CNS Gray Matter and White Matter; PNS Ganglia and NervesAreas of the

CNS made up mainly of myelinated axons are called white matter.Areas

made up mainly of cell bodies are called gray matter. Most of thelocal
synaptic communication between neurons in the CNS occurs in the
graymatter, while axons in the white matter transmit signals over greater
distances.The surface of the cerebral hemispheres is covered by a unique
mantle ofgray matter called the cerebral cortex, which is far more
developed in highermammals than in other species. Beneath this lies the
white matter, which conveyssignals to and from the cortex (Figure 2.7A).
Gray matter is also found inlarge clusters of cells called nuclei located deep
within the cerebral hemispheresand brainstem. Examples include the basal
ganglia, thalamus, and cranial nervenuclei (see Figure 2.7A,B).In the
cerebral hemispheres the gray matter cortex is outside, while thewhite
matter is inside. In the spinal cord the opposite is true: White
matterpathways lie on the outside, while the gray matter is in the center
(see Figure2.7C). In the brainstem, gray matter and white matter regions
are found bothon the inside and on the outside, although most of the
outside surface is whitematter.Several different names with similar
meaning are usedfor white matter pathways in the CNS, including tract
,fascicle , lemniscus , and bundle . A white matter pathwaythat connects
structures on the right and left sides of theCNS is called a commissure .
Axons in the PNS form bundlescalled peripheral nerves , or simply nerves.
Clustersof cell bodies in the PNS are referred to as ganglia .In general,
pathways carrying signals toward a structureare called afferent , while
those carrying signalsaway from a structure are called efferent (a fferents a
rrive,e fferents e xit). Thus, peripheral nerves convey afferentsensory
information about the environment tothe CNS and carry efferent signals for
motor activityfrom the CNS to the periphery.Spinal Cord and Peripheral
Nervous System The human nervous system develops in segments similarto
those of simpler animals, such as segmented worms.As already described,
the segments in the head expandand fuse together, forming the cerebral
hemispheres andbrainstem. Twelve pairs of cranial nerves (see Figure
2.1)exit these segments (these will be discussed further laterin the chapter).
The spinal nerves arise from the segmentsof the spinal cord (Figure 2.8A ).
Each segment gives rise toboth sensory and motor nerve roots on each side
of thebody (Figure 2.8B ).Throughout the nervous system, motor systems
tendto be more ventral, or anterior, and sensory systemsmore dorsal, or
posterior. The same holds true for thespinal cord. Thus, dorsal nerve roots
convey mainly afferentsensory information into the dorsal spinal cord,while
ventral nerve roots carry mainly efferent motorsignals from the ventral
spinal cord to the periphery.The segments and nerve roots of the spinal
cord arenamed according to the level at which they exit thebony vertebral
canal. Thus, there are cervical , thoracic ,lumbar , and sacral nerve roots
(see Figure 2.8A).During development, the bony vertebral canal increases
in length fasterthan the spinal cord. Therefore, the spinal cord ends at the
level of the first orsecond lumbar vertebral bones (L1 or L2). Below this the
spinal canal containsa collection of nerve roots known as the cauda equina
(Latin for “horse’stail”), which continue down to their exit points. The
sensory and motor nerveroots join together a short distance outside the
spinal cord and form a mixedsensory and motor spinal nerve (see Figure
2.8B). Control of the arms andlegs requires much more signal flow than
does control of the chest and abdomen.Thus, the nerves controlling the
extremities give rise to elaboratemeshworks referred to as the brachial
plexus for the arms and the lumbosacralplexus for the legs (see Figure
2.8A). In addition, the spinal cord contains arelatively increased amount of
gray matter in these segments, causing theoverall thickness of the cord to
be greater. These regions of the cord are calledthe cervical enlargement
and the lumbosacral enlargement , respectively.In addition to the sensory
and motor pathways already described, thePNS includes some specialized
neurons that are involved in controlling suchautomatic functions as heart
rate, peristalsis, sweating, and smooth musclecontraction in the walls of
blood vessels, bronchi, sex organs, the pupils, andso on. These neurons are
part of the autonomic nervous system . The autonomicnervous system has
two major divisions (Figure 2.9 ): The sympathetic division arises from
thoracic and lumbar spinal levels T1 to L3 (the thoracolumbar division). It
releases the neurotransmitter norepinephrine onto endorgans and is
involved in such “fight or flight” functions as increased heartrate and blood
pressure, bronchodilation, and increased pupil size. Theparasympathetic
division, in contrast, arises from the cranial nerves andfrom sacral spinal
levels S2 to S4 (the craniosacral division). It releases acetylcholine onto
end organs and is involved in more sedentary functions, suchas increasing
