Neurology 3
Neurology 3
The nervous system is organized into two main subdivisions: the central nervous system and the peripheral
nervous system. Neurology deals with normal functioning and disorders of the nervous system. A neurologist
is a physician who diagnoses and treats disorders of the nervous system.
The central nervous system (CNS) consists of the brain and spinal cord. The brain is the part of the CNS that
is located in the skull and contains about 85 billion neurons. The spinal cord is connected to the brain
through the foramen magnum of the occipital bone and is encircled by the bones of the vertebral column.
The peripheral nervous system (PNS) consists of all nervous tissue outside the CNS. Components of the PNS
include nerves and sensory receptors. A nerve is a bundle of hundreds to thousands of axons plus associated
connective tissue and blood vessels that lies outside the brain and spinal cord.
Twelve pairs of cranial nerves emerge from the brain and thirty-one pairs of spinal nerves emerge from the
spinal cord. Each nerve follows a defined path and serves a specific region of the body. The term sensory
receptor refers to a structure of the nervous system that monitors changes in the external or internal
environment.
The PNS is divided into:
1. sensory or afferent division
conveys input into the CNS from sensory receptors in the body. This division provides the CNS with
sensory information about the somatic senses (tactile, thermal, pain, and proprioceptive sensations)
and special senses (smell, taste, vision, hearing, and equilibrium).
2. motor or efferent division
conveys output from the CNS to effectors (muscles and glands). This division is further subdivided
into a somatic nervous system and an autonomic nervous system.
- The somatic nervous system (SNS) conveys output from the CNS to skeletal muscles only.
Because its motor responses can be consciously controlled, the action of this part of the PNS is
voluntary.
- The autonomic nervous system (ANS) conveys output from the CNS to smooth muscle, cardiac
muscle, and glands. Because its motor responses are not normally under conscious control, the
action of the ANS is involuntary.
The ANS is comprised of two main branches, the sympathetic nervous system and the
parasympathetic nervous system. With a few exceptions, effectors receive innervation from both
of these branches, and usually the two branches have opposing actions.
In general, the parasympathetic nervous system takes care of “rest-and-digest” activities, and the
sympathetic nervous system helps support exercise or emergency actions—the so-called “fight-
or-flight” responses. A third branch of the autonomic nervous system is the enteric nervous
system (ENS), an extensive network of over 100 million neurons confined to the wall of the
gastrointestinal (GI) tract. The ENS helps regulate the activity of the smooth muscle and glands
of the GI tract. Although the ENS can function independently, it communicates with and is
regulated by the other branches of the ANS.
The nervous system carries out a complex array of tasks. These diverse activities can be grouped into three
basic functions:
Sensory function
Sensory receptors detect internal stimuli or external stimuli. This sensory information is then carried
into the brain and spinal cord through cranial and spinal nerves.
Integrative function
The nervous system processes sensory information by analyzing it and making decisions for
appropriate responses—an activity known as integration.
Motor function
Once sensory information is integrated, the nervous system may elicit an appropriate motor response
by activating effectors (muscles and glands) through cranial and spinal nerves. Stimulation of the
effectors causes muscles to contract and glands to secrete.
Neurons
neurons possess electrical excitability, the ability to respond to a stimulus and convert it into an action
potential. A stimulus is any change in the environment that is strong enough to initiate an action potential. An
action potential (nerve impulse) is an electrical signal that propagates (travels) along the surface of the
membrane of a neuron. It begins and travels due to the movement of ions (such as sodium and potassium)
between interstitial fluid and the inside of a neuron through specific ion channels in its plasma membrane.
Once begun, a nerve impulse travels rapidly and at a constant strength. Some neurons are tiny and propagate
impulses over a short distance (less than 1 mm) within the CNS. Others are the longest cells in the body.
Most neurons have three parts:
1. Cell body
contains a nucleus surrounded by cytoplasm that includes typical cellular organelles such as
lysosomes, mitochondria, and a Golgi complex. Neuronal cell bodies also contain free ribosomes
and prominent clusters of rough endoplasmic reticulum, termed Nissl bodies. The ribosomes are the
sites of protein synthesis. Newly synthesized proteins produced by Nissl bodies are used to replace
cellular components, as material for growth of neurons, and to regenerate damaged axons in the
PNS.
Aging neurons also contain lipofuscin, a pigment that occurs as clumps of yellowish brown granules
in the cytoplasm.
A collection of neuron cell bodies outside the CNS is called a ganglion (ganglia is the plural).
A nerve fiber is a general term for any neuronal process (extension) that emerges from the cell body
of a neuron. Most neurons have two kinds of processes: multiple dendrites and a single axon.
2. Dendrites
are the receiving or input portions of a neuron. The plasma membranes of dendrites (and cell bodies)
contain numerous receptor sites for binding chemical messengers from other cells. Dendrites usually
are short, tapering, and highly branched. In many neurons the dendrites form a tree-shaped array of
processes extending from the cell body. Their cytoplasm contains Nissl bodies, mitochondria, and
other organelles.
