakefulness. Thus, in children with ADHD
this subcortical filter may not allow sufficient stimuli
to reach higher cortical regions. This theory and
Medulla Oblongata others will be explored in later chapters.
Secretion of serotonin takes place at the pons,
The medulla is a continuation of the spinal cord and probably in the raphe system. The raphe nuclei are
contains nerve tracts similar to those found in the cells located across the medulla, pons, and mid-
spinal cord. Groups of sensory and motor nuclei are brain regions, with afferent connections to the
arranged in ascending (i.e., afferent-sensory tracts) hypothalamus and limbic system (Brodal, 2004).
or descending (i.e., efferent-motor tracts) cell col- This region also contains the locus ceruleus (LC),
umns. Projections of the major cranial nerves occur which produces 70 percent of norepinephrine in the
at the level of the medulla, including the hypoglossal brain, and serves as a modulator for other neuro-
(tongue), the glossopharyngeal (pharynx and transmitters (Carlson, 2007). The norepinephrine-
larynx), and the accessory (neck muscles) nerves. rich cells in the locus ceruleus connect with the
The sensory and motor tracts cross over into the serotonin-rich cells in the raphe nuclei, and each
opposite side of the brain at the level of the medulla. type has a reciprocal affect on the other. Norepi-
The somatosensory (touch, pressure, pain, and nephrine plays a role in vigilance, arousal, filtra-
temperature) and the motor systems are organized tion of stimuli, and habituation. Finally, the con-
in contralateral fashion, such that sensory informa- tinuation of the RAS at the pontine level appears
tion and movement on the right side of the body to mediate sleep.
are primarily controlled by the left hemisphere. Con- Serotonin inhibits arousal of the RAS, which
versely, the left side of the body is controlled by the then allows the thalamus to bring the cortex to a
right hemisphere. The auditory and visual systems slow-wave sleep state (Carlson, 2007). Anesthetics
also cross in the medulla. These functional systems appear to depress the RAS, which ultimately
will be discussed in more detail later in this chapter. depresses the cortex. Fibers in the RAS also project
The reticular activating system (RAS) comprises to the limbic system and serve behavioral and
a major portion of the medulla, extends into the emotional mechanisms for the control of pain.
midbrain region, and has numerous connections Morphine and opiate-like drugs may produce
and functions (Brodal, 2004). The RAS, considered analgesic actions most likely in the raphe system
the arousal system, plays an important role in main- (Shepherd, 2004).
taining consciousness and attentional states for the
entire brain. The RAS has been hypothesized as one
of the critical mechanisms involved in ADHD
(Sagvolden & Archer, 1989). For example, some Pons
RAS functions control blood pressure, blood
volume in organs, and heart rate, whereas others The pons, between the medulla and midbrain and
regulate sleep and wakefulness. above the cerebellum, serves as a bridge across the
The RAS receives input from most sensory systems right and the left hemispheres. Major sensory and
and connects to all levels of the CNS. Because the motor pathways move through the pons, a conti-
RAS is directly or indirectly connected to much of the nuation from the spinal cord and brainstem regions,
CNS, it can modulate CNS activity. Selective stimuli and enter into higher cortical areas. The pons, in
activate the RAS, which then alerts the cortex to coordination with the cerebellum, receives informa-
incoming stimuli. Researchers espousing a bottom- tion concerning movements from the motor cortex
up model hypothesize that the RAS may be filtering and helps modulate movements (Brodal, 2004).
too much sensory information, thereby not allowing Information from the visual cortex is also received
stimulation to reach the higher cortical regions that at the pontine level, which serves to guide visually
34 2 Functional Neuroanatomy

