The Central Nervous System

LEARNING OBJECTIVES

 Name the major regions of the adult brain

 Describe the connections between the cerebrum and brain stem through the diencephalon, and
from those regions into the spinal cord

 Recognize the complex connections within the subcortical structures of the basal nuclei

 Explain the arrangement of gray and white matter in the spinal cord

The brain and the spinal cord are the central nervous system, and they represent the main organs of the
nervous system. The spinal cord is a single structure, whereas the adult brain is described in terms of
four major regions: the cerebrum, the diencephalon, the brain stem, and the cerebellum. A person’s
conscious experiences are based on neural activity in the brain. The regulation of homeostasis is
governed by a specialized region in the brain. The coordination of reflexes depends on the integration of
sensory and motor pathways in the spinal cord.

The Cerebrum

The iconic gray mantle of the human brain, which appears to make up most of the mass of the brain, is
the cerebrum (Figure 1). The wrinkled portion is the cerebral cortex, and the rest of the structure is
beneath that outer covering. There is a large separation between the two sides of the cerebrum called
the longitudinal fissure. It separates the cerebrum into two distinct halves, a right and left cerebral
hemisphere. Deep within the cerebrum, the white matter of the corpus callosum provides the major
pathway for communication between the two hemispheres of the cerebral cortex.
Figure 1. This figure shows the lateral view on the left panel and anterior view on the right panel of
the brain. The major parts including the cerebrum are labeled.

Many of the higher neurological functions, such as memory, emotion, and consciousness, are the result
of cerebral function. The complexity of the cerebrum is different across vertebrate species. The
cerebrum of the most primitive vertebrates is not much more than the connection for the sense of
smell. In mammals, the cerebrum comprises the outer gray matter that is the cortex (from the Latin
word meaning “bark of a tree”) and several deep nuclei that belong to three important functional
groups. The basal nuclei are responsible for cognitive processing, the most important function being
that associated with planning movements. The basal forebrain contains nuclei that are important in
learning and memory. The limbic cortex is the region of the cerebral cortex that is part of the limbic
system, a collection of structures involved in emotion, memory, and behavior.

Cerebral Cortex

The cerebrum is covered by a continuous layer of gray matter that wraps around either side of the
forebrain—the cerebral cortex. This thin, extensive region of wrinkled gray matter is responsible for the
higher functions of the nervous system. A gyrus (plural = gyri) is the ridge of one of those wrinkles, and
a sulcus (plural = sulci) is the groove between two gyri. The pattern of these folds of tissue indicates
specific regions of the cerebral cortex. The head is limited by the size of the birth canal, and the brain
must fit inside the cranial cavity of the skull. Extensive folding in the cerebral cortex enables more gray
matter to fit into this limited space. If the gray matter of the cortex were peeled off of the cerebrum and
laid out flat, its surface area would be roughly equal to one square meter. The folding of the cortex
maximizes the amount of gray matter in the cranial cavity. During embryonic development, as the
telencephalon expands within the skull, the brain goes through a regular course of growth that results in
everyone’s brain having a similar pattern of folds. The surface of the brain can be mapped on the basis
of the locations of large gyri and sulci. Using these landmarks, the cortex can be separated into four
major regions, or lobes (Figure 2).

Figure 2. Lobes of the
Cerebral Cortex. The cerebral cortex is divided into four lobes. Extensive folding increases the surface
area available for cerebral functions.

The lateral sulcus that separates the temporal lobe from the other regions is one such landmark.
Superior to the lateral sulcus are the parietal lobe and frontal lobe, which are separated from each
other by the central sulcus. The posterior region of the cortex is the occipital lobe, which has no obvious
anatomical border between it and the parietal or temporal lobes on the lateral surface of the brain.
From the medial surface, an obvious landmark separating the parietal and occipital lobes is called
the parieto-occipital sulcus. The fact that there is no obvious anatomical border between these lobes is
consistent with the functions of these regions being interrelated.

Different regions of the cerebral cortex can be associated with particular functions, a concept known as
localization of function. In the early 1900s, a German neuroscientist named Korbinian Brodmann
performed an extensive study of the microscopic anatomy—the cytoarchitecture—of the cerebral
cortex and divided the cortex into 52 separate regions on the basis of the histology of the cortex. His
work resulted in a system of classification known as Brodmann’s areas, which is still used today to
describe the anatomical distinctions within the cortex (Figure 3). The results from Brodmann’s work on
the anatomy align very well with the functional differences within the cortex. Areas 17 and 18 in the
occipital lobe are responsible for primary visual perception. That visual information is complex, so it is
processed in the temporal and parietal lobes as well. The temporal lobe is associated with primary
auditory sensation, known as Brodmann’s areas 41 and 42 in the superior temporal lobe.

Figure 3. Brodmann's Areas of the Cerebral Cortex. Brodmann mapping of functionally distinct regions

of the cortex was based on its cytoarchitecture at a microscopic level.
Because regions of the temporal lobe are part of the limbic system, memory is an important function
associated with that lobe. Memory is essentially a sensory function; memories are recalled sensations
such as the smell of Mom’s baking or the sound of a barking dog. Even memories of movement are
really the memory of sensory feedback from those movements, such as stretching muscles or the
movement of the skin around a joint. Structures in the temporal lobe are responsible for establishing
long-term memory, but the ultimate location of those memories is usually in the region in which the
sensory perception was processed. The main sensation associated with the parietal lobe
is somatosensation, meaning the general sensations associated with the body.

Posterior to the central sulcus is the postcentral gyrus, the primary somatosensory cortex, which is
identified as Brodmann’s areas 1, 2, and 3. All of the tactile senses are processed in this area, including
touch, pressure, tickle, pain, itch, and vibration, as well as more general senses of the body such
as proprioception and kinesthesia, which are the senses of body position and movement, respectively.

Anterior to the central sulcus is the frontal lobe, which is primarily associated with motor functions.
The precentral gyrus is the primary motor cortex. Cells from this region of the cerebral cortex are the
upper motor neurons that instruct cells in the spinal cord to move skeletal muscles.

Anterior to this region are a few areas that are associated with planned movements. The premotor
area is responsible for thinking of a movement to be made. The frontal eye fields are important in
eliciting eye movements and in attending to visual stimuli. Broca’s area is responsible for the production
of language, or controlling movements responsible for speech; in the vast majority of people, it is
located only on the left side. Anterior to these regions is the prefrontal lobe, which serves cognitive
functions that can be the basis of personality, short-term memory, and consciousness. The prefrontal
lobotomy is an outdated mode of treatment for personality disorders (psychiatric conditions) that
profoundly affected the personality of the patient.

Subcortical Structures
 Figure 4. Frontal Section of
Cerebral Cortex and Basal Nuclei. The major components of the basal nuclei, shown in a frontal
section of the brain, are the caudate (just lateral to the lateral ventricle), the putamen (inferior to the
caudate and separated by the large white-matter structure called the internal capsule), and the globus
pallidus (medial to the putamen).

Beneath the cerebral cortex are sets of nuclei known as subcortical nuclei that augment cortical
processes. The nuclei of the basal forebrain serve as the primary location for acetylcholine production,
which modulates the overall activity of the cortex, possibly leading to greater attention to sensory
stimuli. Alzheimer’s disease is associated with a loss of neurons in the basal forebrain.

The hippocampus and amygdala are medial-lobe structures that, along with the adjacent cortex, are
involved in long-term memory formation and emotional responses. The basal nuclei are a set of nuclei in
the cerebrum responsible for comparing cortical processing with the general state of activity in the
nervous system to influence the likelihood of movement taking place. For example, while a student is
sitting in a classroom listening to a lecture, the basal nuclei will keep the urge to jump up and scream
from actually happening. (The basal nuclei are also referred to as the basal ganglia, although that is
potentially confusing because the term ganglia is typically used for peripheral structures.)