gastric secretions and peristalsis, slowing the heart rate, anddecreasing
pupil size. The sympathetic and parasympathetic pathways arecontrolled by
higher centers in the hypothalamus and limbic system as wellas by afferent
sensory information from the periphery.The enteric nervous system is
considered a third autonomic division andconsists of a neural plexus, lying
within the walls of the gut, that is involvedin controlling peristalsis and
gastrointestinal secretions.Cerebral Cortex: Basic Organization and
Primary Sensoryand Motor Areas The cerebral cortex is not a smooth sheet,
but rather has numerous infoldingsor crevices called sulci . The bumps or
ridges of cortex that rise up betweenthe sulci are called gyri . Some sulci
and gyri have particular names and functions,as we will learn shortly. The
cerebral hemispheres have four major lobes:the frontal, temporal, parietal,
and occipital (Figure 2.10 ).Lobes of the Cerebral Hemispheres The frontal
lobes are, appropriately, in the front of the brain and extend backto the
central sulcus of Rolando . The frontal lobes are separated inferiorly
andlaterally from the temporal lobes by an especially deep sulcus called the
Sylvianfissure , or lateral fissure. (The term fissure is sometimes used to
refer todeep sulci.) The parietal lobes are bounded anteriorly by the central
sulcus buthave no sharp demarcation from the temporal lobes or the
occipital lobes whenviewed from the lateral side of the brain (see Figure
2.10A). When viewed fromthe medial aspect, the parieto-occipital sulcus
can be seen more easily, separatingthe parietal from the occipital lobes (see
Figure 2.10B).In addition to these four major lobes, an additional region of
cerebral cortexcalled the insular cortex lies buried within the depths of the
Sylvian fissure.The insula is covered by a lip of frontal cortex anteriorly and
parietalcortex posteriorly, called the frontal operculum and parietal
operculum , respectively(operculum means “covering” or “lid” in Latin)
(see Figure 2.24B).The limbic cortex (see Figure 2.25) was formerly
referred to as the “limbiclobe,” but this terminology is no longer generally
used.The two cerebral hemispheres are separated in the midline by the
interhemisphericfissure , also known as the sagittal or longitudinal fissure
(Figure2.11D ). A large, C-shaped band of white matter called the corpus
callosum (meaning “hard body”) connects both homologous and
heterologous areasin the two hemispheres (see Figure 2.10B).Surface
Anatomy of the Cerebral Hemispheres in Detail Although there is some
variability, the sulci and gyri of the cerebral hemispheresform certain fairly
consistent patterns. We will now briefly reviewthe names of the major sulci,
gyri, and other structures of the cerebral hemispheres(see Figure 2.11).
Functions of these structures will be discussed in thenext section and
throughout the remainder of the book.On the lateral surface (see Figure
2.11A), the frontal lobe is bounded posteriorlyby the central sulcus, as
already noted. The gyrus running in front ofthe central sulcus is called the
precentral gyrus . The remainder of the lateralfrontal surface is divided into
the superior , middle , and inferior frontal gyri bythe superior and inferior
frontal sulci . Similarly, the lateral temporal lobe is dividedinto superior ,
middle , and inferior temporal gyri by the superior andmiddle temporal
sulci . The most anterior portion of the parietal lobe is thepostcentral gyrus
, lying just behind the central sulcus. The intraparietal sulcus divides the
superior parietal lobule from the inferior parietal lobule . The
inferiorparietal lobule consists of the supramarginal gyrus (surrounding
the endof the Sylvian fissure) and the angular gyrus (surrounding the end
of the superiortemporal sulcus).On the medial surface (see Figure 2.11B),
the corpus callosum is clearly visible,consisting of the rostrum , genu ,
body , and splenium . The cingulate gyrus (cingulum means “girdle” or
“belt”) surrounds the corpus callosum, runningfrom the paraterminal gyrus
anteriorly to the isthmus posteriorly. Thecingulate sulcus has a marginal
branch running up to the superior surfacethat forms an important
landmark, since the sulcus immediately in front ofit, on the superior
surface, is the central sulcus. The central sulcus does notusually extend
onto the medial surface, but the region surrounding it iscalled the
paracentral lobule . The portion of the medial occipital lobe belowthe
calcarine fissure is called the lingula (meaning “little tongue”), while
theportion above the calcarine fissure is called the cuneus (meaning
“wedge”).Just in front of the cuneus, the medial parietal lobe is called the
precuneus .On the inferior surface (see Figure 2.11C), the orbital frontal
gyri can beseen, which lay on top of the orbital ridges of the eye. More
medially, the olfactorysulcus (containing the olfactory bulb) separates the
orbital frontal gyrifrom the gyrus rectus (meaning “straight gyrus”). On the
inferior surface ofthe temporal lobe, the inferior temporal sulcus separates
the inferior temporalgyrus from the occipitotemporal , or fusiform , gyri .