determined movements. Finally, information from thalamus); (2) the auditory system (projecting
the hypothalamus and the limbic system converge in into the medial geniculate body), and (3) sensory
the pons and may influence the impact of emotional receptors in the skin for pain, pressure, touch,
and motivational factors on motor activity (Brodal, and temperature.
2004). A number of cranial nerves converge in the The hypothalamus, anterior and inferior to the
pontine region. Cranial nerves innervating the face thalamus, plays a role in controlling the autonomic
and head receive sensory information and transmit nervous system, including eating, sexual functions
signals in the pons for swallowing and chewing and dysfunctions, drinking, sleeping, temperature,
(trigeminal nerve), moving facial muscles, and rage, and violence. With connections to the limbic
affecting the hearing and equilibrium in the inner system, the hypothalamus influences motivational
ear. Cranial nerves innervating the eye muscles mechanisms of behavior. The pituitary, following
(abducens) also pass through the pons. directions from the hypothalamus, secretes hormones
that regulate bodily functions. The internal capsule,
situated lateral to the thalamus, contains fibers
connecting the cortex to lower brain regions including
Midbrain the brainstem and the spinal cord. Major fibers com-
prise the internal capsule and connect the frontal
The most anterior region of the brainstem is the cortical regions to the thalamus and to the pons.
midbrain or mesencephalon. The midbrain serves Finally, the optic nerve converges in the diencephalon
a major relay function for sensory-motor fibers. The and forms the optic chiasma (Brodal, 2004). Fibers
two major divisions in the midbrain are the tegmen- from the optic nerve cross at the chiasma and project
tum, which falls below the ventricle and is separated to the lateral geniculate body in the thalamus via the
by the substantia nigra, and the tectum, which com- optic tract (Brodal, 2004). Figure 2.6 shows these
prises the superior colliculi (upper region involved structures.
in vision) and the inferior colliculi (lower region
involved in the integration of auditory and kines-
thetic impulses). The RAS also continues into the
midbrain region. Several cranial nerves are located Cerebellum
in the midbrain region. The oculomotor nerve
moves the eye (lateral and downward gaze), and
regulates the size of the pupil and the shape of the The cerebellum or hindbrain, behind the brain stem,
lens. The trigeminal nerve also resides in the mid- connects to the midbrain, pons, and medulla. The
brain area and serves as the major sensory nerve of cerebellum receives sensory information about
the face. where the limbs are in space and signals where
muscles should be positioned. The cerebellum
receives information from the semicircular canals
(in the inner ear) concerning orientation in space.
Diencephalon The cerebellum is involved in the unconscious
adjustment of muscles in the body for coordinated,
The diencephalon, the superior region of the smooth, and complex motor activity. Injury of
brain stem, contains major relay and integrative the cerebellum can result in dystaxia (movement
centers for all the sensory systems except smell. disorders), dysarthria (slurred speech), nystagmus
The diencephalon is not clearly demarcated, but (blurred vision and dizziness), and hypotonia
includes the thalamus, the hypothalamus, the (loss of muscle tone) (Swaiman et al., 2006). Though
pituitary gland, the internal capsule, the third still relatively uncommon, subtentorial tumors invol-
ventricle, and the optic nerve (Brodal, 2004). ving the cerebellum and the fourth ventricle are the
The thalamus receives input from several sensory most frequent type of brain tumor affecting young
sources, including: (1) the visual system (project- children (Konczak, Schoch, Dimitrova, Gizewski, &
ing into the lateral geniculate body of the Timmann, 2005).
Cerebral Hemispheres 35

CENTRAL SULCUS
FRONTAL LOBE S PARIETAL LOBE

U
YR
L
NTA

L
LG

A
SU

GY NTR
PA PE

T RA
MIDDLE FRO

S
RI RIO

RU
CE
ET R

CEN
AL

SUPERIOR
FRONTAL

ST
INFERIOR
FRONTAL

PO
PRE

RU R
GY LA
S
GU
MPORAL

AN
SUPERIOR TE
ORBITAL
GYRI
MIDDLE TEMPORAL
SYLVIAN FISSURE
OR TEMPORAL
INFERI
OCCIPITAL LOBE
TEMPORAL LOBE

Fig. 2.6 Surface of the Left Hemisphere Showing Sulci, D. Saklofske (Ed.), International handbook of Personality and
Fissures, and Major Subdivisions of the Cortex Intelligence in Clinical Psychology and Neuropsychology,
Source: Adapted from M. Semrud-Clikeman and P. A. copyright # 1995 by Plenum Press, New York
Teeter, Personality, Intelligence and Neuropsychology, in