The major structures of the basal nuclei that control movement are the caudate, putamen, and globus
pallidus, which are located deep in the cerebrum. The caudate is a long nucleus that follows the basic C-
shape of the cerebrum from the frontal lobe, through the parietal and occipital lobes, into the temporal
lobe. The putamen is mostly deep in the anterior regions of the frontal and parietal lobes. Together, the
caudate and putamen are called the striatum. The globus pallidus is a layered nucleus that lies just
medial to the putamen; they are called the lenticular nuclei because they look like curved pieces fitting
together like lenses. The globus pallidus has two subdivisions, the external and internal segments, which
are lateral and medial, respectively. These nuclei are depicted in a frontal section of the brain
in Figure 4.

Figure 5. Connections of
Basal Nuclei. Input to the basal nuclei is from the cerebral cortex, which is an excitatory connection
releasing glutamate as a neurotransmitter. This input is to the striatum, or the caudate and putamen.
In the direct pathway, the striatum projects to the internal segment of the globus pallidus and the
substantia nigra pars reticulata (GPi/SNr). This is an inhibitory pathway, in which GABA is released at
the synapse, and the target cells are hyperpolarized and less likely to fire. The output from the basal
nuclei is to the thalamus, which is an inhibitory projection using GABA.

The basal nuclei in the cerebrum are connected with a few more nuclei in the brain stem that together
act as a functional group that forms a motor pathway. Two streams of information processing take place
in the basal nuclei. All input to the basal nuclei is from the cortex into the striatum (Figure 5).

The direct pathway is the projection of axons from the striatum to the globus pallidus internal segment
(GPi) and the substantia nigra pars reticulata (SNr). The GPi/SNr then projects to the thalamus, which
projects back to the cortex. The indirect pathway is the projection of axons from the striatum to the
globus pallidus external segment (GPe), then to the subthalamic nucleus (STN), and finally to GPi/SNr.
The two streams both target the GPi/SNr, but one has a direct projection and the other goes through a
few intervening nuclei. The direct pathway causes the disinhibition of the thalamus (inhibition of one
cell on a target cell that then inhibits the first cell), whereas the indirect pathway causes, or reinforces,
the normal inhibition of the thalamus. The thalamus then can either excite the cortex (as a result of the
direct pathway) or fail to excite the cortex (as a result of the indirect pathway).

The switch between the two pathways is the substantia nigra pars compacta, which projects to the
striatum and releases the neurotransmitter dopamine. Dopamine receptors are either excitatory (D1-
type receptors) or inhibitory (D2-type receptors). The direct pathway is activated by dopamine, and the
indirect pathway is inhibited by dopamine.

When the substantia nigra pars compacta is firing, it signals to the basal nuclei that the body is in an
active state, and movement will be more likely. When the substantia nigra pars compacta is silent, the
body is in a passive state, and movement is inhibited. To illustrate this situation, while a student is sitting
listening to a lecture, the substantia nigra pars compacta would be silent and the student less likely to
get up and walk around. Likewise, while the professor is lecturing, and walking around at the front of the
classroom, the professor’s substantia nigra pars compacta would be active, in keeping with his or her
activity level.

Watch this video to learn about the basal nuclei (also known as the basal ganglia), which have two
pathways that process information within the cerebrum.

As shown in this video, the direct pathway is the shorter pathway through the system that results in
increased activity in the cerebral cortex and increased motor activity. The direct pathway is described as
resulting in “disinhibition” of the thalamus. What does disinhibition mean? What are the two neurons
doing individually to cause this?

EVERYDAY CONNECTIONS: THE MYTH OF LEFT BRAIN/RIGHT BRAIN

There is a persistent myth that people are “right-brained” or “left-brained,” which is an

oversimplification of an important concept about the cerebral hemispheres. There is some lateralization
of function, in which the left side of the brain is devoted to language function and the right side is
devoted to spatial and nonverbal reasoning. Whereas these functions are predominantly associated with
those sides of the brain, there is no monopoly by either side on these functions. Many pervasive
functions, such as language, are distributed globally around the cerebrum. Some of the support for this
misconception has come from studies of split brains.
A drastic way to deal with a rare and devastating neurological condition (intractable epilepsy) is to
separate the two hemispheres of the brain. After sectioning the corpus callosum, a split-brained patient
will have trouble producing verbal responses on the basis of sensory information processed on the right
side of the cerebrum, leading to the idea that the left side is responsible for language function. However,
there are well-documented cases of language functions lost from damage to the right side of the brain.

The deficits seen in damage to the left side of the brain are classified as aphasia, a loss of speech
function; damage on the right side can affect the use of language. Right-side damage can result in a loss
of ability to understand figurative aspects of speech, such as jokes, irony, or metaphors. Nonverbal
aspects of speech can be affected by damage to the right side, such as facial expression or body
language, and right-side damage can lead to a “flat affect” in speech, or a loss of emotional expression in
speech—sounding like a robot when talking.

The Diencephalon

Figure 6. The
Diencephalon. The diencephalon is composed primarily of the thalamus and hypothalamus, which
together define the walls of the third ventricle. The thalami are two elongated, ovoid structures on
either side of the midline that make contact in the middle. The hypothalamus is inferior and anterior
to the thalamus, culminating in a sharp angle to which the pituitary gland is attached.

The diencephalon is the one region of the adult brain that retains its name from embryologic
development. The etymology of the word diencephalon translates to “through brain.” It is the
connection between the cerebrum and the rest of the nervous system, with one exception. The rest of
the brain, the spinal cord, and the PNS all send information to the cerebrum through the diencephalon.
Output from the cerebrum passes through the diencephalon. The single exception is the system
associated with olfaction, or the sense of smell, which connects directly with the cerebrum. In the
earliest vertebrate species, the cerebrum was not much more than olfactory bulbs that received
peripheral information about the chemical environment (to call it smell in these organisms is imprecise
because they lived in the ocean). The diencephalon is deep beneath the cerebrum and constitutes the
walls of the third ventricle. The diencephalon can be described as any region of the brain with
“thalamus” in its name. The two major regions of the diencephalon are the thalamus itself and the
hypothalamus (Figure 6). There are other structures, such as the epithalamus, which contains the pineal
gland, or the subthalamus, which includes the subthalamic nucleus that is part of the basal nuclei.

Thalamus

The thalamus is a collection of nuclei that relay information between the cerebral cortex and the
periphery, spinal cord, or brain stem. All sensory information, except for the sense of smell, passes
through the thalamus before processing by the cortex. Axons from the peripheral sensory organs, or
intermediate nuclei, synapse in the thalamus, and thalamic neurons project directly to the cerebrum. It
is a requisite synapse in any sensory pathway, except for olfaction. The thalamus does not just pass the
information on, it also processes that information. For example, the portion of the thalamus that
receives visual information will influence what visual stimuli are important, or what receives attention.
The cerebrum also sends information down to the thalamus, which usually communicates motor
commands. This involves interactions with the cerebellum and other nuclei in the brain stem. The
cerebrum interacts with the basal nuclei, which involves connections with the thalamus. The primary
output of the basal nuclei is to the thalamus, which relays that output to the cerebral cortex. The cortex
also sends information to the thalamus that will then influence the effects of the basal nuclei.

Hypothalamus

Inferior and slightly anterior to the thalamus is the hypothalamus, the other major region of the
diencephalon. The hypothalamus is a collection of nuclei that are largely involved in regulating
homeostasis. The hypothalamus is the executive region in charge of the autonomic nervous system and
the endocrine system through its regulation of the anterior pituitary gland. Other parts of the
hypothalamus are involved in memory and emotion as part of the limbic system.