3. Axon
propagates nerve impulses toward another neuron, a muscle fiber, or a gland cell. An axon is a long,
thin, cylindrical projection that often joins to the cell body at a cone-shaped elevation called the axon
hillock.
The part of the axon closest to the axon hillock is the initial
segment. In most neurons, nerve impulses arise at the junction
of the axon hillock and the initial segment, an area called the
trigger zone, from which they travel along the axon to their
destination. An axon contains mitochondria, microtubules, and
neurofibrils. Because rough endoplasmic reticulum is not
present, protein synthesis does not occur in the axon. The
cytoplasm of an axon, called axoplasm, is surrounded by a
plasma membrane known as the axolemma.
Along the length of an axon, side branches called axon
collaterals may branch off, typically at a right angle to the axon.
The axon and its collaterals end by dividing into many fine
processes called axon terminals.
The site of communication between two neurons or between a
neuron and an effector cell is called a synapse. The tips of some
axon terminals swell into bulb-shaped structures called synaptic
end bulbs; others exhibit a string of swollen bumps called
varicosities. Both synaptic end bulbs and varicosities
contain many tiny membrane-enclosed sacs called synaptic
vesicles that store a chemical called a neurotransmitter.
Both structural and functional features are used to classify the various neurons in the body. Structurally,
neurons are classified according to the number of processes extending from the cell body (multipolar
neurons, bipolar neurons or unipolar neurons).
Functionally, neurons are classified according to the direction in which the nerve impulse (action potential) is
conveyed with respect to the CNS.
- Sensory neurons or afferent neurons either contain sensory receptors at their distal ends
(dendrites) or are located just after sensory receptors that are separate cells. Once an appropriate
stimulus activates a sensory receptor, the sensory neuron forms an action potential in its axon
and the action potential is conveyed into the CNS through cranial or spinal nerves.
- Motor neurons or efferent neurons convey action potentials away from the CNS to effectors
(muscles and glands) in the periphery (PNS) through cranial or spinal nerves
- Interneurons or association neurons are mainly located within the CNS between sensory and
motor neurons. Interneurons integrate (process) incoming sensory information from sensory
neurons and then elicit a motor response by activating the appropriate motor neurons.
Neuroglia
Neuroglia or glia make up about half the volume of the CNS. Generally, neuroglia are smaller than neurons,
and they are 5 to 25 times more numerous. In contrast to neurons, glia do not generate or propagate action
potentials, and they can multiply and divide in the mature nervous system. In cases of injury or disease,
neuroglia multiply to fill in the spaces formerly occupied by neurons. Brain tumors derived from glia, called
gliomas, tend to be highly malignant and to grow rapidly. Of the six types of neuroglia, four – astrocytes,
oligodendrocytes, microglia, and ependymal cells – are found only in the CNS. The remaining two types –
Schwann cells and satellite cells – are present in the PNS.
Neuroglia of the CNS can be classified on the basis of size, cytoplasmic processes, and intracellular
organization into four types:
Astrocytes
These star-shaped cells have many processes and are the largest and most numerous of the neuroglia.
There are two types of astrocytes. Protoplasmic astrocytes have many short branching processes and
are found in gray matter. Fibrous astrocytes have many long unbranched processes and are located
mainly in white matter. The processes of astrocytes make contact with blood capillaries, neurons,
and the pia mater.
Processes of astrocytes wrapped around blood capillaries isolate neurons of the CNS from various
potentially harmful substances in blood by secreting chemicals that maintain the unique selective
permeability characteristics of the endothelial cells of the capillaries. In the embryo, astrocytes
secrete chemicals that appear to regulate the growth, migration, and interconnection among neurons
in the brain. Astrocytes help to maintain the appropriate chemical environment for the generation of
nerve impulses.
Oligodendrocytes
These resemble astrocytes but are smaller and contain fewer processes. Processes of
oligodendrocytes
are responsible for forming and maintaining the myelin sheath around CNS axons. As you will see
shortly, the myelin sheath is a multilayered lipid and protein covering around some axons that
insulates them and increases the speed of nerve impulse conduction. Such axons are said to be
myelinated.
Microglia
These neuroglia are small cells with slender processes that give off numerous spinelike projections.
Microglial cells or microglia function as phagocytes. Like tissue macrophages, they remove cellular
debris formed during normal development of the nervous system and phagocytize microbes and
damaged nervous tissue.
Ependymal cells
cuboidal to columnar cells arranged in a single layer that possess microvilli and cilia. These cells line
the ventricles of the brain and central canal of the spinal cord (spaces filled with cerebrospinal fluid,
which protects and nourishes the brain and spinal cord). Functionally, ependymal cells produce,
possibly monitor, and assist in the circulation of cerebrospinal fluid.
Neuroglia of the PNS completely surround axons and cell bodies. The two types of glial cells in the PNS are:
Schwann cells
These cells encircle PNS axons. Like oligodendrocytes, they form the myelin sheath around axons. A
single oligodendrocyte myelinates several axons, but each Schwann cell myelinates a single axon. A
single Schwann cell can also enclose as many as 20 or more unmyelinated axons (axons that lack a
myelin sheath). Schwann cells participate in axon regeneration, which is more easily accomplished
in the PNS than in the CNS.