More medially, the collateralsulcus , continuing anteriorly as the rhinal
sulcus , separates the fusiformgyri from the parahippocampal gyrus
.Finally, on the superior surface (see Figure 2.11D), many of the same
landmarksseen on the lateral surface are again visible.Primary Sensory and
Motor AreasThe primary sensory and motor areas of the cortex are shown
in Figure 2.12. Theprimary motor cortex lies in the precentral gyrus in the
frontal lobe (see Figure2.11A). This area controls movement of the opposite
side of the body. The primarysomatosensory cortex is in the postcentral
gyrus in the parietal lobe and isinvolved in sensation for the opposite side
of the body. Note that the precentraland postcentral gyri are separated by
the central sulcus and that (as in the spinalcord) motor areas lie anterior to
somatosensory areas. The primary visual cortexis in the occipital lobes
along the banks of a deep sulcus called the calcarinefissure (see Figures
2.11B and 2.12). The primary auditory cortex is composed ofthe transverse
gyri of Heschl, which are two fingerlike gyri that lie inside the
Sylvianfissure on the superior surface of each temporal lobe (see Figures
2.12and 2.24B). Higher-order sensory and motor information processing
takes placein the association cortex, as will be discussed later in this
chapter.Sensory and motor pathways are usually topographically organized.
Thismeans that adjacent areas on the receptive (or motor) surface are
mapped toadjacent fibers in white matter pathways and to adjacent regions
of cortex.For example, in primary motor and primary somatosensory
cortex, regionsrepresenting the hand are adjacent to regions representing
the arm, and soon (Figure 2.13). These somatotopic maps on the cortex are
sometimes calledthe motor or sensory homunculus (“little man”). Similarly,
adjacent retinalareas are mapped in a retinotopic fashion onto the primary
visual cortex,and adjacent regions of the cochlea, sensing different
frequencies, have atonotopic representation on the primary auditory
cortex.Interestingly, the primary somatosensory cortex and primary motor
cortexrepresent sensation and movement, respectively, for the opposite
side ofthe body. This relationship was first noted by physicians in ancient
Greece,including Hippocrates, who observed that patients with head
injuries haddeficits affecting the side of the body opposite to the side of the
injury.Knowledge of the levels at which the somatosensory and motor
pathwayscross over in the nervous system can be helpful for clinical
neuroanatomicallocalization and will be discussed later in this chapter. The
primary visualcortex represents visual inputs from the opposite visual field.