Brain Stem
 Figure 7. The Brain
Stem. The brain stem comprises three regions: the midbrain, the pons, and the medulla.

The midbrain and hindbrain (composed of the pons and the medulla) are collectively referred to as the
brain stem (Figure 7). The structure emerges from the ventral surface of the forebrain as a tapering cone
that connects the brain to the spinal cord. Attached to the brain stem, but considered a separate region
of the adult brain, is the cerebellum. The midbrain coordinates sensory representations of the visual,
auditory, and somatosensory perceptual spaces. The pons is the main connection with the cerebellum.
The pons and the medulla regulate several crucial functions, including the cardiovascular and respiratory
systems and rates.

The cranial nerves connect through the brain stem and provide the brain with the sensory input and
motor output associated with the head and neck, including most of the special senses. The major
ascending and descending pathways between the spinal cord and brain, specifically the cerebrum, pass
through the brain stem.

Midbrain

One of the original regions of the embryonic brain, the midbrain is a small region between the thalamus
and pons. It is separated into the tectum and tegmentum, from the Latin words for roof and floor,
respectively. The cerebral aqueduct passes through the center of the midbrain, such that these regions
are the roof and floor of that canal. The tectum is composed of four bumps known as the colliculi
(singular = colliculus), which means “little hill” in Latin. The inferior colliculus is the inferior pair of these
enlargements and is part of the auditory brain stem pathway. Neurons of the inferior colliculus project
to the thalamus, which then sends auditory information to the cerebrum for the conscious perception of
sound. The superior colliculus is the superior pair and combines sensory information about visual space,
auditory space, and somatosensory space. Activity in the superior colliculus is related to orienting the
eyes to a sound or touch stimulus. If you are walking along the sidewalk on campus and you hear
chirping, the superior colliculus coordinates that information with your awareness of the visual location
of the tree right above you. That is the correlation of auditory and visual maps. If you suddenly feel
something wet fall on your head, your superior colliculus integrates that with the auditory and visual
maps and you know that the chirping bird just relieved itself on you. You want to look up to see the
culprit, but do not. The tegmentum is continuous with the gray matter of the rest of the brain stem.
Throughout the midbrain, pons, and medulla, the tegmentum contains the nuclei that receive and send
information through the cranial nerves, as well as regions that regulate important functions such as
those of the cardiovascular and respiratory systems.

Pons

The word pons comes from the Latin word for bridge. It is visible on the anterior surface of the brain
stem as the thick bundle of white matter attached to the cerebellum. The pons is the main connection
between the cerebellum and the brain stem. The bridge-like white matter is only the anterior surface of
the pons; the gray matter beneath that is a continuation of the tegmentum from the midbrain. Gray
matter in the tegmentum region of the pons contains neurons receiving descending input from the
forebrain that is sent to the cerebellum.

Medulla

The medulla is the region known as the myelencephalon in the embryonic brain. The initial portion of
the name, “myel,” refers to the significant white matter found in this region—especially on its exterior,
which is continuous with the white matter of the spinal cord. The tegmentum of the midbrain and pons
continues into the medulla because this gray matter is responsible for processing cranial nerve
information. A diffuse region of gray matter throughout the brain stem, known as the reticular
formation, is related to sleep and wakefulness, such as general brain activity and attention.

The Cerebellum

The cerebellum, as the name suggests, is the “little brain.” It is covered in gyri and sulci like the
cerebrum, and looks like a miniature version of that part of the brain (Figure 8). The cerebellum is
largely responsible for comparing information from the cerebrum with sensory feedback from the
periphery through the spinal cord. It accounts for approximately 10 percent of the mass of the brain.

Figure 8. The Cerebellum. The cerebellum is situated on the posterior surface of the brain stem.
Descending input from the cerebellum enters through the large white matter structure of the pons.
Ascending input from the periphery and spinal cord enters through the fibers of the inferior olive.
Output goes to the midbrain, which sends a descending signal to the spinal cord.

Descending fibers from the cerebrum have branches that connect to neurons in the pons. Those
neurons project into the cerebellum, providing a copy of motor commands sent to the spinal cord.
Sensory information from the periphery, which enters through spinal or cranial nerves, is copied to a
nucleus in the medulla known as the inferior olive. Fibers from this nucleus enter the cerebellum and
are compared with the descending commands from the cerebrum. If the primary motor cortex of the
frontal lobe sends a command down to the spinal cord to initiate walking, a copy of that instruction is
sent to the cerebellum. Sensory feedback from the muscles and joints, proprioceptive information about
the movements of walking, and sensations of balance are sent to the cerebellum through the inferior
olive and the cerebellum compares them. If walking is not coordinated, perhaps because the ground is
uneven or a strong wind is blowing, then the cerebellum sends out a corrective command to
compensate for the difference between the original cortical command and the sensory feedback. The
output of the cerebellum is into the midbrain, which then sends a descending input to the spinal cord to
correct the messages going to skeletal muscles.
The Spinal Cord

The description of the CNS is concentrated on the structures of the brain, but the spinal cord is another
major organ of the system. Whereas the brain develops out of expansions of the neural tube into
primary and then secondary vesicles, the spinal cord maintains the tube structure and is only specialized
into certain regions. As the spinal cord continues to develop in the newborn, anatomical features mark
its surface. The anterior midline is marked by the anterior median fissure, and the posterior midline is
marked by the posterior median sulcus. Axons enter the posterior side through the dorsal (posterior)
nerve root, which marks the posterolateral sulcus on either side. The axons emerging from the anterior
side do so through the ventral (anterior) nerve root. Note that it is common to see the terms dorsal
(dorsal = “back”) and ventral (ventral = “belly”) used interchangeably with posterior and anterior,
particularly in reference to nerves and the structures of the spinal cord. You should learn to be
comfortable with both.

On the whole, the posterior regions are responsible for sensory functions and the anterior regions are
associated with motor functions. This comes from the initial development of the spinal cord, which is
divided into the basal plate and the alar plate. The basal plate is closest to the ventral midline of the
neural tube, which will become the anterior face of the spinal cord and gives rise to motor neurons. The
alar plate is on the dorsal side of the neural tube and gives rise to neurons that will receive sensory input
from the periphery. The length of the spinal cord is divided into regions that correspond to the regions
of the vertebral column. The name of a spinal cord region corresponds to the level at which spinal
nerves pass through the intervertebral foramina.

Immediately adjacent to the brain stem is the cervical region, followed by the thoracic, then the lumbar,
and finally the sacral region. The spinal cord is not the full length of the vertebral column because the
spinal cord does not grow significantly longer after the first or second year, but the skeleton continues
to grow. The nerves that emerge from the spinal cord pass through the intervertebral formina at the
respective levels. As the vertebral column grows, these nerves grow with it and result in a long bundle of
nerves that resembles a horse’s tail and is named the cauda equina. The sacral spinal cord is at the level
of the upper lumbar vertebral bones. The spinal nerves extend from their various levels to the proper
level of the vertebral column.

Gray Horns

In cross-section, the gray matter of the spinal cord has the appearance of an ink-blot test, with the
spread of the gray matter on one side replicated on the other—a shape reminiscent of a bulbous capital
“H.” As shown in Figure 9, the gray matter is subdivided into regions that are referred to as horns.
The posterior horn is responsible for sensory processing. The anterior horn sends out motor signals to
the skeletal muscles. The lateral horn, which is only found in the thoracic, upper lumbar, and sacral
regions, is the central component of the sympathetic division of the autonomic nervous system. Some of
the largest neurons of the spinal cord are the multipolar motor neurons in the anterior horn. The fibers
that cause contraction of skeletal muscles are the axons of these neurons. The motor neuron that
causes contraction of the big toe, for example, is located in the sacral spinal cord. The axon that has to
reach all the way to the belly of that muscle may be a meter in length. The neuronal cell body that
maintains that long fiber must be quite large, possibly several hundred micrometers in diameter, making
it one of the largest cells in the body.