Satellite cells
These flat cells surround the cell bodies of neurons of PNS ganglia*. Besides providing structural
support, satellite cells regulate the exchanges of materials between neuronal cell bodies and
interstitial fluid.
*a ganglion (plural is ganglia) refers to a cluster of neuronal cell bodies located in the PNS. As mentioned
earlier, ganglia are closely associated with cranial and spinal nerves. By contrast, a nucleus is a cluster of
neuronal cell bodies located in the CNS.
nerve is a bundle of axons that is located in the PNS. Cranial nerves connect the brain to the periphery,
whereas spinal nerves connect the spinal cord to the periphery. A tract is a bundle of axons that is located in
the CNS. Tracts interconnect neurons in the spinal cord and brain.
Myelination
axons surrounded by a multilayered lipid and protein covering, called the myelin sheath, are said to be
myelinated. The sheath electrically insulates the axon of a neuron and increases the speed of nerve impulse
conduction. Axons without such a covering are said to be unmyelinated.
Two types of neuroglia produce myelin sheaths: Schwann cells (in the PNS) and oligodendrocytes (in the
CNS). Schwann cells begin to form myelin sheaths around axons during fetal development.
The outer nucleated cytoplasmic layer of the Schwann cell, which encloses the myelin sheath, is the
neurolemma (sheath of Schwann). A neurolemma is found only around axons in the PNS. When an axon is
injured, the neurolemma aids regeneration by forming a regeneration tube that guides and stimulates
regrowth of the axon. Gaps in the myelin sheath, called nodes of Ranvier, appear at intervals along the axon.
Each Schwann cell wraps one axon segment between two nodes.
In the CNS, an oligodendrocyte myelinates parts of several axons. A neurolemma is not present, however,
because the oligodendrocyte cell body and nucleus do not envelop the axon. Nodes of Ranvier are present,
but they are fewer in number. Axons in the CNS display little regrowth after injury. This is thought to be due,
in part, to the absence of a neurolemma, and in part to an inhibitory influence exerted by the
oligodendrocytes on axon regrowth. The amount of myelin increases from birth to maturity, and its presence
greatly increases the speed of nerve impulse conduction.
Graded potential
A graded potential is a small deviation from the resting membrane potential that makes the membrane either
more polarized (inside more negative) or less polarized (inside less negative). When the response makes the
membrane more polarized (inside more negative), it is termed a hyperpolarizing graded potential. When the
response makes the membrane less polarized (inside less negative), it is termed a depolarizing graded
potential. A graded potential occurs when a stimulus causes mechanically-gated or ligand-gated channels to
open or close in an excitable cell’s plasma membrane.
To say that these electrical signals are graded means that they vary in amplitude (size), depending on the
strength of the stimulus. They are larger or smaller depending on how many ligand-gated or mechanically-
gated channels have opened (or closed) and how long each remains open. The opening or closing of these ion
channels alters the flow of specific ions across the membrane, producing a flow of current that is localized,
which means that it spreads to adjacent regions along the plasma membrane in either direction from the
stimulus source for a short distance and then gradually dies out.
This mode of travel by which graded potentials die out as they spread along the membrane is known as
decremental conduction. Because they die out within a few millimeters of their point of origin, graded
potentials are useful for short-distance communication only.
Action potential
An action potential (AP) or impulse is a sequence of rapidly occurring events that decrease and reverse the
membrane potential and then eventually restore it to the resting state. An action potential has two main
phases:
depolarizing phase
negative membrane potential becomes less negative, reaches zero, and then becomes positive
repolarizing phase
the membrane potential is restored to the resting state of −70 mV
Following the repolarizing phase there may be an after-hyperpolarizing phase, during which the membrane
potential temporarily becomes more negative than the resting level.
Two types of voltage-gated channels open and then close during an action potential. The first channels that
open, the voltage-gated Na+ channels, allow Na+ to rush into the cell, which causes the depolarizing phase.
Then voltage-gated K+ channels open, allowing K+ to flow out, which produces the repolarizing phase. The
after-hyperpolarizing phase occurs when the voltage-gated K+ channels remain open a' er the repolarizing
phase ends.
An action potential occurs in the membrane of the axon of a neuron when depolarization reaches a certain
level termed the threshold. The generation of an action potential depends on whether a particular stimulus is
able to bring the membrane potential to threshold. An action potential will not occur in response to a
subthreshold stimulus, a weak depolarization that cannot bring the membrane potential to threshold.
However, an action potential will occur in response to a threshold stimulus, a stimulus that is just strong
enough to depolarize the membrane to threshold. Several action potentials will form in response to a
suprathreshold stimulus, a stimulus that is strong enough to depolarize the membrane above threshold.
The period of time a' er an action potential begins during which an excitable cell cannot generate another
action potential in response to a normal threshold stimulus is called the refractory period. During the absolute
refractory period, even a very strong stimulus cannot initiate a second action potential. This period coincides
with the period of Na+ channel activation and inactivation. Inactivated Na+ channels cannot reopen; they
first must return to the resting state. In contrast to action potentials, graded potentials do not exhibit a
refractory period.