Structure and Function of the Forebrain brain tumors and head trauma (Chapter 10) and in
the discussion of psychopharmacology (Chapter 11).
Neocortex In the following sections, the basic divisions of the
nervous system will be explored.
The neocortex, often referred to simply as the
cortex, comprises the highest functional division of
the forebrain and makes up about 80 percent of the
human brain. The cortex is wrinkled in appearance, Cerebral Hemispheres
with various elevated ridges and convolutions.
Ridges are referred to as gyri, the deepest indenta- The cerebrum comprises the right and left hemi-
tions are called fissures, and the shallower indenta- spheres, which appear to have anatomical (asymme-
tions are called sulci. The configuration of fissures try) as well as functional (lateralization) differences
and large sulci can be identified on visual inspection (Brodal, 2004). Asymmetry typically refers to the struc-
of the cortex (see Fig. 2.6). The lateral or Sylvian tural or morphological differences between the two
fissure separates the frontal lobe from the temporal hemispheres (Rosen, Galaburda, & Sherman, 1990).
lobe, and the central sulcus (fissure of Rolando) Although neuroanatomical differences may underlie
separates the frontal from the parietal lobe. The behavioral variations documented for each hemi-
central sulcus is a prominent landmark separating sphere, it is not known whether chemical as well as
the motor cortex (anterior to the central sulcus) structural differences between the hemispheres also
from the sensory cortex (posterior to the central account for functional asymmetries (Witelson &
sulcus). The surface areas of posterior temporal Kigar, 1988). Cerebral lateralization refers to the
and parietal locations are not clearly defined degree to which each hemisphere is specialized for
from the occipital regions. Finally, the calcarine processing specific tasks. The right and left hemi-
sulcus extends from the occipital pole below to the spheres appear to differ in terms of their efficiency
splenium of the corpus callosum. The following sec- in processing certain stimuli, such that both hemi-
tions will describe the structures and functions of the spheres are not equally good at all tasks (Brodal,
cortex. This brief overview of the structure, function, 2004). Goldberg and Costa (1981) indicate that
and development of neurons serves as a foundation significant cytoarchitectural differences exist between
for understanding the basic structure of the CNS and the two hemispheres that may be related to neurobe-
will be explored in more detail in a discussion of havioral differences. The left hemisphere has a greater
36 2 Functional Neuroanatomy

ratio of gray matter to white matter, particularly in the be related to the difficulty in encoding letters and
frontal, parietal, and temporal regions, compared to words (Galaburda, Sherman, Rosen, Aboitiz, &
the right hemisphere. Conversely, the right hemi- Geschwind, 1985; Hynd, Semrud-Clikeman, Lorys,
sphere has greater white-to-gray matter ratios than Novey, & Eliopulos, 1990; Larsen, Hoeien, &
the left hemisphere. Oedegaard, 1992).
Major anatomical and functional differences Early accounts of cerebral lateralization often
observed in the two hemispheres are described as listed specific functions for each hemisphere in a
follows: dichotomous, all-or-nothing fashion, implying that
all aspects of a given task were carried out by one
1. The left hemisphere has more neuronal represen-
hemisphere. This all-or-nothing approach is prob-
tations in modality-specific regions in the three
ably overly simplistic because both hemispheres
sensory cortices.
generally play a role in most complex tasks. One
2. The right hemisphere has greater association
hemisphere, however, is usually considered domi-
zones, where sensory modalities converge.
nant or most important for a specific task, while the
3. The left hemisphere is structurally conducive
other hemisphere is recessive or nondominant.
to single modality processing, distinct motor
Table 2.2 summarizes the developmental milestones
activity, and intraregional integration.
for anatomical and functional asymmetries.
4. The right hemisphere is structurally conducive to
Witelson (1990) suggests that it is unclear whether
multiple modality and intraregional integration.
functional differences between the two hemispheres are
5. The right hemisphere has a greater capacity for
relative or absolute, in such a way that each hemi-
handling informational complexity because of
sphere is able to process tasks, but does so less effi-
its intraregional connections, whereas the left
ciently. Others have proposed that the two hemispheres
hemisphere seems best suited for processing
operate in a domain-specific fashion, whereby each
unimodal stimuli.
hemisphere acts in an autonomous manner with
The right hemisphere appears better able to pro- restricted access to information processed by the
cess novel information, whereas the left hemisphere other hemisphere.
seems able to work more efficiently with information
with preexisting codes, such as those found in lan-
Table 2.2 Major Division of the Nervous System
guage activities. These differences will be further Functional
explored in a later discussion regarding nonverbal Brain Divisions Brain Structures Divisions
learning disabilities. Although the correlations Telencephalon Neocortex Forebrain
between structure and function are not perfect, (endbrain) Basal ganglia
cerebral asymmetry has been of great interest to Limbic system
child neuropsychologists (Baron, 2004). Further, Olfactory bulb
particular anatomical asymmetries between the two Lateral ventricles
hemispheres are present at birth (Kolb & Whishaw, Diencephalon Thalamus
(between-brain) Epithalamus
2003). Measurable differences have been observed in
Hypothalamns
the left planum temporale (near the auditory cortex) Pineal gland
by 39 weeks gestation, leading some to suggest that Third ventricle
the functional lateralization of language in the left Mesencephalon Tectum Brain stem
hemisphere is determined prenatally (Witelson & (midbrain) Tegmentum
Kigar, 1988). In adults, approximately 70 percent Cerebral aqueduct
of right-handed individuals show larger planum tem- Metencephalon Cerebellum
(across-brain) Pons
porale in the left hemisphere. The planum temporale
Fourth ventricle
has been related to phonological coding, a process
Myelencephalon Medulla oblongata Spinal cord
very important in reading (Semrud-Clikeman, (spinal brain) Fourth Ventricle
Hynd, Novey, & Eliopulos, 1991). The typical asym- Source: Adapted with permission from Fundamentals of
metry of the left hemisphere has not been observed Human Neuropsychology, 3rd edition, by B. Kolb and I. Q.
in those with developmental dyslexia and, thus, may Whishaw (1990). San Francisco: W. H. Freeman.
Cerebral Hemispheres 37