Thus, the lefthalf of the visual field for each eye is mapped to the right
primary visual cortex(see Figure 11.15). Information reaching the primary
auditory cortex isless lateralized and represents more of a mixture of inputs
from both ears(the input from the opposite ear is slightly stronger, but this
is usually notclinically detectable).Cell Layers and Regional Classification of
the Cerebral Cortex The majority of the cerebral cortex is composed of
neocortex , which has six celllayers , labeled I through VI, counting from
the surface inward (Figure 2.14 ; Table2.3 ). In a few regions associated
with the limbic system, less than six layers arepresent. Neocortical circuitry
is quite complex, and we will describe only a fewof the major connections of
each layer here. Layer I contains mainly dendritesof neurons from deeper
layers as well as axons. Layers II and III contain neuronsthat project
mainly to other areas of cortex. Layer IV receives the majority of
inputsfrom the thalamus. Layer V projects mostly to subcortical structures
otherthan the thalamus, such as the brainstem, spinal cord, and basal
ganglia. LayerVI projects primarily to the thalamus. In addition to these
connections, numerousother circuits exist between and within the cortical
layers that are beyondthe scope of this discussion. The cortical layers I
through VI have alternativenames that will not be used here but are
included in Table 2.3 for reference.The relative thickness of the cell layers
varies according to the main functionof that area of cortex. For example,
the primary motor cortex has large efferentprojections to the brainstem and
spinal cord, which control movement.It receives relatively little direct
sensory information from thalamic relay centers.Therefore, in the primary
motor cortex, layer V is thicker and has manymore cell bodies than layer IV
(see Figure 2.14B). The opposite holds for primaryvisual cortex, where layer
IV contains many cell bodies and layer V isrelatively cell poor (see Figure
2.14C). Association cortex has a cellular structurethat is intermediate
between these types (see Figure 2.14A).Avariety of classification schemes
exist for different regions of the cerebralcortex based on microscopic
appearance and function. The most widelyknown of these was published by
Korbinian Brodmann in 1909. Based on hisstudies conducted with the
microscope, Brodmann parceled the cortex into 52cytoarchitectonic areas ,
each assigned a number corresponding to the order inwhich he prepared
the slides (Figure 2.15 ; Table 2.4 ). It turns out that many ofthe areas
identified by Brodmann correlate fairly well with various functionalareas of
the cortex, and therefore his nomenclature is still often used today.Motor
SystemsMotor control involves a delicate balance between multiple parallel
pathwaysand recurrent feedback loops. We will now provide an overview of
the mostimportant motor pathways and of the cerebellum and basal
ganglia, which aremajor feedback systems.Main Motor PathwaysThe most
important motor pathway in humans is the corticospinal tract.
Thecorticospinal tract begins mainly in the primary motor cortex, where
neuroncell bodies project via axons down through the cerebral white matter
and brainstemto reach the spinal cord (Figure 2.16). The corticospinal tract
is sometimescalled the pyramidal tract because of its triangular shape in
the medulla (seeFigures 6.11Aand 14.5B). The majority of fibers in the
corticospinal tract (about85%) cross over to control movement of the
opposite side of the body. Thiscrossing over, known as the pyramidal
decussation, occurs at the junction betweenthe medulla and the spinal cord.
Thus, lesions occurring above the pyramidaldecussation produce
contralateral (opposite-sided) weakness with respectto the lesion, while
lesions below the pyramidal decussation will produceipsilateral (same-
sided) weakness. Several other descending motor pathwaysthat exist in
addition to the corticospinal tract will be discussed in Chapter 6.Motor
neurons that project from the cortex down to the spinal cord orbrainstem
are referred to as upper motor neurons (UMNs ). UMNs formsynapses
onto the lower motor neurons (LMNs ), which are located in the
anteriorhorns of the central gray matter of the spinal cord (see Figure
2.16B) or inbrainstem motor nuclei . The axons of LMNs project out of the
CNS via theanterior spinal roots or via the cranial nerves to finally reach
muscle cells inthe periphery. Lesions affecting UMNs and LMNs have
certain distinct clinicalfeatures, which we will learn about in Chapter
3.Cerebellum and Basal GangliaThe motor system is called upon to perform
many delicate and complicatedtasks. Therefore, in order to refine the
output of the motorsystem, multiple feedback systems are employed,
including the cerebellumand basal ganglia (Figure 2.17). These two systems
do notthemselves project directly to the LMNs. Instead, the cerebellumand
basal ganglia act by modulating the output of the corticospinaland other
descending motor systems. The cerebellum and basal gangliaboth receive
major inputs from the motor cortex. The cerebellumalso receives significant
inputs from the brainstem and spinalcord, as discussed in Chapter 15. The
cerebellum and basal ganglia,in turn, project back to the motor cortex via
the thalamus.Lesions in the cerebellum lead to disorders in
coordinationand balance, often referred to as ataxia. Lesions in the basal
gangliacause hypokinetic movement disorders, such as Parkinsonism,in
which movements are infrequent, slow, and rigid, and
hyperkineticmovement disorders, such as Huntington’s disease, which
ischaracterized by dancelike, involuntary movements.Somatosensory
SystemsSensation from the body is conveyed by parallel pathways
mediating differentsensory modalities that travel to the central nervous
system. We will nowreview the most important sensory pathways and
introduce the thalamus, amajor relay center for signals of all kinds (sensory
and other) that travel tothe cerebral cortex.Main Somatosensory
PathwaysSomatic sensation refers to the conscious perceptions of touch,
pain, temperature,vibration, and proprioception (limb or joint position
sense). There are twomain pathways in the spinal cord for somatic
sensation:1. The posterior column pathways (Figure 2.18) convey
proprioception,vibration sense, and fine, discriminative touch.2. The
anterolateral pathways (Figure 2.19) convey pain, temperature sense,and
crude touch.Some aspects of touch sensation are carried by both pathways,
so touch sensationis not eliminated in isolated lesions of either pathway.