In contrast to the graded potential, an action potential is not decremental (it does not die out). Instead, an
action potential keeps its strength as it spreads along the membrane. This mode of conduction is called
propagation, and it depends on positive feedback.
There are two types of propagation: continuous conduction and saltatory conduction. The type of action
potential propagation described so far is continuous conduction, which involves step-by-step depolarization
and repolarization of each adjacent segment of the plasma membrane. Continuous conduction occurs in
unmyelinated axons and in muscle fibers. Saltatory conduction, the special mode of action potential
propagation that occurs along myelinated axons, occurs because of the uneven distribution of voltage-gated
channels. Few voltage-gated channels are present in regions where a myelin sheath covers the axolemma. By
contrast, at the nodes of Ranvier (where there is no myelin sheath), the axolemma has many voltage-gated
channels. Hence, current carried by Na+ and K+ flows across the membrane mainly at the nodes.
The speed of propagation of an action potential is affected by three major factors: amount of myelination,
axon diameter, and temperature.
1. Amount of myelination
action potentials propagate more rapidly along myelinated axons than along unmyelinated axons
2. Axon diameter
larger diameter axons propagate action potentials faster than smaller ones due to their larger surface
areas
3. Temperature
axons propagate action potentials at lower speeds when cooled
Although the plasma membranes of presynaptic and postsynaptic neurons in a chemical synapse are close,
they do not touch. They are separated by the synaptic cleft, a space of 20–50 nm that is filled with interstitial
fluid. Nerve impulses cannot conduct across the synaptic cleft, so an alternative, indirect form of
communication occurs. In response to a nerve impulse, the presynaptic neuron releases a neurotransmitter
that diffuses through the fluid in the synaptic cleft and binds to receptors in the plasma membrane of the
postsynaptic neuron. The postsynaptic neuron receives the chemical signal and in turn produces a
postsynaptic potential, a type of graded potential. Thus, the presynaptic neuron converts an electrical signal
(nerve impulse) into a chemical signal (released neurotransmitter). The postsynaptic neuron receives the
chemical signal and in turn generates an electrical signal (postsynaptic potential). The time required for these
processes at a chemical synapse, a synaptic delay of about 0.5 msec.
A typical chemical synapse transmits a signal as follows:
1) A nerve impulse arrives at a synaptic end bulb of a presynaptic axon
2) The depolarizing phase of the nerve impulse opens voltage-gated Ca2+ channels, which are present
in the membrane of synaptic end bulbs. Because calcium ions are more concentrated in the
extracellular fluid, Ca2+ flows inward through the opened channels.
3) An increase in the concentration of Ca2+ inside the presynaptic neuron serves as a signal that
triggers exocytosis of the synaptic vesicles. The neurotransmitter molecules within the vesicles are
released into the synaptic cleft. Each synaptic vesicle contains several thousand molecules of
neurotransmitter.
4) The neurotransmitter molecules diffuse across the synaptic cleft and bind to neurotransmitter
receptors in the postsynaptic neuron’s plasma membrane.
5) Binding of neurotransmitter molecules to their receptors on ligand-gated channels opens the
channels and allows particular ions to flow across the membrane.
6) As ions flow through the opened channels, the voltage across the membrane changes. This change in
membrane voltage is a postsynaptic potential. Depending on which ions the channels admit, the
postsynaptic potential may be a depolarization (excitation) or a hyperpolarization (inhibition).
7) When a depolarizing postsynaptic potential reaches threshold, it triggers an action potential in the
axon of the postsynaptic neuron.
Neurotransmitters released from a presynaptic neuron bind to neurotransmitter receptors in the plasma
membrane of a postsynaptic cell. Each type of neurotransmitter receptor has one or more neurotransmitter
binding sites where its specific neurotransmitter binds. Neurotransmitter receptors are classified as either
ionotropic receptors or metabotropic receptors based on whether the neurotransmitter binding site and the ion
channel are components of the same protein or are components of different proteins.
The same neurotransmitter can be excitatory at some synapses and inhibitory at other synapses, depending
on the structure of the neurotransmitter receptor to which it binds. For example, at some excitatory synapses
acetylcholine (ACh) binds to ionotropic receptors that contain cation channels that open and subsequently
generate EPSPs in the postsynaptic cell. By contrast, at some inhibitory synapses ACh binds to metabotropic
receptors coupled to G proteins that open K+ channels, resulting in the formation of IPSPs in the
postsynaptic cell.
Removal of the neurotransmitter from the synaptic cleft is essential for normal synaptic function. If a
neurotransmitter could linger in the synaptic cleft, it would influence the postsynaptic neuron, muscle fiber,
or gland cell indefinitely. Neurotransmitter is removed in three ways:
- Diffusion
- Enzymatic degradation
- Uptake by cells
-neurotransmitters-
Neurotransmitters can be divided into two classes based on size: small-molecule neurotransmitters and
neuropeptides.