Zaidal, Clark, and Suyenobu (1990) suggest the price to be paid when one hemisphere assumes the
following: function of the other, usually involving the loss or
compromise of higher level functions. These more
(1) the two hemispheres can operate independently
complex functions also may be more dependent on
of one another, which reinforces the concept
the anatomical differences generally found between
of hemispheric specialization, in some domain-
the two hemispheres that exist early in the develop-
specific functions;
ing brain. This difference is most likely a result of the
(2) hemispheric specialization is hard-wired and
differential ratio of gray-to-white matter between the
is apparently innately directed;
two hemispheres described by Goldberg and Costa
(3) developmental patterns of the two hemispheres
(1981). Recovery and loss of functions will be cov-
may differ, and
ered in more detail in subsequent chapters.
(4) while the two hemispheres may share processing
resources, they can remain autonomous at any
stage of processing.
Functional neuroimaging techniques will help Interhemispheric Connections
answer these questions and will no doubt add to our
understanding of the relative contribution of the two Large bundles of myelinated fibers connect various
hemispheres, as well as specific structures, during intra- and interhemispheric regions. The two hemi-
certain activities. Some findings have implicated spheres are connected via several transverse com-
parts of the right hemisphere (particularly the poster- missures or pathways, including the corpus callo-
ior portion) to be important for visual-spatial and sum, the anterior commissure, and the posterior
mental rotation tasks (Perez-Fabello, Campos, & commissure. The corpus callosum, comprising the
Gomez-Juncal, 2007). Others have found more rostrum, the genu, the body, and the splenium,
activation in the left hemisphere for processing of contains approximately 300 million nerve fibers
language and verbal comprehension (Booth et al., for rapid interhemispheric communication
1999). In addition, studies are beginning to show a (Carlson, 2007). The genu connects rostral portions
right hemispheric preference for processing of emo- of the right and left frontal lobes, while the body has
tional information. Facial expression processing interconnections between the frontal and parietal
has been found to be bilateral and to involve the regions across the two hemispheres. The splenium
fusiform gyrus of the temporal lobe (Pierce, Muller, connects temporal and occipital regions and is
Ambrose, Allen, & Courchesne, 2001). While ana- reportedly larger in females (Semrud-Clikeman,
tomical differences appear early in development, Fine, & Bledsoe, 2008). The splenium has been
there is insufficient evidence to conclude that mor- implicated in various childhood disorders, includ-
phological variations between the two hemispheres ing ADHD (Hynd, Semrud-Clikeman, Lorys,
predict functional capabilities in any perfect sense Novey, & Eliopulos, 1991; Semrud-Clikeman et al.,
(Kinsbourne, 2003). 1994) and dyslexia (Fine, Semrud-Clikeman, Keith,
Damage to the left hemisphere can result in a Stapleton, & Hynd, 2006). The anterior commissure
shift of language functions to the right hemisphere, is smaller than the corpus callosum and connects the
particularly if both the posterior and anterior speech temporal lobes of the right and left hemispheres
zones are damaged (Kolb & Whishaw, 2003). While (Kolb & Whishaw, 2003).
language functions can be assumed by the right
hemisphere, complex visuospatial functions appear
to be in jeopardy (Kolb & Whishaw, 2003); further,
complex syntactic processing appears vulnerable. lntrahemispheric Connections
So, although the left hemisphere might be better
organized anatomically to deal with the language Association fibers connect cortical regions within
process, as suggested by the Goldberg and Costa each hemisphere (Kandel, Schwartz, & Jessell,
(1981) model, the right hemisphere is able to do 2000b). Association pathways allow for rapid
so under specific conditions. However, there is a communication within hemispheric regions for the
38 2 Functional Neuroanatomy

