Chapter13 Anatomy of the Nervous System

Cerebral Cortex

The cerebrum is covered by a continuous layer of grey matter that wraps around either side of the forebrain—the cerebral cortex. This thin, extensive region of wrinkled grey 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 grey matter to fit into this limited space. If the grey matter of the cortex were peeled off the cerebrum and laid out flat, its surface area would be roughly equal to one square metre.

The folding of the cortex maximizes the amount of grey matter in the cranial cavity. 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 13.2). 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 the 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.

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. All 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

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 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, the direct pathway, and the indirect pathway. 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 can then either excite the cortex (as a result of the direct pathway) or fail to excite the cortex (as a result of the indirect pathway).

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 being processed 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, like body temperature. 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.

Midbrain

The cerebral aqueduct passes through the centre 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.

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 bridgelike white matter is only the anterior surface of the pons; the grey matter beneath that is a continuation of the floor of the cerebral aqueduct from the midbrain. Grey matter in the floor of the cerebral aqueduct region of the pons contains neurons receiving descending input from the forebrain that is sent to the cerebellum.

Medulla

The medulla (also called the medulla oblongata) 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 floor of the cerebral aqueduct of the midbrain and pons continues into the medulla because this grey matter is responsible for processing cranial nerve information. A diffuse region of grey matter throughout the brain stem, known as the reticular formation, is related to sleep and wakefulness, such as general brain activity and attention.

Grey Horns

In cross-section, the grey matter of the spinal cord has the appearance of an ink-blot test, with the spread of the grey matter on one side replicated on the other—a shape reminiscent of a bulbous capital “H.” As shown in Figure 13.6, the grey 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, contains cell bodies of motor neurons of the autonomic nervous system. In the centre of the grey matter is a small space called the central canal; it extends the entire length of the spinal cord and is filled with cerebrospinal fluid. At its superior end, the central canal is continuous with the fourth ventricle (a space that contains cerebrospinal fluid) in the medulla oblongata of the brain.

Some of the largest neurons of the spinal cord are the multipolar motor neurons in the anterior horn. The fibres 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 metre in length. The neuronal cell body that maintains that long fibre must be quite large, possibly several hundred micrometres in diameter, making it one of the largest cells in the body.

White Columns

Just as the grey matter is separated into horns, the white matter of the spinal cord is separated into columns. Ascending tracts of nervous system fibres 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 grey matter are the posterior columns. Between the two anterior horns and bounded by the axons of motor neurons emerging from that grey matter area are the anterior columns. The areas of white matter on either side of the spinal cord, between the posterior horn and the axons of the anterior horn neurons, are the lateralcolumns. 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.

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 (Figure 13.7).

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. 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 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.

Venous Return

After passing through the CNS, blood returns to the circulation through a series of dural sinuses and veins (Figure 13.8). 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.

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 shelflike 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 spiderweb-like trabeculae between it and the pia mater. The arachnoid defines a saclike enclosure around the CNS. The trabeculae are found in the subarachnoid space, which is filled with circulating cerebrospinal fluid (CSF).

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 by the thin fibrous membrane of the pia mater. It is thought to have a continuous layer of cells providing a fluid-impermeable membrane. 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 the 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 nourish the central nervous tissue are between the pia mater and the nervous tissue.

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 right and left 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 13.10). 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 centre of 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 aqueduct opens up into the fourth ventricle. The floor of the fourth ventricle is the dorsal surface of the pons and upper medulla. 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 millilitres 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. As with elsewhere in its circulation, the CSF picks up metabolic wastes from the nervous tissue and moves them 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 13.1).

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 from 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 fibres; five are strictly motor; and the remaining four are mixed nerves.

The names of the cranial nerves are listed in Table 13.2 along with a brief description of their function, their source (sensory ganglion or motor nucleus), and their target (sensory nucleus or skeletal muscle). They are listed here with a brief explanation of each nerve (Figure 13.13).

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 controls 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 (auditory) 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.

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

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 13.2). 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 fibres. They are the trigeminal (CNV), facial (CNVII), glossopharyngeal (CNIX), and vagus (CNX) nerves.

The nerves that convey both are often related to one another. 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 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 fibres, 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 8 pairs of cervical nerves designated C1 to C8, 12 thoracic nerves designated T1 to T12, 5 pairs of lumbar nerves designated L1 to L5, 5 pairs of sacral nerves designated S1 to S5, and 1 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.

Spinal nerves extend outward from the vertebral column to innervate 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 fibres 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 13.14). 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.

These plexuses are described as arising from spinal nerves and giving rise to certain systemic nerves, but they contain fibres that serve sensory functions or fibres that serve motor functions. This means that some fibres 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 fibres 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 fibres of cutaneous sensation in the arm, as well as motor fibres 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.