The small-molecule neurotransmitters include:
Acetylcholine
ACh slows heart rate at inhibitory synapses made by parasympathetic neurons of the vagus (X)
nerve. The enzyme acetylcholinesterase (AChE) inactivates ACh by splitting it into acetate and
choline fragments.
Amino acids
Several amino acids are neurotransmitters in the CNS. Glutamate (glutamic acid) and aspartate
(aspartic acid) have powerful excitatory effects. Most excitatory neurons in the CNS and perhaps
half of the synapses in the brain communicate via glutamate.
Gamma-aminobutyric acid (GABA) and glycine are important inhibitory neurotransmitters. At many
synapses, the binding of GABA to ionotropic receptors opens Cl− channels. GABA is found only in
the CNS, where it is the most common inhibitory neurotransmitter. As many as one-third of all brain
synapses use GABA. Antianxiety drugs such as diazepam (Valium®) enhance the action of GABA.
Like GABA, the binding of glycine to ionotropic receptors opens Cl− channels. About half of the
inhibitory synapses in the spinal cord use the amino acid glycine; the rest use GABA.
Biogenic amines
Certain amino acids are modified and decarboxylated (carboxyl group removed) to produce biogenic
amines. Those that are prevalent in the nervous system include norepinephrine, epinephrine,
dopamine, and serotonin.
Most biogenic amines bind to metabotropic receptors; Biogenic amines may cause either excitation
or inhibition, depending on the type of metabotropic receptor at the synapse.
Norepinephrine (NE) plays roles in arousal (awakening from deep sleep), dreaming, and regulating
mood. A smaller number of neurons in the brain use epinephrine as a neurotransmitter. Both
epinephrine and norepinephrine also serve as hormones. Cells of the adrenal medulla, the inner
portion of the adrenal gland, release them into the blood.
Brain neurons containing the neurotransmitter dopamine (DA) are active during emotional
responses,
addictive behaviors, and pleasurable experiences. In addition, dopamine-releasing neurons help
regulate skeletal muscle tone and some aspects of movement due to contraction of skeletal muscles.
The muscular stiffness that occurs in Parkinson’s disease is due to degeneration of neurons that
release
dopamine.
Norepinephrine, dopamine, and epinephrine are classified chemically as catecholamines. They
all have an amino group (-NH2) and a catechol ring composed of six carbons and two adjacent
hydroxyl (-OH) groups.
Serotonin, which is also known as 5-hydroxytryptamine (5-HT), is concentrated in the neurons in a
part of the brain called the raphe nucleus. It is thought to be involved in sensory perception,
temperature regulation, control of mood, appetite, and the induction of sleep.
ATP and other purines
Nitric oxide
is an important excitatory neurotransmitter secreted in the brain, spinal cord, adrenal glands, and
nerves to the penis and has widespread effects throughout the body. it is formed
on demand and acts immediately. Its action is brief because NO is a
highly reactive free radical. It exists for less than 10 seconds before it
combines with oxygen and water to form inactive nitrates and nitrites.
Because NO is lipid-soluble, it di( uses from cells that produce it into
neighboring cells, where it activates an enzyme for production of a
second messenger called cyclic GMP. Some research suggests that NO
plays a role in memory and learning.
Carbon monoxide
Carbon monoxide (CO), like NO, is not produced in advance and packaged into synaptic vesicles. It
too is formed as needed and diffuses out of cells that produce it into adjacent cells. CO is an
excitatory neurotransmitter produced in the brain and in response to some neuromuscular and
neuroglandular functions. CO might protect against excess neuronal activity and might be related to
dilation of blood vessels, memory, olfaction (sense of smell), vision, thermoregulation, insulin
release, and anti-inflammatory activity.
Neurotransmitters consisting of 3 to 40 amino acids linked by peptide bonds called neuropeptides are
numerous and widespread in both the CNS and PNS. Neuropeptides bind to metabotropic receptors and have
excitatory or inhibitory actions, depending on the type of metabotropic receptor at the synapse.
Neuropeptides are formed in the neuron cell body, packaged into vesicles, and transported to axon terminals.
Besides their role as neurotransmitters, many neuropeptides serve as hormones that regulate physiological
responses elsewhere in the body.
- Substance P
Found in sensory neurons, spinal cord pathways, and parts of brain associated with pain;
enhances perception of pain.
- Enkephalins
Inhibit pain impulses by suppressing release of substance P; may have role in memory and
learning, control of body temperature, sexual activity, and mental illness
- Endorphins
Inhibit pain by blocking release of substance P; may have role in memory and learning, sexual
activity, control of body temperature, and mental illness
- Dynorphins
May be related to controlling pain and registering emotions
- Hypothalamic releasing and inhibiting hormones
Produced by hypothalamus; regulate release of hormones by anterior pituitary
- Angiotensin II
Stimulates thirst; may regulate blood pressure in brain. As a hormone, causes vasoconstriction
and promotes release of aldosterone, which increases rate of salt and water reabsorption by
kidneys
- Cholecystokinin (CCK)
Found in brain and small intestine; may regulate feeding as a “stop eating” signal. As a hormone,
regulates pancreatic enzyme secretion during digestion, and contraction of smooth muscle in
gastrointestinal tract
- Neuropeptide Y
Stimulates food intake; may play a role in the stress response
-the spinal cord-
The spinal cord and spinal nerves contribute to homeostasis by providing
quick, reflexive responses to many stimuli. The spinal cord is the pathway for
sensory input to the brain and motor output from the brain. A spinal cord
reflex—a quick, automatic response to certain kinds of stimuli that involves
neurons only in the spinal nerves and spinal cord.