perception and integration of stimuli and to organize Structure and Function of the Cortex
complex output (e.g., emotional responses to stimuli).
Short association fibers connect one to another, and The forebrain (telencephalon) comprises the four
longer fibers connect one lobe to another. For exam- lobes, the lateral ventricles, the olfactory bulb, the
ple, the arcuate fasciculus connects the frontal and limbic system, the basal ganglia, and the neocor-
temporal lobes; the longitudinal fasciculus connects tex. Some textbooks also place the thalamus in the
the temporal and the occipital lobes with the frontal forebrain region, while others refer to this as a
lobe; the occipitofrontal fasciculus connects the diencephalic structure (Brodal, 2004). The cortex
frontal, temporal, and occipital lobes, and the angular comprises the right and left hemispheres, each
gyms connects the parietal and the occipital lobes with four major lobes: (1) frontal, motor cortex;
(Kandel, Schwartz, & Jessell, 2000a). Dysfunction of (2) parietal, somatosensory cortex; (3) occipital,
these pathways can result in a variety of behavioral, visual cortex, and (4) temporal, auditory cortex.
cognitive, and personality manifestations including (See Fig. 2.6 for a view of the cortical regions.)
reading, spelling, and computational disorders in Figure 2.7 illustrates the various functions of
children (Zaidel, Iacoboni, Zaidel, & Bogen, 2003). the lobes.

Movement fo muscles Sensory information

from body
Primary motor
cortex

Perceptions and memories Primary somatosensory

are translated into plans and cortex
actions by the frontal lobes
Sensory
association
cortex

Motor association
cortex Visual
information

Primary auditory cortex

Auditory information
Temporal lobe
pulled down Primary visual cortex
to show primary (mostly on inner surface)
auditory cortex

Sensory
association
cortex

Fig. 2.7 Major Structures and Functions of the Cortex

Source: From Neil R. Carlson, Physiology of Behavior, 5th edition, p. 91. Copyright # 1994 by Allyn and Bacon. Reprinted
with permission
Structure and Function of the Cortex 39

Frontal Lobes (midbrain structure), the pons, and the spinal cord
for the production of movement. The primary
The frontal lobes are the most anterior cortical cortex controls movements to the opposite side of
structures and comprise the primary motor cortex, the body and is arranged in a homuncular fashion.
the premotor cortex, an area of expressive language A homunculus is a schematic of brain function
(Brocas area), the medial cortex, and the prefrontal mapping onto specific body structures. Thus, there
cortex (Damasio & Anderson, 2003). Whereas the is a specific region that is responsible for movement
primary and premotor areas of the frontal lobes of the thumb, or the ankle, or the nose that maps
have major motor functions, the prefrontal cortex onto the primary motor cortex.
mediates reasoning and planning and monitors Specific muscle groups of the body are repre-
other cortical and subcortical functions. The pre- sented in an inverted pattern stretching across the
frontal cortex matures the most slowly of all of the primary motor area. Stimulation to specific areas of
areas of the lobes. the primary motor cortex produces contractions of
Lesions or damage to the primary motor cortex highly localized muscle areas. For example, Brocas
can result in paralysis to the contralateral side of area resides near the primary motor area in the
the body, whereas lesions to the premotor cortex left hemisphere, controls facial musculature, and
can produce more complex coordination problems mediates speech production (Kolb & Whishaw,
because this region directs the execution of the pri- 2003).
mary motor area (Kolb & Whishaw, 2003). Lesions
or damage to the prefrontal cortex, with its intricate
connections to other brain regions, including thala- Premotor Cortex
mic, hypothalamic, and limbic areas, often result in
affective dissociations, impaired executive functions The premotor cortex, anterior to the primary motor
and judgment, and intellectual deficits (Lezak, cortex, plays a role in controlling limb and body
Howieson, & Loring, 2004). movements. More complex, coordinated move-
ments appear to be regulated at this level, especially
fluid sequential movements. The premotor cortex
Primary Motor Cortex directs the primary cortex in the execution and main-
tenance of simple movements. The limbic system also
The motor system comprises the primary motor, the influences the motor cortex, directly and indirectly,
premotor, and to a lesser degree, the prefrontal, primarily in terms of attentional and motivational
with each region assuming differentiated motor aspects of motor functions (Damasio & Anderson,
functions. The primary motor cortex is involved 2003).
with the execution and maintenance of simple
motor functions; the premotor cortex directs the
primary motor cortex; and the prefrontal cortex Prefrontal Cortex
influences motor planning and adds flexibility to
motor behavior as a result of input from internal The prefrontal cortex, the most anterior region of
and external factors (Lezak et al., 2004). the frontal lobe, receives incoming signals from the
The primary motor cortex resides immediately thalamus, which then project to the hypothalamus.
anterior to the central sulcus and contains giant pyr- Further, connections to the limbic system allow the
amidal cells (Betz), which control fine motor and prefrontal cortex to mediate, regulate, and control
highly skilled voluntary movements (Damasio & affective, emotional behavior. Prefrontal connec-
Anderson, 2003). The primary motor cortex tions to the temporal, parietal, and occipital associa-
receives afferent (incoming sensory) signals from tion regions allow for a comparison of past and
the parietal lobe, the cerebellum, and the thalamus present sensory experiences (Gazzaniga et al.,
for the integration of sensory-motor signals, while 2002). These intricate connections of the prefrontal
efferent (outgoing motor) signals are transmitted to cortex with cortical and subcortical regions allow for
the reticular activating system, the red nucleus highly integrative, complex functions. Judgments
40 2 Functional Neuroanatomy