Note that the primarysensory neuron cell bodies are located outside the
CNS in the dorsal rootganglia and that they have bifurcating axons, with
one long process extendingto the periphery and one into the spinal cord
(see Figures 2.18 and 2.19). Forlocalizing clinical lesions, the knowledge of
where in the CNS the two mainsensory pathways cross over is equally as
important as the knowledge that thecorticospinal tract crosses over at the
pyramidal decussation. Therefore, wewill briefly outline the two main
sensory pathways here.POSTERIOR COLUMN PATHWAY Primary sensory
neuron axons carrying informationabout proprioception, vibration sense,
and fine touch enter first thespinal cord via the dorsal roots and then the
ipsilateral white matter dorsal(posterior) columns to ascend all the way to
the dorsal column nuclei in themedulla (see Figure 2.18). Here they make
synapses onto the secondary sensoryneurons, which send out axons that
cross over to the other side of themedulla. These axons continue to ascend,
now on the contralateral side, andsynapse in the thalamus, and from there
neurons project to the primary somatosensorycortex in the postcentral
gyrus.ANTEROLATERAL PATHWAY Primary sensory neurons carrying
informationabout pain, temperature sense, and crude touch also enter the
spinal cordvia the dorsal roots (see Figure 2.19). However, these axons
make their firstsynapses immediately in the gray matter of the spinal cord.
Axons from thesecondary sensory neurons cross over to the other side of
the spinal cordand ascend in the anterolateral white matter, forming the
spinothalamictract. After synapsing in the thalamus, the pathway again
continues to theprimary somatosensory cortex.Thalamus The thalamus is
an important relay center. Nearly all pathways that project tothe cerebral
cortex do so after synapsing in the thalamus. The thalami are graymatter
structures located deep within the cerebral white matter just abovethe
brainstem and behind the basal ganglia (see Figure 2.17 and Figure 2.20
).They are shaped somewhat like eggs, with their posterior ends angled
outward,together forming an inverted V in horizontal sections (see Figures
2.20Band 16.2). The thalamus consists of multiple nuclei. Each sensory
modality, includingvision, hearing, taste, and somatic sensation, has a
different nucleararea where synapses occur before the information is
relayed to the cortex (olfactoryinputs are an exception and do not pass
directly through the thalamus).Nonsensory pathways also relay in the
thalamus. For example, there are thalamicnuclei that process information
coming from the basal ganglia, cerebellum,limbic pathways, and brainstem
reticular formation on the way to the cortex(see Figure 2.20A). An
important feature of thalamic circuits is the reciprocalnature of cortical–
thalamic connections. Thus, virtually all cortical regions projectstrongly via
layer VI (see Table 2.3) back to the thalamic areas from whichtheir major
inputs arise.As we discussed earlier in this chapter, the thalamus, together
with thehypothalamus and epithalamus, form the diencephalon (see Figure
2.2). Thehypothalamus is an important region for control of autonomic,
neuroendocrine,limbic, and other circuits. The epithalamus encompasses
severalsmall nuclei, including the pineal body (see Figure 2.11B), habenula,
andparts of the pretectum.Stretch Reflex The monosynaptic stretch reflex
is a well-studied reflex arc that provides rapidlocal feedback for motor
control. The reflex arc begins with specialized receptorscalled muscle
spindles , which detect the amount and rate of stretch in muscles(Figure
2.21 ). This information is transmitted to the distal processes of
sensoryneurons and is then conveyed via the dorsal roots into the spinal
graymatter. In the spinal gray matter the sensory neurons form multiple
synapses,including some direct synapses onto LMNs in the anterior horn.