The spinal cord extends from the foramen magnum to approximately the
level of the disc between vertebrae LI and LII in adults, although it can end
as high as vertebra TXII or as low as the disc between vertebrae LII and LIII.
In neonates, the spinal cord extends approximately to vertebra LIII but can
reach as low as vertebra LIV. The distal end of the cord (the conus
medullaris) is cone shaped. A fine filament of connective tissue (the pial part
of the filum terminale) continues inferiorly from the apex of the conus
medullaris.
The spinal cord is not uniform in diameter along its length. It has two major
swellings or enlargements in regions associated with the origin of spinal nerves that innervate the upper and
lower limbs. A cervical enlargement occurs in the region associated with the origins of spinal nerves C5 to
Tl, which innervate the upper limbs. A lumbosacral enlargement occurs in the region associated with the
origins of spinal nerves L1 to S3, which innervate the lower limbs.
The external surface of the spinal cord is marked by a number of fissures and sulci:
The anterior median fissure extends the length of the anterior surface.
The posterior median sulcus extends along the posterior surface.
The posterolateral sulcus on each side of the posterior surface marks where the posterior rootlets of
spinal nerves enter the cord.
Internally, the cord has a small central canal surrounded by gray and white matter:
- The gray matter is rich in nerve cell bodies, which form longitudinal columns along the cord,
and in cross section these columns form a characteristic H-shaped appearance in the central
regions of the cord.
- The white matter surrounds the gray matter and is rich in nerve cell processes , which form large
bundles or tracts that ascend and descend in the cord to other spinal cord levels or carry
information to and from the brain.
The arterial supply to the spinal cord comes from two sources. It consists of:
1. longitudinally oriented vessels, arising superior to the cervical portion of the cord, which descend on
the surface of the cord; and
2. feeder arteries that enter the vertebral canal through the intervertebral foramina at every level; these
feeder vessels, or segmental spinal arteries, arise predominantly from the vertebral and deep cervical
arteries in the neck, the posterior intercostal arteries in the thorax, and the lumbar arteries in the
abdomen
After entering an intervertebral foramen, the segmental spinal arteries give rise to anterior and posterior
radicular arteries. This occurs at every vertebral level. The radicular arteries follow, and supply, the anterior
and posterior roots. At various vertebral levels, the segmental spinal arteries also give off segmental
medullary arteries. These vessels pass directly to the longitudinally oriented vessels, reinforcing these.
The longitudinal vessels consist of:
• a single anterior spinal artery, which originates within the cranial cavity as the union of two vessels that
arise from the vertebral arteries-the resulting single anterior spinal artery passes inferiorly, approximately
parallel to the anterior median fissure, along the surface of the spinal cord; and
• two posterior spinal arteries, which also originate in the cranial cavity, usually arising directly from a
terminal branch of each vertebral artery (the posterior inferior cerebellar artery)-the right and left posterior
spinal arteries descend along the spinal cord, each as two branches that bracket the posterolateral sulcus and
the connection of posterior roots with the spinal cord.
The anterior and posterior spinal arteries are reinforced along their length by eight to ten segmental
medullary arteries. The largest of these is the arteria radicularis magna or the artery of Adamkiewicz. This
vessel arises in the lower thoracic or upper lumbar region, usually on the left side, and reinforces the arterial
supply to the lower portion of the spinal cord, including the lumbar enlargement.
Veins that drain the spinal cord form a number of longitudinal channels:
• Two pairs of veins on each side bracket the connections of the posterior and anterior roots to the cord.
• One midline channel parallels the
anterior median fissure
• One midline channel passes along
the posterior median sulcus
These longitudinal channels drain into
an extensive internal vertebral plexus
in the extradural (epidural) space of
the vertebral canal, which then drains
into segmentally arranged vessels that
connect with major systemic veins,
such as the azygos system in the
thorax. The internal vertebral plexus
also communicates with intracranial
veins.
The meninges are three protective, connective tissue coverings that encircle the spinal cord and brain. From
superficial to deep they are the (1) dura mater, (2) arachnoid mater, and (3) pia mater. The spinal meninges
surround the spinal cord and are continuous with the cranial meninges, which encircle the brain.
- The spinal dura mater is the outermost meningeal membrane and is separated from the bones
forming the vertebral canal by an extradural space. Superiorly, it is continuous with the inner
meningeal layer of cranial dura mater at the foramen magnum of the skull. Inferiorly, the dural
sac dramatically narrows at the level of the lower border of vertebra SII and forms an investing
sheath for the pial part of the filum terminale of the spinal cord. This terminal cord-like
extension of dura mater (the dural part of the filum terminale) attaches to the posterior surface of
the vertebral bodies of the coccyx.