and insights arise out of prefrontal activity, whereas pressure and vibration; (3) recognition of fine touch
motor planning, consequential thinking, and ongoing (proprioception) and (4) awareness of the position
monitoring of behavior also appear to be regulated and movement of body parts (kinesthetic sense)
by prefrontal regions. The limbic system also seems (Lezak et al., 2004). Numerous fibers converge in
to play a role in complex, intentional, or volitional the primary sensory projection area, including
motor behaviors, though this is not considered part afferents coming from the thalamus, skin, muscles,
of the motor area. The development of executive joints, and tendons from the opposite side of the
control functions is discussed in more detail in later body. Lesions to the primary parietal regions can
sections of this chapter. Also see Chapter 6 for a produce sensory deficits to the contralateral (oppo-
discussion of neuropsychiatric disorders (e.g., site) side of the body, and other more complex
ADHD and Tourette syndrome) associated with deficits can occur when the temporoparietal and/
frontal lobe and executive control damage or or inferior parietal regions are involved (Tranel,
dysfunction. 1992).
Like the primary frontal cortex, the primary
sensory projection area is arranged in a homuncu-
Parietal Lobes lar fashion, with the proportion of cortical repre-
sentation related to the need for sensitivity in
a particular body region (Brodal, 2004). For
The parietal lobe is separated from the frontal
example, the region representing the face, lips,
regions by the central sulcus and from the temporal
and tongue is quite large because speech produc-
lobe by the lateral fissure. The parietal lobes play a
tion requires multiple sensory input from these
central role in the perception of tactile sensory
various muscles to provide sensory feedback to
information, including the recognition of pain,
orchestrate a complex series of movements needed
pressure, touch, proprioception, and kinesthetic
for speaking. The proximity of the primary parietal
sense. The parietal lobe is comprised of three
region to the primary motor regions allows for
areas: the primary sensory projection area, the sec-
the rapid cross-communication between sensory-
ondary somatosensory area, and the tertiary or
motor systems that is necessary for the execution
association area (Carlson, 2007).
of motor behavior.

Primary Sensory Cortex

Secondary and Association Cottices
The primary sensory projection area is immediately
posterior to the central sulcus, adjacent to the pri- Input from the primary sensory projection regions
mary motor cortex. Some have argued that there is a is synthesized into more complex sensory forms by
great deal of functional overlap between the sensory secondary parietal regions. The tertiary or associa-
and motor cortical areas with approximately one- tion region, the most posterior area of the parietal
quarter of the points in the motor area also showing lobe, receives input from the primary sensory
sensory capabilities and one-quarter of the points in projection area and sends efferents into the thala-
the sensory area also showing motor capabilities mus. The association region is involved with the
(Brodal, 2004). Thus, regions posterior to the sulcus integration and utilization of complex sensory
have been labeled as the sensory-motor area, while information. Gazzaniga et al. (2002) indicate that
regions anterior to the sulcus are labeled the motor- the association regions synthesize information,
sensory area. What seems most evident from this whereas the primary areas are involved with finer
research is that the sensory-motor regions are highly distinctions and analysis of information. The
interrelated, which probably results in increased association region overlaps with other cortical
functional efficiency. structures, including temporal and occipital areas
The primary sensory projection area has four for the integration of sensory information from
major functions: (1) recognition of the source, qual- different modalities. Although damage to the asso-
ity, and severity of pain; (2) discrimination of light ciation region does not produce visual, auditory,
Structure and Function of the Cortex 41