Figure 9. Cross-section of Spinal Cord. The cross-section of a thoracic spinal cord segment shows the
posterior, anterior, and lateral horns of gray matter, as well as the posterior, anterior, and lateral
columns of white matter. LM × 40. (Micrograph provided by the Regents of University of Michigan
Medical School © 2012)

White Columns

Just as the gray matter is separated into horns, the white matter of the spinal cord is separated into
columns. Ascending tracts of nervous system fibers in these columns carry sensory information up to
the brain, whereas descending tracts carry motor commands from the brain. Looking at the spinal cord
longitudinally, the columns extend along its length as continuous bands of white matter. Between the
two posterior horns of gray matter are the posterior columns. Between the two anterior horns, and
bounded by the axons of motor neurons emerging from that gray matter area, are the anterior columns.
The white matter on either side of the spinal cord, between the posterior horn and the axons of the
anterior horn neurons, are the lateral columns. The posterior columns are composed of axons of
ascending tracts. The anterior and lateral columns are composed of many different groups of axons of
both ascending and descending tracts—the latter carrying motor commands down from the brain to the
spinal cord to control output to the periphery.
Circulation and the Central Nervous System

LEARNING OBJECTIVES

 Describe the vessels that supply the CNS with blood

 Name the components of the ventricular system and the regions of the brain in which each is
located

 Explain the production of cerebrospinal fluid and its flow through the ventricles

 Explain how a disruption in circulation would result in a stroke

The CNS is crucial to the operation of the body, and any compromise in the brain and spinal cord can
lead to severe difficulties. The CNS has a privileged blood supply, as suggested by the blood-brain
barrier. The function of the tissue in the CNS is crucial to the survival of the organism, so the contents of
the blood cannot simply pass into the central nervous tissue. To protect this region from the toxins and
pathogens that may be traveling through the blood stream, there is strict control over what can move
out of the general systems and into the brain and spinal cord. Because of this privilege, the CNS needs
specialized structures for the maintenance of circulation. This begins with a unique arrangement of
blood vessels carrying fresh blood into the CNS. Beyond the supply of blood, the CNS filters that blood
into cerebrospinal fluid (CSF), which is then circulated through the cavities of the brain and spinal cord
called ventricles.

Blood Supply to the Brain

 Figure 1. Circle of Willis. The blood supply to
the brain enters through the internal carotid arteries and the vertebral arteries, eventually giving rise
to the circle of Willis.

A lack of oxygen to the CNS can be devastating, and the cardiovascular system has specific regulatory
reflexes to ensure that the blood supply is not interrupted. There are multiple routes for blood to get
into the CNS, with specializations to protect that blood supply and to maximize the ability of the brain to
get an uninterrupted perfusion.

Arterial Supply
The major artery carrying recently oxygenated blood away from the heart is the aorta. The very first
branches off the aorta supply the heart with nutrients and oxygen. The next branches give rise to
the common carotid arteries, which further branch into the internal carotid arteries. The external
carotid arteries supply blood to the tissues on the surface of the cranium. The bases of the common
carotids contain stretch receptors that immediately respond to the drop in blood pressure upon
standing. The orthostatic reflex is a reaction to this change in body position, so that blood pressure is
maintained against the increasing effect of gravity (orthostatic means “standing up”). Heart rate
increases—a reflex of the sympathetic division of the autonomic nervous system—and this raises blood
pressure.

The internal carotid artery enters the cranium through the carotid canal in the temporal bone. A second
set of vessels that supply the CNS are the vertebral arteries, which are protected as they pass through
the neck region by the transverse foramina of the cervical vertebrae. The vertebral arteries enter the
cranium through the foramen magnum of the occipital bone.

Branches off the left and right vertebral arteries merge into the anterior spinal artery supplying the
anterior aspect of the spinal cord, found along the anterior median fissure. The two vertebral arteries
then merge into the basilar artery, which gives rise to branches to the brain stem and cerebellum. The
left and right internal carotid arteries and branches of the basilar artery all become the circle of Willis, a
confluence of arteries that can maintain perfusion of the brain even if narrowing or a blockage limits
flow through one part (Figure 1).

Watch this animation to see how blood flows to the brain and passes through the circle of Willis before
being distributed through the cerebrum. The circle of Willis is a specialized arrangement of arteries that
ensure constant perfusion of the cerebrum even in the event of a blockage of one of the arteries in the
circle. The animation shows the normal direction of flow through the circle of Willis to the middle
cerebral artery. Where would the blood come from if there were a blockage just posterior to the middle
cerebral artery on the left?

Venous Return

After passing through the CNS, blood returns to the circulation through a series of dural sinuses and
veins (Figure 2). The superior sagittal sinus runs in the groove of the longitudinal fissure, where it
absorbs CSF from the meninges. The superior sagittal sinus drains to the confluence of sinuses, along
with the occipital sinuses and straight sinus, to then drain into the transverse sinuses. The transverse
sinuses connect to the sigmoid sinuses, which then connect to the jugular veins. From there, the blood
continues toward the heart to be pumped to the lungs for reoxygenation.
Figure 2. Dural Sinuses and Veins. Blood drains from the brain through a series of sinuses that connect
to the jugular veins.

Protective Coverings of the Brain and Spinal Cord

The outer surface of the CNS is covered by a series of membranes composed of connective tissue called
the meninges, which protect the brain. The dura mater is a thick fibrous layer and a strong protective
sheath over the entire brain and spinal cord. It is anchored to the inner surface of the cranium and
vertebral cavity. The arachnoid mater is a membrane of thin fibrous tissue that forms a loose sac around
the CNS. Beneath the arachnoid is a thin, filamentous mesh called the arachnoid trabeculae, which
looks like a spider web, giving this layer its name. Directly adjacent to the surface of the CNS is the pia
mater, a thin fibrous membrane that follows the convolutions of gyri and sulci in the cerebral cortex and
fits into other grooves and indentations (Figure 3).
Figure 3. Meningeal Layers of Superior Sagittal Sinus. The layers of the meninges in the longitudinal
fissure of the superior sagittal sinus are shown, with the dura mater adjacent to the inner surface of
the cranium, the pia mater adjacent to the surface of the brain, and the arachnoid and subarachnoid
space between them. An arachnoid villus is shown emerging into the dural sinus to allow CSF to filter
back into the blood for drainage.

Dura Mater

Like a thick cap covering the brain, the dura mater is a tough outer covering. The name comes from the
Latin for “tough mother” to represent its physically protective role. It encloses the entire CNS and the
major blood vessels that enter the cranium and vertebral cavity. It is directly attached to the inner
surface of the bones of the cranium and to the very end of the vertebral cavity. There are infoldings of
the dura that fit into large crevasses of the brain. Two infoldings go through the midline separations of
the cerebrum and cerebellum; one forms a shelf-like tent between the occipital lobes of the cerebrum
and the cerebellum, and the other surrounds the pituitary gland. The dura also surrounds and supports
the venous sinuses.

Arachnoid Mater
The middle layer of the meninges is the arachnoid, named for the spider-web–like trabeculae between it
and the pia mater. The arachnoid defines a sac-like enclosure around the CNS. The trabeculae are found
in the subarachnoid space, which is filled with circulating CSF. The arachnoid emerges into the dural
sinuses as the arachnoid granulations, where the CSF is filtered back into the blood for drainage from
the nervous system. The subarachnoid space is filled with circulating CSF, which also provides a liquid
cushion to the brain and spinal cord. Similar to clinical blood work, a sample of CSF can be withdrawn to
find chemical evidence of neuropathology or metabolic traces of the biochemical functions of nervous
tissue.