The dura mater is also continuous with the epineurium, the outer covering of spinal and cranial
nerves.
- The arachnoid mater is a thin delicate membrane against, but not adherent to, the deep surface of
the dura mater (between the dura mater and the arachnoid mater is a thin) subdural space, which
contains interstitial fluid. It is separated from the pia mater by the subarachnoid space. The
arachnoid mater ends at the level of vertebra SII
- The pia mater is the innermost meninx and it’s a thin transparent connective tissue layer that
adheres to the surface of the spinal cord and brain. Within the pia mater are many blood vessels
that supply oxygen and nutrients to the spinal cord. Triangular-shaped membranous extensions
of the pia mater suspend the spinal cord in the middle of its dural sheath. These extensions,
called denticulate ligaments, are thickenings of the pia mater. They project laterally and fuse
with the arachnoid mater and inner surface of the dura mater between the anterior and posterior
nerve roots of spinal nerves on either side. Extending along the entire length of the spinal cord,
the denticulate ligaments protect the spinal cord against sudden displacement that could result in
shock. Between the arachnoid mater and pia mater is a space, the subarachnoid space, which
contains shock-absorbing cerebrospinal fluid.
Spinal tap: In a spinal tap (lumbar puncture), a local anesthetic is given, and a long hollow needle is
inserted into the subarachnoid space to withdraw cerebrospinal fluid (CSF) for diagnostic purposes; to
introduce antibiotics, contrast media for myelography, or anesthetics; to administer chemotherapy; to
measure CSF pressure; and/or to evaluate the e& ects of treatment for diseases such as meningitis. The
spinal cord ends around the second lumbar vertebra (L2); however, the spinal meninges and circulating
cerebrospinal fluid extend to the second sacral vertebra (S2). Between vertebrae L2 and S2 the spinal
meninges are present, but the spinal cord is absent. Consequently, a spinal tap is normally performed in
adults between the L3 and L4 or L4 and L5 lumbar vertebrae because this region provides safe access to the
subarachnoid space without the risk of damaging the spinal cord. (A line drawn across the highest points of
the iliac crests, called the supracristal line, passes through the spinous process of the fourth lumbar vertebra
and is used as a landmark for administering a spinal tap.)
There are approximately 31 pairs of spinal nerves, named according to their position with respect to
associated vertebrae:
• eight cervical nerves - C1 to C8,
• twelve thoracic nerves - T1 to T 12,
• five lumbar nerves - L1 to L5,
• five sacral nerves - S1 to S5,
• one coccygeal nerve - Co.
The first cervical nerve (C1) emerges from the vertebral canal between the skull and vertebra CI. Therefore
cervical nerves C2 to C7 also emerge from the vertebral canal above their respective vertebrae. Because
there are only seven cervical vertebrae, C8 emerges between vertebrae CVII and TI. As a consequence, all
remaining spinal nerves, beginning with T1 , emerge from the vertebral canal below their respective
vertebrae.
A transverse section of the spinal cord reveals regions of white matter that surround an inner core of gray
matter. In the gray matter of the spinal cord and brain, clusters of neuronal cell bodies form functional groups
called nuclei. Sensory nuclei receive input from receptors via sensory neurons, and motor nuclei provide
output to effector tissues via motor neurons. The gray matter on each side of the spinal cord is subdivided
into regions called horns. The posterior (dorsal) gray horns contain axons of incoming sensory neurons as
well as cell bodies and axons of interneurons. The anterior (ventral) gray horns contain somatic motor nuclei,
which are clusters of cell bodies of somatic motor neurons that provide nerve impulses for contraction of
skeletal muscles. Between the posterior and anterior gray horns are the lateral gray horns, which are present
only in thoracic and upper lumbar segments of the spinal cord. The lateral gray horns contain autonomic
motor nuclei, which are clusters of cell bodies of autonomic motor neurons that regulate the activity of
cardiac muscle, smooth muscle, and glands. The white matter of the spinal cord, like the gray matter, is
organized into regions. The anterior and posterior gray horns divide the white matter on each side into three
broad areas called columns: (1) anterior (ventral) white columns, (2) posterior (dorsal) white columns, and
(3) lateral white columns. Each column in turn contains distinct bundles of axons having a common origin or
destination and carrying similar information.
The internal organization of the spinal cord allows sensory input and motor output to be processed by the
spinal cord in the following way:
1. Sensory receptors detect a sensory stimulus
2. Sensory neurons convey this sensory input in the form of nerve impulses along their axons, which
extend from sensory receptors into the spinal nerve and then into the posterior root. From the
posterior root, axons of sensory neurons may proceed along three possible paths:
- Axons of sensory neurons may extend into the white matter of the spinal cord and ascend to the
brain as part of a sensory tract
- Axons of sensory neurons may enter the posterior gray horn and synapse with interneurons
whose axons extend into the white matter of the spinal cord and then ascend to the brain as part
of a sensory tract.