The LMNsproject via the ventral roots back out to the muscle, causing it to
contract. Damageanywhere in this pathway (see Figure 2.21) can cause the
reflex to be diminishedor absent.In addition to the monosynaptic pathway,
the afferent sensory neuronforms synapses onto excitatory and inhibitory
interneurons in the spinalgray matter, which in turn make synapses onto
LMNs. Thus, local circuits inthe spinal cord can use sensory information to
regulate the activity of LMNswithout conscious input from higher centers.
There are, nevertheless, descendingpathways that modulate the activity of
the stretch reflex. As wewill learn, if these higher centers or their
descending pathways are damaged,the stretch reflex may become
hyperactive or hypoactive . Thus, by testingthe stretch reflex on neurologic
exam, one obtains information aboutmultiple pathways, including sensory
neurons and motor neurons in thePNS and descending modulatory
pathways in the CNS. Since the stretch reflexis often tested on neurologic
exam by tapping over a tendon with the reflexhammer (see Chapter 3 or
www.neuroexam.com ), this is also commonlyknown as a deep tendon
reflex .Brainstem and Cranial Nerves The overall structure of the brainstem
can be seen in Figure 2.22 . As discussedearlier, the brainstem is composed
of the midbrain, pons, and medulla. It isconnected to the diencephalon
rostrally, the cerebellum dorsally, and the spinalcord caudally (see Figure
2.22C). Most of the cranial nerves arise from the brainstem.The cranial
nerves are analogous in some ways to the spinal nerves, hav- ing both
sensory and motor functions. However, they also carry out more
specializedfunctions relating to the organs of the head (Table 2.5 ).
Examination ofthe cranial nerves provides crucial information about the
functioning of thenervous system, as we will discuss in the next chapter.In
addition to the cranial nerve nuclei and pathways, the brainstem istightly
packed with numerous other important nuclei and white matter tracts.All
information passing between the cerebral hemispheres and the spinal
cordmust pass through the brainstem. Therefore, lesions in the brainstem
can havea devastating effect on sensory and motor function. In addition,
the brainstemcontains nuclei that play important roles in the motor system;
nuclei that producenausea and vomiting in response to certain chemicals;
modulatory nucleicontaining the neurotransmitters norepinephrine,
serotonin, dopamine, andacetylcholine (see Table 2.2), which project
widely throughout the CNS; nuclearareas involved in pain modulation; and
nuclei controlling heart rate,blood pressure, and respiration, among other
functions.An important region of the brainstem that contains many of these
nuclei isthe reticular formation . Named for the network-like appearance of
its fibers inhistological sections, the reticular formation extends throughout
the centralportions of the brainstem from the medulla to the midbrain. The
more caudalportions of the reticular formation in the medulla and lower
pons tend to beinvolved mainly in motor and autonomic functions. The
rostral reticular formationin the upper pons and midbrain plays an
important role in regulatingthe level of consciousness , influencing higher
areas through modulation ofthalamic and cortical activity (Figure 2.23 ).
Thus, lesions that affect the pontomesencephalicreticular formation can
cause lethargy and coma.Cortical, thalamic, and other forebrain networks
are also important inmaintaining consciousness. Therefore, the level of
consciousness can also beimpaired in bilateral lesions of the thalami or in
bilateral (or large unilateral)lesions of the cerebral hemispheres. Note that
mass lesions above the brainstemoften cause impaired consciousness
indirectly when they exert pressureon the brainstem through mass effect,
thus distorting or compressingthe reticular formation and thalamus.Limbic
System Several structures in the brain are referred to collectively as the
limbic system because they are located near the medial edge or fringe
(limbus in Latin) ofthe cerebral cortex (Figure 2.24 ). These structures
have evolved, from a systemdevoted mainly to olfaction in simpler animals,
to perform diverse functions,including the regulation of emotions, memory,
appetitive drives, and autonomicand neuroendocrine control. The limbic
system includes certaincortical areas located in the medial and anterior
temporal lobes (see Figure2.24A), anterior insula (see Figure 2.24B),
inferior medial frontal lobes, and cingulategyri. It also includes deeper
structures, such as the hippocampal formation and the amygdala , located
within the medial temporal lobes (see Figure2.24A), several nuclei in the
medial thalamus, hypothalamus, basal ganglia,septal area, and brainstem.