Pia Mater

The outer surface of the CNS is covered in the thin fibrous membrane of the pia mater. It is thought to
have a continuous layer of cells providing a fluid-impermeable membrane. The name pia mater comes
from the Latin for “tender mother,” suggesting the thin membrane is a gentle covering for the brain. The
pia extends into every convolution of the CNS, lining the inside of the sulci in the cerebral and cerebellar
cortices. At the end of the spinal cord, a thin filament extends from the inferior end of CNS at the upper
lumbar region of the vertebral column to the sacral end of the vertebral column. Because the spinal cord
does not extend through the lower lumbar region of the vertebral column, a needle can be inserted
through the dura and arachnoid layers to withdraw CSF. This procedure is called a lumbar puncture and
avoids the risk of damaging the central tissue of the spinal cord. Blood vessels that are nourishing the
central nervous tissue are between the pia mater and the nervous tissue.

DISORDERS OF THE MENINGES

Meningitis is an inflammation of the meninges, the three layers of fibrous membrane that surround the
CNS. Meningitis can be caused by infection by bacteria or viruses. The particular pathogens are not
special to meningitis; it is just an inflammation of that specific set of tissues from what might be a
broader infection. Bacterial meningitis can be caused by Streptococcus, Staphylococcus, or the
tuberculosis pathogen, among many others. Viral meningitis is usually the result of common
enteroviruses (such as those that cause intestinal disorders), but may be the result of the herpes virus or
West Nile virus. Bacterial meningitis tends to be more severe.

The symptoms associated with meningitis can be fever, chills, nausea, vomiting, light sensitivity,
soreness of the neck, or severe headache. More important are the neurological symptoms, such as
changes in mental state (confusion, memory deficits, and other dementia-type symptoms). A serious risk
of meningitis can be damage to peripheral structures because of the nerves that pass through the
meninges. Hearing loss is a common result of meningitis.

The primary test for meningitis is a lumbar puncture. A needle inserted into the lumbar region of the
spinal column through the dura mater and arachnoid membrane into the subarachnoid space can be
used to withdraw the fluid for chemical testing. Fatality occurs in 5 to 40 percent of children and 20 to
50 percent of adults with bacterial meningitis. Treatment of bacterial meningitis is through antibiotics,
but viral meningitis cannot be treated with antibiotics because viruses do not respond to that type of
drug. Fortunately, the viral forms are milder.

Watch this video that describes the procedure known as the lumbar puncture, a medical procedure used
to sample the CSF. Because of the anatomy of the CNS, it is a relative safe location to insert a needle.
Why is the lumbar puncture performed in the lower lumbar area of the vertebral column?

The Ventricular System

Cerebrospinal fluid (CSF) circulates throughout and around the CNS. In other tissues, water and small
molecules are filtered through capillaries as the major contributor to the interstitial fluid. In the brain,
CSF is produced in special structures to perfuse through the nervous tissue of the CNS and is continuous
with the interstitial fluid. Specifically, CSF circulates to remove metabolic wastes from the interstitial
fluids of nervous tissues and return them to the blood stream. The ventricles are the open spaces within
the brain where CSF circulates. In some of these spaces, CSF is produced by filtering of the blood that is
performed by a specialized membrane known as a choroid plexus. The CSF circulates through all of the
ventricles to eventually emerge into the subarachnoid space where it will be reabsorbed into the blood.

The Ventricles

There are four ventricles within the brain, all of which developed from the original hollow space within
the neural tube, the central canal. The first two are named the lateral ventricles and are deep within
the cerebrum. These ventricles are connected to the third ventricle by two openings called
the interventricular foramina. The third ventricle is the space between the left and right sides of the
diencephalon, which opens into the cerebral aqueduct that passes through the midbrain. The aqueduct
opens into the fourth ventricle, which is the space between the cerebellum and the pons and upper
medulla (Figure 4).
Figure 4. Cerebrospinal Fluid Circulation. The choroid plexus in the four ventricles produce CSF, which
is circulated through the ventricular system and then enters the subarachnoid space through the
median and lateral apertures. The CSF is then reabsorbed into the blood at the arachnoid
granulations, where the arachnoid membrane emerges into the dural sinuses.

As the telencephalon enlarges and grows into the cranial cavity, it is limited by the space within the
skull. The telencephalon is the most anterior region of what was the neural tube, but cannot grow past
the limit of the frontal bone of the skull. Because the cerebrum fits into this space, it takes on a C-
shaped formation, through the frontal, parietal, occipital, and finally temporal regions.

The space within the telencephalon is stretched into this same C-shape. The two ventricles are in the left
and right sides, and were at one time referred to as the first and second ventricles. The interventricular
foramina connect the frontal region of the lateral ventricles with the third ventricle. The third ventricle is
the space bounded by the medial walls of the hypothalamus and thalamus. The two thalami touch in the
center in most brains as the massa intermedia, which is surrounded by the third ventricle. The cerebral
aqueduct opens just inferior to the epithalamus and passes through the midbrain. The tectum and
tegmentum of the midbrain are the roof and floor of the cerebral aqueduct, respectively. The aqueduct
opens up into the fourth ventricle. The floor of the fourth ventricle is the dorsal surface of the pons and
upper medulla (that gray matter making a continuation of the tegmentum of the midbrain). The fourth
ventricle then narrows into the central canal of the spinal cord.

The ventricular system opens up to the subarachnoid space from the fourth ventricle. The single median
aperture and the pair of lateral apertures connect to the subarachnoid space so that CSF can flow
through the ventricles and around the outside of the CNS. Cerebrospinal fluid is produced within the
ventricles by a type of specialized membrane called a choroid plexus. Ependymal cells (one of the types
of glial cells described in the introduction to the nervous system) surround blood capillaries and filter
the blood to make CSF. The fluid is a clear solution with a limited amount of the constituents of blood. It
is essentially water, small molecules, and electrolytes. Oxygen and carbon dioxide are dissolved into the
CSF, as they are in blood, and can diffuse between the fluid and the nervous tissue.

Cerebrospinal Fluid Circulation

The choroid plexuses are found in all four ventricles. Observed in dissection, they appear as soft, fuzzy
structures that may still be pink, depending on how well the circulatory system is cleared in preparation
of the tissue. The CSF is produced from components extracted from the blood, so its flow out of the
ventricles is tied to the pulse of cardiovascular circulation. From the lateral ventricles, the CSF flows into
the third ventricle, where more CSF is produced, and then through the cerebral aqueduct into the fourth
ventricle where even more CSF is produced.

A very small amount of CSF is filtered at any one of the plexuses, for a total of about 500 milliliters daily,
but it is continuously made and pulses through the ventricular system, keeping the fluid moving. From
the fourth ventricle, CSF can continue down the central canal of the spinal cord, but this is essentially a
cul-de-sac, so more of the fluid leaves the ventricular system and moves into the subarachnoid space
through the median and lateral apertures. Within the subarachnoid space, the CSF flows around all of
the CNS, providing two important functions.

Watch this animation that shows the flow of CSF through the brain and spinal cord, and how it
originates from the ventricles and then spreads into the space within the meninges, where the fluids
then move into the venous sinuses to return to the cardiovascular circulation. What are the structures
that produce CSF and where are they found? How are the structures indicated in this animation?