or sensory deficits, damage to the association area further divided into dorsal (superior) and ventral
can result in disorders of the integration of complex (inferior) areas. The inferior and superior regions
sensory information. Cross-modal matching of are divided by the lateral-occipital sulcus, while the
visual with auditory and sensory stimuli takes calcarine fissure extends from the occipital pole into
place in the association region, which is considered the splenium of the corpus callosum (see Fig. 2.7).
to be the highest level of sensory analysis. Some The visual cortex receives projections from the
argue that this region regulates much of what is retina in each eye via the lateral geniculate nucleus
measured by intelligence tests, including cognitive in the thalamus (see Fig. 2.8). The rods and cones in
and mental functions such as thinking, reasoning, the retina respond to photic stimulation, and photo-
and perception (Kolb & Whishaw, 2003). chemical processes result in nerve impulses in the
optic nerve (Carlson, 2007). Once inside the cere-
brum, the optic nerve forms the optic chiasm. The
optic chiasm is where nerve fibers partially cross,
Occipital Lobes project to the lateral geniculate in the thalamus, and
converge in the visual cortex. Damage anywhere
The most posterior region of the cortex comprises along this pathway can produce a variety of visual
the occipital lobe (primary visual cortex), which is defects.

Lateral geniculate Information from

nucleus right half of
visual field

Primary visual Optic nerve

cortex

Information from
left half of
visual field

Field of view
of right eye

Field of view
of left eye

Fig. 2.8 Visual Fields and Cortical Visual Pathways

Source: From Neil R. Carlson, Physiology of Behavior, 5th edition, p. 149. Copyright # 1994 by Allyn and Bacon. Reprinted
by permission
42 2 Functional Neuroanatomy

The occipital lobe comprises primary, secondary, (prosopagnosia) and object recognition (Bauer &
and tertiary or association regions (Kolb & Demery, 2003).
Whishaw, 2003). The primary occipital cortex The primary temporal cortex is involved with the
receives afferent input from the thalamus, which perception of speech sounds, particularly in the left
passes through the temporal cortex. Damage to hemisphere, and nonverbal tonal sequences, parti-
this tract, even if it occurs in the temporal lobe, cularly in the right hemisphere, while the secondary
can produce visual field defects. The association and association regions are more complex and var-
region is involved with complex visual perception, ied in function. The secondary and association
relating past visual stimuli to present stimuli for the regions add the affective quality to stimuli that is
recognition and appreciation of what is being seen. essential for learning to take place. When positive,
Damage to the association region, particularly in negative, or neutral affective qualities are attached
the right hemisphere, can produce a variety of visual to stimuli, information takes on motivational or
deficits, including recognition of objects, faces, and emotional importance to the learner. Without this
drawings. association, all stimuli would be judged as equal
and we would respond to all stimuli with the same
affect or emotion (Heilman, Blonder, Bowers, &
Valenstein, 2003).
Temporal Lobes The mesial temporal region, including the
adjoining hippocampus and amygdala, appears to
The temporal lobe has three major divisions: (1) the be linked to memory processes and plays a role in
posterior region of the superior temporal gyrus, learning or acquiring new information (Tranel,
which is referred to as Wernickes area in the left 1992). Lesions in this area result in impaired reten-
hemisphere; (2) the inferior temporal region, includ- tion of new memories, as this region appears to be
ing the occipitotemporal association region, and related to the process by which new memories are
(3) the mesial temporal aspect, including the hippo- stored or are retrieved from storage (Lezak et al.,
campal and amygdala regions (Tranel, 1992). The 2004). Asymmetry of functions is evident in the
temporal lobe has complex interconnections, with temporal lobes. Memory functions appear to be
afferent fibers coming from the parietal lobe, effer- lateralized. The recall of verbal information, includ-
ent fibers projecting into the parietal and frontal ing stories and word lists presented either orally or
lobes, and the corpus callosum and the anterior visually, is stored in the left hemisphere, whereas
commissure connecting the right and left temporal nonverbal recall for geometric drawings, faces,
lobes (Kolb & Whishaw, 2003). Three major path- and tunes is stored in the right hemisphere (Kolb &
ways connect the temporal lobe with other cortical Whishaw, 2003).
regions for complex integrated functions. The arcu-
ate fasciculus connects the frontal and temporal
lobes, the superior longitudinal fasciculus connects
the temporal to the occipital and frontal cortices Olfactory Bulb
(i.e., the sensory and motor regions of Wernickes
and Brocas areas), and the occipitofrontal fascicu- The olfactory system is the only sensory system that
lus connects the frontal-temporal-occipital regions. converges in the telencephalon. The olfactory bulb
The anatomical complexity, including large asso- receives sensory information concerning smell
ciation regions, suggests that the temporal lobes directly from the olfactory nerve and converges
have diverse functions, including the perception of with the olfactory tract; at this juncture, axons
auditory sensations, the analysis of affective tone in cross to the bulb in the opposite hemisphere via the
auditory stimuli, and long-term memory storage. anterior commissure. The olfactory tract projects to
Although the temporal lobe has primary auditory the primary olfactory cortex to a small region called
perception and association functions related to the uncus, close to the end of the temporal lobe
speech and language processing, it also plays a (Brodal, 2004). Although olfactory assessment is
significant role in memory functions and in facial often ignored, the sense of smell is frequently
Summary 43