These areas are interconnected by a variety of pathways,including the
fornix —a paired, arch-shaped white matter structurethat connects the
hippocampal formation to the hypothalamus and septalnuclei (see Figure
2.24A).Lesions in the limbic system can cause deficits in the consolidation
of immediaterecall into longer-term memories. Thus, patients with lesions
inthese areas may have no trouble recalling remote events but have
difficultyforming new memories. In addition, limbic dysfunction can cause
behavioralchanges and may underlie a number of psychiatric disorders.
Finally,epileptic seizures most commonly arise from the limbic structures
of the medialtemporal lobe, resulting in seizures that may begin with
emotions suchas fear, memory distortions such as déjà vu, or olfactory
hallucinations.Association Cortex In addition to the primary motor and
sensory areas described earlier, the cerebralcortex contains a large amount
of association cortex (Figure 2.25 ), whichcarries out higher-order
information processing. In unimodal association cortex,higher-order
processing takes place mostly for a single sensory or motormodality.
Unimodal association cortex is usually located adjacent to a primarymotor
or sensory area; for example, unimodal visual association cortex is
locatedadjacent to the primary visual cortex, and unimodal motor
associationcortex (premotor cortex and supplementary motor area) is
located adjacent tothe primary motor cortex (see Figure 2.25).
Heteromodal association cortex isinvolved in integrating functions from
multiple sensory and/or motor modalities.We will now briefly review the
functions of several clinically importantareas of the association cortex,
particularly since we will describe how to examinethese structures with the
Mental Status exam in the next chapter.Language is usually perceived first
by the primary auditory cortex in thesuperior temporal lobe when we are
listening to speech or by the primaryvisual cortex in the occipital lobes
when we are reading. From here, cortical–cortical association fibers convey
information to Wernicke’s area in thedominant (usually left) hemisphere
(see Figure 2.25). Lesions in Wernicke’sarea cause deficits in language
comprehension, also sometimes called receptive or sensory aphasia , or
Wernicke’s aphasia . Broca’s area is located in thefrontal lobe, also in the
left hemisphere, adjacent to the areas of primarymotor cortex involved in
moving the lips, tongue, face, and larynx. Lesionsin Broca’s area cause
deficits in the production of language, with relativesparing of language
comprehension. This is called expressive or motor aphasia ,or Broca’s
aphasia , which can be remembered by the mnemonic “Broca’sbroken boca
” (boca means “mouth” in Spanish).The parietal lobe is divided by the
intraparietal sulcus into a superior andinferior parietal lobule (see Figures
2.11A,D). Lesions in the inferior parietallobule in the left hemisphere can
produce an interesting constellation of abnormalities,including difficulty
with calculations, right–left confusion, inabilityto identify fingers by name
(finger agnosia ), and difficulties with writtenlanguage. This group of
abnormalities is called Gerstmann’s syndrome .Performance of complex
motor tasks, such as brushing one’s teeth orthrowing a baseball, requires
higher-order planning before the primarymotor cortex can be activated.
Motor planning appears to be distributed inmany different areas of cortex.
Thus, diffuse lesions of the cortex, or sometimesmore focal lesions affecting
the frontal or left parietal lobe, can produceabnormalities in motor
conceptualization, planning, and executioncalled apraxia .The parietal
lobes also play an important role in spatial awareness. Thus,lesions in the
parietal lobe, especially in the nondominant (usually right)hemisphere ,
often cause a distortion of perceived space and neglect of thecontralateral
side. For example, right parietal lesions can cause left hemineglect .With
this syndrome, patients will often ignore objects in their left visualfield, but
they may see them if their attention is strongly drawn to thatside. They may
draw a clock face without filling in any numbers on the leftside of the clock.