As with elsewhere in its circulation, the CSF picks up metabolic wastes from the nervous tissue and
moves it out of the CNS. It also acts as a liquid cushion for the brain and spinal cord. By surrounding the
entire system in the subarachnoid space, it provides a thin buffer around the organs within the strong,
protective dura mater. The arachnoid granulations are outpocketings of the arachnoid membrane into
the dural sinuses so that CSF can be reabsorbed into the blood, along with the metabolic wastes. From
the dural sinuses, blood drains out of the head and neck through the jugular veins, along with the rest of
the circulation for blood, to be reoxygenated by the lungs and wastes to be filtered out by the kidneys
(Table 1).
Table 1. Components of CSF Circulation

Lateral Third Cerebral

Fourth ventricle
ventricles ventricle aqueduct

Between pons/upper medulla and

Location in CNS Cerebrum Diencephalon Midbrain
cerebellum

Blood vessel Choroid

Choroid plexus None Choroid plexus
structure plexus
The Peripheral Nervous System

LEARNING OBJECTIVES

 Describe the structures found in the PNS

 Distinguish between somatic and autonomic structures, including the special peripheral
structures of the enteric nervous system

 Name the twelve cranial nerves and explain the functions associated with each

 Describe the sensory and motor components of spinal nerves and the plexuses that they pass
through

The PNS is not as contained as the CNS because it is defined as everything that is not the CNS. Some
peripheral structures are incorporated into the other organs of the body. In describing the anatomy of
the PNS, it is necessary to describe the common structures, the nerves and the ganglia, as they are
found in various parts of the body. Many of the neural structures that are incorporated into other
organs are features of the digestive system; these structures are known as the enteric nervous
system and are a special subset of the PNS.

Ganglia

Figure 1. Dorsal Root
Ganglion. The cell bodies of sensory neurons, which are unipolar neurons by shape, are seen in this
photomicrograph. Also, the fibrous region is composed of the axons of these neurons that are passing
through the ganglion to be part of the dorsal nerve root (tissue source: canine). LM × 40. (Micrograph
provided by the Regents of University of Michigan Medical School © 2012)
A ganglion is a group of neuron cell bodies in the periphery. Ganglia can be categorized, for the most
part, as either sensory ganglia or autonomic ganglia, referring to their primary functions. The most
common type of sensory ganglion is a dorsal (posterior) root ganglion. These ganglia are the cell bodies
of neurons with axons that are sensory endings in the periphery, such as in the skin, and that extend
into the CNS through the dorsal nerve root. The ganglion is an enlargement of the nerve root. Under
microscopic inspection, it can be seen to include the cell bodies of the neurons, as well as bundles of
fibers that are the posterior nerve root (Figure 1). The cells of the dorsal root ganglion are unipolar cells,
classifying them by shape. Also, the small round nuclei of satellite cells can be seen surrounding—as if
they were orbiting—the neuron cell bodies.

Another type of sensory ganglion is a cranial nerve ganglion. This is analogous to the dorsal root
ganglion, except that it is associated with a cranial nerve instead of a spinal nerve. The roots of cranial
nerves are within the cranium, whereas the ganglia are outside the skull. For example, the trigeminal
ganglion is superficial to the temporal bone whereas its associated nerve is attached to the mid-pons
region of the brain stem. The neurons of cranial nerve ganglia are also unipolar in shape with associated
satellite cells. The other major category of ganglia are those of the autonomic nervous system, which is
divided into the sympathetic and parasympathetic nervous systems.

Figure 2. Spinal Cord and

Root Ganglion. The slide includes both a cross-section of the lumbar spinal cord and a section of the
dorsal root ganglion (see also Figure 1) (tissue source: canine). LM × 1600. (Micrograph provided by
the Regents of University of Michigan Medical School © 2012)

The sympathetic chain ganglia constitute a row of ganglia along the vertebral column that receive
central input from the lateral horn of the thoracic and upper lumbar spinal cord. Superior to the chain
ganglia are three paravertebral ganglia in the cervical region. Three other autonomic ganglia that are
related to the sympathetic chain are the prevertebral ganglia, which are located outside of the chain
but have similar functions. They are referred to as prevertebral because they are anterior to the
vertebral column. The neurons of these autonomic ganglia are multipolar in shape, with dendrites
radiating out around the cell body where synapses from the spinal cord neurons are made. The neurons
of the chain, paravertebral, and prevertebral ganglia then project to organs in the head and neck,
thoracic, abdominal, and pelvic cavities to regulate the sympathetic aspect of homeostatic mechanisms.

Another group of autonomic ganglia are the terminal ganglia that receive input from cranial nerves or
sacral spinal nerves and are responsible for regulating the parasympathetic aspect of homeostatic
mechanisms. These two sets of ganglia, sympathetic and parasympathetic, often project to the same
organs—one input from the chain ganglia and one input from a terminal ganglion—to regulate the
overall function of an organ. For example, the heart receives two inputs such as these; one increases
heart rate, and the other decreases it.

The terminal ganglia that receive input from cranial nerves are found in the head and neck, as well as
the thoracic and upper abdominal cavities, whereas the terminal ganglia that receive sacral input are in
the lower abdominal and pelvic cavities. Terminal ganglia below the head and neck are often
incorporated into the wall of the target organ as a plexus. A plexus, in a general sense, is a network of
fibers or vessels. This can apply to nervous tissue (as in this instance) or structures containing blood
vessels (such as a choroid plexus). For example, the enteric plexus is the extensive network of axons and
neurons in the wall of the small and large intestines. The enteric plexus is actually part of the enteric
nervous system, along with the gastric plexuses and the esophageal plexus. Though the enteric nervous
system receives input originating from central neurons of the autonomic nervous system, it does not
require CNS input to function. In fact, it operates independently to regulate the digestive system.

View the University of Michigan WebScope to explore the tissue sample in greater detail. If you zoom in
on the dorsal root ganglion, you can see smaller satellite glial cells surrounding the large cell bodies of
the sensory neurons. From what structure do satellite cells derive during embryologic development?

Nerves

Bundles of axons in the PNS are referred to as nerves. These structures in the periphery are different
than the central counterpart, called a tract. Nerves are composed of more than just nervous tissue. They
have connective tissues invested in their structure, as well as blood vessels supplying the tissues with
nourishment. The outer surface of a nerve is a surrounding layer of fibrous connective tissue called
the epineurium. Within the nerve, axons are further bundled into fascicles, which are each surrounded
by their own layer of fibrous connective tissue called perineurium. Finally, individual axons are
surrounded by loose connective tissue called the endoneurium (Figure 3).
Figure 3. Nerve Structure. The structure of a nerve is organized by the layers of connective tissue on
the outside, around each fascicle, and surrounding the individual nerve fibers (tissue source: simian).
LM × 40. (Micrograph provided by the Regents of University of Michigan Medical School © 2012)

These three layers are similar to the connective tissue sheaths for muscles. Nerves are associated with
the region of the CNS to which they are connected, either as cranial nerves connected to the brain or
spinal nerves connected to the spinal cord.
Figure 4. Close-Up of Nerve Trunk. Zoom in on this slide of a nerve trunk to examine the endoneurium,
perineurium, and epineurium in greater detail (tissue source: simian). LM × 1600. (Micrograph
provided by the Regents of University of Michigan Medical School © 2012)

View the University of Michigan WebScope to explore the tissue sample in greater detail. With what
structures in a skeletal muscle are the endoneurium, perineurium, and epineurium comparable?