- Axons of sensory neurons may enter the posterior gray horn and synapse with interneurons that
in turn synapse with somatic motor neurons that are involved in spinal reflex pathways
3. Motor output from the spinal cord to skeletal muscles involves somatic motor neurons of the anterior
gray horn. Many somatic motor neurons are regulated by the brain. Axons from higher brain centers
form motor tracts that descend from the brain into the white matter of the spinal cord. There they
synapse with somatic motor neurons either directly or indirectly by first synapsing with interneurons
that in turn synapse with somatic motor neurons.
4. When activated, somatic motor neurons convey motor output in the form of nerve impulses along
their axons, which sequentially pass through the anterior gray horn and anterior root to enter the
spinal nerve. From the spinal nerve, axons of somatic motor neurons extend to skeletal muscles of
the body
5. Motor output from the spinal cord to cardiac muscle, smooth muscle, and glands involves autonomic
motor neurons of the lateral gray horn. When activated, autonomic motor neurons convey motor
output in the form of nerve impulses along their axons, which sequentially pass through the lateral
gray horn, anterior gray horn, and anterior root to enter the spinal nerve
6. From the spinal nerve, axons of autonomic motor neurons from the spinal cord synapse with another
group of autonomic motor neurons located in the peripheral nervous system (PNS). The axons of this
second group of autonomic motor neurons in turn synapse with cardiac muscle, smooth muscle, and
glands
Plexuses
Axons from the anterior rami of spinal nerves, except for thoracic nerves T2–T12, do not go directly to the
body structures they supply. Instead, they form networks on both the left and right sides of the body by
joining with various numbers of axons from anterior rami of adjacent nerves. Such a network of axons is
called a plexus. The principal plexuses are the cervical plexus, brachial plexus, lumbar plexus, and sacral
plexus. A smaller coccygeal plexus is also present. Emerging from the plexuses are nerves bearing names
that are often descriptive of the general regions they serve or the course they take. Each of the nerves in turn
may have several branches named for the specific structures they innervate.
The anterior rami of spinal nerves T2–T12 do not enter into the formation of plexuses and are known as
intercostal nerves or thoracic nerves. These nerves directly connect to the structures they supply in the
intercostal spaces. After leaving its intervertebral foramen, the anterior ramus of nerve T2 innervates the
intercostal muscles of the second intercostal space and supplies the skin of the axilla and posteromedial
aspect of the arm. Nerves T3–T6 extend along the costal grooves of the ribs and then to the intercostal
muscles and skin of the anterior and lateral chest wall. Nerves T7–T12 supply the intercostal muscles and
abdominal muscles, along with the overlying skin. The posterior rami of the intercostal nerves supply the
deep back muscles and skin of the posterior aspect of the thorax.
Dermatomes
A dermatome is an area of skin that provides sensory input to the CNS via the posterior roots of one pair of
spinal nerves or via the trigeminal (V) nerve, which serves most of the skin of the face and sculp.
Knowing which spinal cord segments supply each dermatome makes it possible to locate damaged regions of
the spinal cord. If the skin in a particular region is stimulated but the sensation is not perceived, the nerves
supplying that dermatome are probably damaged. In regions where the overlap is considerable, little loss of
sensation may result if only one of the nerves supplying the dermatome is damaged.
Cervical plexus
The cervical plexus is formed by the roots (anterior rami) of the first four cervical nerves (C1–C4), with
contributions from C5. There is one on each side of the neck alongside the first four cervical vertebrae.
The cervical plexus supplies the skin and muscles of the head, neck, and superior part of the shoulders and
chest. The phrenic nerves arise from the cervical plexuses and supply motor fibers to the diaphragm.
Branches of the cervical plexus also run parallel to two cranial nerves, the accessory (XI) nerve and
hypoglossal (XII) nerve.
Brachial plexus
The roots (anterior rami) of spinal nerves C5–C8 and T1 form the
brachial plexus, which extends inferiorly and laterally on either side of
the last four cervical and first thoracic vertebrae. It passes above the first
rib posterior to the clavicle and then enters the axilla.
As with the cervical and other plexuses, the roots are the anterior rami of
the spinal nerves. The roots of several spinal nerves unite to form trunks
in the inferior part of the neck. These are the superior, middle, and
inferior trunks. Posterior to the clavicles, the trunks diverge into
divisions, called the anterior and posterior divisions. In the axillae, the
divisions unite to form cords called the lateral, medial, and posterior
cords. The cords are named for their relationship to the axillary artery, a
large artery that supplies blood to the upper limb. The branches of the
brachial plexus form the principal nerves of the brachial plexus. The
brachial plexus provides almost the entire nerve supply of the shoulders
and upper limbs.
Five large terminal branches arise from the brachial plexus: (1) The
axillary nerve supplies the deltoid and teres minor muscles. (2) The
musculocutaneous nerve supplies the anterior muscles of the arm. (3) The
radial nerve supplies the muscles on the posterior aspect of the arm and
forearm. (4) The median nerve supplies most of the muscles of the
anterior forearm and some of the muscles of the hand. (5) The ulnar nerve
supplies the anteromedial muscles of the forearm and most of the muscles
of the hand.