They may also be completely unaware of the left side oftheir body, thinking,
for example, that their left arm belongs to someoneelse, and they may be
unaware of any weakness or other deficits on thatside. Unawareness of a
deficit is called anosognosia (from the Greek a ,“lack”; nosos , “disease”;
gnosis , “knowledge”).Patients may also display a phenomenon called
extinction , in which a tactileor visual stimulus is perceived normally when
it is presented to one sideonly, but when it is presented on the side opposite
the lesion simultaneously with an identical stimulus on the normal side, the
patient neglects the stimuluson the side opposite the lesion. These severe
abnormalities in spatialorientation and awareness are less common in
lesions of the dominant (usuallyleft) parietal lobe, possibly because the
dominant hemisphere is morespecialized for language than it is for
visuospatial functions.The frontal lobes are the largest hemispheres and
contain vast areas of associationcortex (see Figure 2.25). Lesions in the
frontal lobes cause a varietyof disorders in personality and cognitive
functioning. Frontal release signs are “primitive” reflexes that are normal in
infants, such as grasp , root , suck ,and snout reflexes , but that can also be
seen in adults with frontal lobe lesions.In addition, patients with frontal
lobe lesions may have particular difficultywhen asked to perform a
sequence of actions repeatedly or to changefrom one activity to another. In
doing these tasks they tend to perseverate ,meaning that they repeat a
single action over and over without moving onto the next one. Personality
changes with frontal lobe lesions may includeimpaired judgment, a cheerful
lack of concern about one’s illness, inappropriatejoking, and other
disinhibited behaviors. Other patients with frontallobe lesions may be
abulic (opposite of ebullient), with a tendency to starepassively and to
respond to commands only after a long delay. Frontal lesionscan also cause
a characteristic unsteady magnetic gait , in which the feetshuffle close to the
floor, and urinary incontinence .Lesions in the visual association cortex in
the parieto-occipital and inferiortemporal lobes can produce a variety of
interesting phenomena, includingprosopagnosia (inability to recognize
faces), achromatopsia (inability torecognize colors), palinopsia
(persistence or reappearance of an objectviewed earlier), and other
phenomena. Seizures in the visual association cortexcan cause elaborate
visual hallucinations.Blood Supply to the Brain and Spinal Cord There are
two pairs of arteries that carry all the blood supply to the brain andone pair
of draining veins (Figure 2.26A,B ). The internal carotid arteries form
theanterior blood supply, and the vertebral arteries , which join together in
a singlebasilar artery , form the posterior blood supply (see Figure 2.26A).
The anteriorand posterior blood supplies from the carotid and
vertebrobasilar systems,respectively, join together in an anastomotic ring at
the base of the braincalled the circle of Willis (see Figure 2.26C). The main
arteries supplying thecerebral hemispheres arise from the circle of Willis.
Ordinarily, however, theanterior and middle cerebral arteries derive their
main blood supply from theinternal carotid arteries (anterior circulation),
while the posterior cerebral arteries derive their main supply from the
vertebrobasilar system (posteriorcirculation). The main arteries supplying
the brainstem and cerebellum alsoarise from the vertebral and basilar
arteries. These include the superior , anteriorinferior , and posterior
inferior cerebellar arteries . Venous drainage for thebrain is provided
almost entirely by the internal jugular veins (see Figure 2.26B).The spinal
cord receives its blood supply from the anterior spinal artery ,which runs
along the ventral surface of the cord in the midline and from thepaired
posterior spinal arteries , which run along the right and left dorsal
surfacesof the cord (see Figure 6.5). The anterior and posterior spinal
arteriesare supplied in the cervical region mainly by branches arising from
the vertebralarteries (see Figure 2.26C). In the thoracic and lumbar regions,
thespinal arteries are supplied by radicular arteries arising from the
aorta.Conclusions In this chapter we have reviewed the overall structure
and organization of thenervous system. In addition, we have discussed in
general terms the functionsof each of the main areas in the brain, spinal
cord, and peripheral nervous system,and we have briefly reviewed the
blood supply to the brain. This overviewshould provide a framework upon
which important details will be filled inand reinforced through clinical
cases in the chapters that follow. It also providesthe necessary background
for the next chapter, in which we will introducethe neurologic exam and
explore what each portion of the exam can teachus about neuroanatomy.
Thus, you, the reader, have completed the roughsketch and now have
before you an undertaking that will require innumerablefiner and finer
brush strokes, each subtly enhancing the final composition.