Cranial Nerves

The nerves attached to the brain are the cranial nerves, which are primarily responsible for the sensory
and motor functions of the head and neck (one of these nerves targets organs in the thoracic and
abdominal cavities as part of the parasympathetic nervous system). There are twelve cranial nerves,
which are designated CNI through CNXII for “Cranial Nerve,” using Roman numerals for 1 through 12.
They can be classified as sensory nerves, motor nerves, or a combination of both, meaning that the
axons in these nerves originate out of sensory ganglia external to the cranium or motor nuclei within the
brain stem. Sensory axons enter the brain to synapse in a nucleus. Motor axons connect to skeletal
muscles of the head or neck. Three of the nerves are solely composed of sensory fibers; five are strictly
motor; and the remaining four are mixed nerves.

Learning the cranial nerves is a tradition in anatomy courses, and students have always used mnemonic
devices to remember the nerve names. A traditional mnemonic is the rhyming couplet, “On Old
Olympus’ Towering Tops/A Finn And German Viewed Some Hops,” in which the initial letter of each
word corresponds to the initial letter in the name of each nerve. The names of the nerves have changed
over the years to reflect current usage and more accurate naming. An exercise to help learn this sort of
information is to generate a mnemonic using words that have personal significance. The names of the
cranial nerves are listed in Table 1 along with a brief description of their function, their source (sensory
ganglion or motor nucleus), and their target (sensory nucleus or skeletal muscle).

Figure 5. The Cranial Nerves

The anatomical arrangement of the roots of the cranial nerves observed from an inferior view of the
brain.

Figure 5 shows where the nerves are located in the brain. The olfactory nerve and optic nerve are
responsible for the sense of smell and vision, respectively. The oculomotor nerve is responsible for eye
movements by controlling four of the extraocular muscles. It is also responsible for lifting the upper
eyelid when the eyes point up, and for pupillary constriction. The trochlear nerve and the abducens
nerve are both responsible for eye movement, but do so by controlling different extraocular muscles.
The trigeminal nerve is responsible for cutaneous sensations of the face and controlling the muscles of
mastication. The facial nerve is responsible for the muscles involved in facial expressions, as well as part
of the sense of taste and the production of saliva. The vestibulocochlear nerve is responsible for the
senses of hearing and balance. The glossopharyngeal nerve is responsible for controlling muscles in the
oral cavity and upper throat, as well as part of the sense of taste and the production of saliva. The vagus
nerve is responsible for contributing to homeostatic control of the organs of the thoracic and upper
abdominal cavities. The spinal accessory nerve is responsible for controlling the muscles of the neck,
along with cervical spinal nerves. The hypoglossal nerve is responsible for controlling the muscles of the
lower throat and tongue.
 Table 1. Cranial Nerves

 Mnemoni
# Name Function (S/M/B) Central connection (nuclei)
c

On I Olfactory Smell (S) Olfactory bulb

Old II Optic Vision (S) Hypothalamus/thalamus/midbrain

Olympus III Oculomotor Eye movements (M) Oculomotor nucleus

Towering IV Trochlear Eye movements (M) Trochlear nucleus

Sensory/motor—face Trigeminal nuclei in the midbrain, pons,

Tops V Trigeminal
(B) and medulla

A VI Abducens Eye movements (M) Abducens nucleus

Facial nucleus, solitary nucleus, superior

Finn VII Facial Motor—face, Taste (B)
salivatory nucleus

Auditory Cochlearn nucleus, Vestibular

And VIII Hearing/balance (S)
(Vestibulocochlear) nucleus/cerebellum

Motor—throat Taste Solitary nucleus, inferior salivatory

German IX Glossopharyngeal
(B) nucleus, nucleus ambiguus

Motor/sensory—
Viewed X Vagus Medulla
viscera (autonomic)

Some XI Spinal Accessory Motor—head and neck Spinal accessory nucleus

  Table 1. Cranial Nerves

 Mnemoni
# Name Function (S/M/B) Central connection (nuclei)
c

(M)

Motor—lower throat
Hops XII Hypoglossal Hypoglossal nucleus
(M)

Three of the cranial nerves also contain autonomic fibers, and a fourth is almost purely a component of
the autonomic system. The oculomotor, facial, and glossopharyngeal nerves contain fibers that contact
autonomic ganglia. The oculomotor fibers initiate pupillary constriction, whereas the facial and
glossopharyngeal fibers both initiate salivation. The vagus nerve primarily targets autonomic ganglia in
the thoracic and upper abdominal cavities.

Visit this site to read about a man who wakes with a headache and a loss of vision. His regular doctor
sent him to an ophthalmologist to the vision loss. The ophthalmologist recognizes a greater problem and
immediately sends him to the emergency room. Once there, the patient undergoes a large battery of
tests, but a definite cause cannot be found. A specialist recognizes the problem as meningitis, but the
question is what caused it originally. How can that be cured? The loss of vision comes from swelling
around the optic nerve, which probably presented as a bulge on the inside of the eye. Why is swelling
related to meningitis going to push on the optic nerve?

Another important aspect of the cranial nerves that lends itself to a mnemonic is the functional role
each nerve plays. The nerves fall into one of three basic groups. They are sensory, motor, or both
(see Table 1). The sentence, “Some Say Marry Money But My Brother Says Brains Beauty Matter More,”
corresponds to the basic function of each nerve.

The first, second, and eighth nerves are purely sensory: the olfactory (CNI), optic (CNII), and
vestibulocochlear (CNVIII) nerves. The three eye-movement nerves are all motor: the oculomotor (CNIII),
trochlear (CNIV), and abducens (CNVI). The spinal accessory (CNXI) and hypoglossal (CNXII) nerves are
also strictly motor. The remainder of the nerves contain both sensory and motor fibers. They are the
trigeminal (CNV), facial (CNVII), glossopharyngeal (CNIX), and vagus (CNX) nerves.

The nerves that convey both are often related to each other. The trigeminal and facial nerves both
concern the face; one concerns the sensations and the other concerns the muscle movements. The
facial and glossopharyngeal nerves are both responsible for conveying gustatory, or taste, sensations as
well as controlling salivary glands. The vagus nerve is involved in visceral responses to taste, namely the
gag reflex. This is not an exhaustive list of what these combination nerves do, but there is a thread of
relation between them.

Spinal Nerves

The nerves connected to the spinal cord are the spinal nerves. The arrangement of these nerves is much
more regular than that of the cranial nerves. All of the spinal nerves are combined sensory and motor
axons that separate into two nerve roots. The sensory axons enter the spinal cord as the dorsal nerve
root. The motor fibers, both somatic and autonomic, emerge as the ventral nerve root. The dorsal root
ganglion for each nerve is an enlargement of the spinal nerve.

There are 31 spinal nerves, named for the level of the spinal cord at which each one emerges. There are
eight pairs of cervical nerves designated C1 to C8, twelve thoracic nerves designated T1 to T12, five pairs
of lumbar nerves designated L1 to L5, five pairs of sacral nerves designated S1 to S5, and one pair of
coccygeal nerves. The nerves are numbered from the superior to inferior positions, and each emerges
from the vertebral column through the intervertebral foramen at its level. The first nerve, C1, emerges
between the first cervical vertebra and the occipital bone. The second nerve, C2, emerges between the
first and second cervical vertebrae. The same occurs for C3 to C7, but C8 emerges between the seventh
cervical vertebra and the first thoracic vertebra. For the thoracic and lumbar nerves, each one emerges
between the vertebra that has the same designation and the next vertebra in the column. The sacral
nerves emerge from the sacral foramina along the length of that unique vertebra.
 Figure 6. Nerve Plexuses of
the Body There are four main nerve plexuses in the human body. The cervical plexus supplies nerves
to the posterior head and neck, as well as to the diaphragm. The brachial plexus supplies nerves to the
arm. The lumbar plexus supplies nerves to the anterior leg. The sacral plexus supplies nerves to the
posterior leg.
Spinal nerves extend outward from the vertebral column to enervate the periphery. The nerves in the
periphery are not straight continuations of the spinal nerves, but rather the reorganization of the axons
in those nerves to follow different courses. Axons from different spinal nerves will come together into
a systemic nerve. This occurs at four places along the length of the vertebral column, each identified as
a nerve plexus, whereas the other spinal nerves directly correspond to nerves at their respective levels.
In this instance, the word plexus is used to describe networks of nerve fibers with no associated cell
bodies. Of the four nerve plexuses, two are found at the cervical level, one at the lumbar level, and one
at the sacral level (Figure 6).