associated with various neuropsychiatric distur- involving learning and memory. Seizure activity in
bances found in adults, particularly schizophrenia, limbic structures, particularly the hippocampus,
Parkinsons disease, multiple sclerosis, subfrontal sometimes includes temporal lobe structures as
tumors, and some brain injuries (Heilman & well (Lockman, 1994). Seizures at this site may
Valenstein, 2003). result in a temporary loss of consciousness and
loss of memory.

Limbic System Basal Ganglia

The limbic system is a complex deep collection of The term basal ganglia refers to all or some of
structures in the forebrain comprising the hippo- the masses of gray matter within the cerebral hemi-
campus, amygdale, septum, and cingulate gyms spheres, including the caudate nucleus, putamen,
(Kolb & Whishaw, 2003). The limbic system has and globus pallidus. The corpus striatum connects
widespread connections with the neocortex and to the neocortex and to the thalamus, and has
with the autonomic and endocrinological systems, ascending and descending pathways to midbrain
and is considered a primitive brain structure structures (red nucleus and substantia nigra), and
involved with the olfactory senses. It resides to the spinal cord (Brodal, 2004). Serotonin-rich
between two brain regions (diencephalon and connections from the raphe nuclei also reach the
telencephalon) and serves as an intermediary to striatum, the prefrontal regions, and the limbic
cognitive and emotional functions (Wilkinson, system. These serotonin pathways serve to inhibit
1986). In humans the limbic system has less to do motor actions and emotional responses. The basal
with the olfactory system than with emotional and ganglia are intimately involved with motor func-
memory functions that are essential for the survival tions and, when damaged, can produce postural
of the species. It also has preservation functions for changes, increases or decreases in muscle tone, and
the individual. movement changes (e.g., twitches, tremors or jerks).
Wilkinson (1986) describes a number of important Sydenhams chorea, a childhood disease resulting
functions of the limbic system, including: from poststreptoccal rheumatic fever involving
the corpus striatum, is characterized by irregular
(1) analyzing and responding to fearsome, threa-
and purposeless movements. This disease usually
tening situations;
appears insidiously, gradually worsening with
(2) monitoring sexual responses, including repro-
symptoms of hyperkinetic movement disorder,
ducing and nurturing offspring;
emotional lability, and hypotonia. Rheumatic
(3) remembering recent and past events, and
heart disease is often found in conjunction with
(4) sensing and responding to feeling states,
Sydenhams chorea and is the cause of mortality
including pleasure.
in this disorder. The chorea generally dissipates
Autonomic responses (e.g., heart rate, breathing, six months after onset, but the emotional lability
blood pressure, and digestive functions) can be remains (Cardona et al., in press).
influenced by limbic structures, especially the
cingulate gyrus. Aggressive reactions and social
indifference have been associated with the cingulate
gyrus. The cingulate gyrus has also been associated Summary
with error checking and self-monitoring of behavior
(Semrud-Clikeman, Pliszka, Lancaster, & Liotti, This chapters goal was to provide an introduction
2006). Feelings of anxiety, deja vu experiences, to the field of neuropsychology. Defining neurop-
rage, and fear have been associated with functions sychology includes the study of the brain and
of the amygdala (Gazzaniga et al., 2002). With the transactional effect of neurology and the
its connections with other limbic and cortical environment. Neuropsychologists are specialists
structures, the hippocampus has broad functions in assessment and intervention, and specific and
44 2 Functional Neuroanatomy

intensive training is necessary for practice in this Booth, J. R., MacWhinney, B., Thulborn, K. R., Sacco, K.,
field. Child (pediatric) neuropsychologists require Voyvodic, J. T., & Feldman, H. M. (1999). Functional
organization of activation patterns in children: Whole
additional training in child development as well as brain fMRI imaging during three different cognitive
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