The cervical plexus is composed of axons from spinal nerves C1 through C5 and branches into nerves in
the posterior neck and head, as well as the phrenic nerve, which connects to the diaphragm at the base
of the thoracic cavity. The other plexus from the cervical level is the brachial plexus.

Spinal nerves C4 through T1 reorganize through this plexus to give rise to the nerves of the arms, as the
name brachial suggests. A large nerve from this plexus is the radial nerve from which the axillary
nerve branches to go to the armpit region. The radial nerve continues through the arm and is paralleled
by the ulnar nerve and the median nerve. The lumbar plexus arises from all the lumbar spinal nerves
and gives rise to nerves enervating the pelvic region and the anterior leg. The femoral nerve is one of
the major nerves from this plexus, which gives rise to the saphenous nerve as a branch that extends
through the anterior lower leg.

The sacral plexus comes from the lower lumbar nerves L4 and L5 and the sacral nerves S1 to S4. The
most significant systemic nerve to come from this plexus is the sciatic nerve, which is a combination of
the tibial nerve and the fibular nerve. The sciatic nerve extends across the hip joint and is most
commonly associated with the condition sciatica, which is the result of compression or irritation of the
nerve or any of the spinal nerves giving rise to it.

These plexuses are described as arising from spinal nerves and giving rise to certain systemic nerves, but
they contain fibers that serve sensory functions or fibers that serve motor functions. This means that
some fibers extend from cutaneous or other peripheral sensory surfaces and send action potentials into
the CNS. Those are axons of sensory neurons in the dorsal root ganglia that enter the spinal cord
through the dorsal nerve root. Other fibers are the axons of motor neurons of the anterior horn of the
spinal cord, which emerge in the ventral nerve root and send action potentials to cause skeletal muscles
to contract in their target regions. For example, the radial nerve contains fibers of cutaneous sensation
in the arm, as well as motor fibers that move muscles in the arm. Spinal nerves of the thoracic region, T2
through T11, are not part of the plexuses but rather emerge and give rise to the intercostal
nerves found between the ribs, which articulate with the vertebrae surrounding the spinal nerve.
The Function of Nervous Tissue

LEARNING OBJECTIVES

 Distinguish the major functions of the nervous system: sensation, integration, and response

 List the sequence of events in a simple sensory receptor–motor response pathway

Having looked at the components of nervous tissue, and the basic anatomy of the nervous system, next
comes an understanding of how nervous tissue is capable of communicating within the nervous system.
Before getting to the nuts and bolts of how this works, an illustration of how the components come
together will be helpful.

Imagine you are about to take a shower in the morning before going to school. You have turned on the
faucet to start the water as you prepare to get in the shower. After a few minutes, you expect the water
to be a temperature that will be comfortable to enter. So you put your hand out into the spray of water.
What happens next depends on how your nervous system interacts with the stimulus of the water
temperature and what you do in response to that stimulus. Figure 1 shows one possible path you may
follow.
Figure 1. Testing the Water

Here's a more detailed breakdown of the processes found in Figure 1:

1. The sensory neuron has endings in the skin that sense a stimulus such as water temperature.
The strength of the signal that starts here is dependent on the strength of the stimulus.

2. The graded potential from the sensory endings, if strong enough, will initiate an action potential
at the initial segment of the axon (which is immediately adjacent to the sensory endings in the
skin).

3. The axon of the peripheral sensory neuron enters the spinal cord and contacts another neuron
in the gray matter. The contact is a synapse where another graded potential is caused by the
release of a chemical signal from the axon terminals.
 4. An action potential is initiated at the initial segment of this neuron and travels up the sensory
pathway to a region of the brain called the thalamus. Another synapse passes the information
along to the next neuron.

5. The sensory pathway ends when the signal reaches the cerebral cortex.

6. After integration with neurons in other parts of the cerebral cortex, a motor command is sent
from the precentral gyrus of the frontal cortex.

7. The upper motor neuron sends an action potential down to the spinal cord. The target of the
upper motor neuron is the dendrites of the lower motor neuron in the gray matter of the spinal
cord.

8. The axon of the lower motor neuron emerges from the spinal cord in a nerve and connects to a
muscle through a neuromuscular junction to cause contraction of the target muscle.

Found in the skin of your fingers or toes is a type of sensory receptor that is sensitive to temperature,
called a thermoreceptor. When you place your hand under the shower (Figure 2), the cell membrane of
the thermoreceptors changes its electrical state (voltage).

Figure 2. The Sensory
Input Receptors in the skin sense the temperature of the water.

The amount of change is dependent on the strength of the stimulus (how hot the water is). This is called
a graded potential. If the stimulus is strong, the voltage of the cell membrane will change enough to
generate an electrical signal that will travel down the axon. You have learned about this type of signaling
before, with respect to the interaction of nerves and muscles at the neuromuscular junction. The
voltage at which such a signal is generated is called the threshold, and the resulting electrical signal is
called an action potential. In this example, the action potential travels—a process known
as propagation—along the axon from the axon hillock to the axon terminals and into the synaptic end
bulbs. When this signal reaches the end bulbs, it causes the release of a signaling molecule called
a neurotransmitter.

The neurotransmitter diffuses across the short distance of the synapse and binds to a receptor protein
of the target neuron. When the molecular signal binds to the receptor, the cell membrane of the target
neuron changes its electrical state and a new graded potential begins. If that graded potential is strong
enough to reach threshold, the second neuron generates an action potential at its axon hillock. The
target of this neuron is another neuron in the thalamus of the brain, the part of the CNS that acts as a
relay for sensory information. At another synapse, neurotransmitter is released and binds to its
receptor. The thalamus then sends the sensory information to the cerebral cortex, the outermost layer
of gray matter in the brain, where conscious perception of that water temperature begins.

Figure 3. The Motor Response On

the basis of the sensory input and the integration in the CNS, a motor response is formulated and
executed.
Within the cerebral cortex, information is processed among many neurons, integrating the stimulus of
the water temperature with other sensory stimuli, with your emotional state (you just aren't ready to
wake up; the bed is calling to you), memories (perhaps of the lab notes you have to study before a quiz).
Finally, a plan is developed about what to do, whether that is to turn the temperature up, turn the
whole shower off and go back to bed, or step into the shower. To do any of these things, the cerebral
cortex has to send a command out to your body to move muscles (Figure 3).
A region of the cortex is specialized for sending signals down to the spinal cord for movement.
The upper motor neuron is in this region, called the precentral gyrus of the frontal cortex, which has an
axon that extends all the way down the spinal cord. At the level of the spinal cord at which this axon
makes a synapse, a graded potential occurs in the cell membrane of a lower motor neuron. This second
motor neuron is responsible for causing muscle fibers to contract. In the manner described in the
chapter on muscle tissue, an action potential travels along the motor neuron axon into the periphery.
The axon terminates on muscle fibers at the neuromuscular junction. Acetylcholine is released at this
specialized synapse, which causes the muscle action potential to begin, following a large potential
known as an end plate potential. When the lower motor neuron excites the muscle fiber, it contracts. All
of this occurs in a fraction of a second, but this story is the basis of how the nervous system functions.