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Anatomy of The Sympathetic Nervous System

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Anatomy of The Sympathetic Nervous System

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Chapter 36

Anatomy of the Sympathetic


Nervous System
Toral R. Patel
University of Alabama School of Medicine, Birmingham, Alabama, USA.
e-mail: [email protected]

AUTONOMIC NERVOUS SYSTEM nervous system (CNS). Its axon synapses on the cell body of a
postsynaptic neuron located outside the CNS in autonomic
The autonomic nervous system (ANS), also known as the ganglia. The fibers from the postsynaptic neuron terminate
visceral motor system, is a complex network that is part on effector organs. This construction allows for a collection
of the peripheral nervous system that maintains internal of neuronal bodies, also known as ganglia, outside of the
physiologic homeostasis. The ANS consists of motor fibers CNS. The postganglionic fibers of the parasympathetic
that stimulate smooth involuntary muscle, modified cardiac system are generally very short compared to the longer post-
muscles, and glandular secretory cells. The afferent com- ganglionic fibers of the sympathetic nervous system.
ponent of the autonomic reflexes conducts visceral pain Peripheral nerve fibers are also classified by function; efferent
impulses and regulates visceral functions. Receptors of fibers are motor fibers that innervate skeletal muscle and
the afferent component reside in the viscera and are sen- afferent fibers are sensory fibers. Other fibers consist of motor
sitive to mechanical, chemical, or thermal stimuli. The and sensory, such as those innervating viscera and glands.
afferent nerves (sensory) conduct signals along somatic Preganglionic axons in both systems utilize the neuro-
and autonomic nerves, which enter the spinal cord through transmitter acetylcholine, which acts on nicotinic receptors
the dorsal roots or the brain stem through cranial nerves. on the postganglionic neuron. Postganglionic axons in the
The efferent component (motor) is divided into two parasympathetic system also utilize acetylcholine while
divisions: the parasympathetic nervous system (PNS) and acting on muscarinic receptors at the target organ. Postgan-
the sympathetic nervous system (SNS). The PNS is also glionic axons in the sympathetic system use norepinephrine,
considered the craniosacral division, while the SNS is which acts on adrenergic receptors. An exception to this is
considered the thoracolumbar division. Both subsystems the postganglionic sympathetic axons to the sweat glands,
operate independently in some functions and interact coop- which have muscarinic receptors and require acetylcholine
eratively in others. An older hypothesis of the sympathetic as a neurotransmitter (Hare & Hinsey, 1942).
and parasympathetic nervous system functions charac-
terized them as “excitatory” and “inhibitory” of physio-
logical responses. A modern reconsideration characterizes
Anatomy and Physiology
the SNS as a “quick response exciting a system” and the
PNS as “slower activated but inhibiting a system.” Many As mentioned earlier, the ANS innervation is divided into the
also label the PNS as the “rest and digest” system and the sympathetic nervous system and the parasympathetic
SNS as the “fight or flight” system. The third subsystem nervous system. The sympathetic division has thoracolumbar
of the ANS is the enteric system. The enteric division lines outflow, meaning that the neurons begin at the thoracic and
the esophagus, stomach, intestines, gallbladder, and pan- lumbar (T1-L2/3) portions of the spinal cord. The parasym-
creas and operates almost entirely independently; however, pathetic division has craniosacral outflow, meaning that the
its functions can be overridden by the sympathetic and para- neurons begin with cranial nerves III, VII, IX, and X, as well
sympathetic divisions. as the sacral spinal roots.
The autonomic nervous system differs from the somatic Peripheral nerves may be divided into myelinated
system in that two neurons are involved in sending signals and unmyelinated nerves. Classically, the large fibers are
from the source to the target. The nerve cell body of the first myelinated and the small ones are unmyelinated. The
presynaptic neuron is located in the grey matter of the central central core of the nerve fiber is the axon, and the membrane

Nerves and Nerve Injuries, Vol. 1. http://dx.doi.org/10.1016/B978-0-12-410390-0.00038-X


© 2015 Elsevier Ltd. All rights reserved. 495
496 PART IV Anatomy of the Spinal Nerves and Cervical Plexus

FIGURE 36.1 Arrangement of nerve fibers.

of the axon conducts the action potential. The axon is filled Axonal transport is further divided into fast and slow
with axoplasm, a viscous intracellular fluid that contains the transport. Vesicular cargo moves relatively fast (50-
microtubules, neurofilaments, mitochondria, endoplasmic 400 mm/day) whereas soluble (cytosolic) and cytoskeletal
reticulum, and granular or vesicular structures. The axo- proteins move much slower (moving at less than 8 mm/
plasm is surrounded by a multilayered sheath known as day) during transport. Anterograde axonal transport has a
the myelin sheath (Figure 36.1). The myelin sheath is major fast and a slow component. The slow component is
formed by Schwann cells of the peripheral nervous system, divided into “slow component a” and “slow component b”
which form chains of neurolemmal cells along the nerves. at rates of approximately 0.1 and 6 mm/day, respectively.
Unmyelinated fibers conduct continuous or uninterrupted The fast component of transport demonstrates maximal rates
impulses. On myelinated nerve fibers, every 1-3 mm is a for anterograde (200-400 mm/day) and retrograde (100-
node of the Ranvier, an uninsulated area of 2-3 μm in length 200 mm/day) transport (Oztas, 2003). The mechanisms of
filled with sodium channels, responsible for an increase in slow axonal transport are only recently becoming clear, as
speed in conduction of action potentials through a neuron. a result of advanced imaging techniques such as fluorescent
The impulse conduction is saltatory or jumping as the labeling techniques in fluorescence microscopy. Recent
impulse jumps from node to node. The speed of conduction studies have revealed that the movement of cytoskeletal
is much greater along myelinated fibers. Conduction veloc- “slow” cargo is actually rapid but, unlike fast transport, they
ities depend on axon and total fiber diameter, myelin pause frequently, making the overall transit rate much
thickness, myelin structure, and intermodal length. slower. The mechanism is known as the “stop and go” model
Hundreds to thousands of fibers are present in each nerve. of slow axonal transport (Brown, 2003; Kuznetsov, 2011).
A nerve as a whole is surrounded by the epineurium, a con- The rapid transport system is dependent on temperature
nective tissue sheath. Connective tissue fibers run inward (axonal transport is reduced at low temperatures (9-11  C))
from the sheath and enclose bundles of nerve fibers. Such with nerve blocks at temperatures lower than this. This is
bundles are termed fasciculi (funiculi); the connective tissue attributed to the disruption of microtubules unstable at low
that encloses them is called perineurium. Each nerve fiber is temperatures; age (decrease in trafficking as the systems
enclosed by a connective tissue sheath termed endoneurium. innervated mature); Ca ions; oxidative metabolism (an ade-
(Figure 36.2) In order to develop and function during growth quate supply of blood-transported oxygen and ATP is
and survival, the neurons require fast and slow transport required for fast transport); and the integrity of microtubules
systems through retrograde and anterograde transport (agents such as colchicines and vinblastine disrupt neurotu-
(Schwartz, 1979). The retrograde system is comprised of bules and block both fast and slow transport).
materials traveling centrally to the cell body, whereas anter-
ograde transport includes material traveling to the synapse.
Types of Nerve Fibers
Microtubules made of tubulin in the axon provide cyto-
skeletal tracks for transportation. The motor protein kinesin Motor, sensory, and sympathetic nerve fibers form the
is involved in anterograde transport, while the motor protein peripheral nerve trunks or faniculi (Shields, 1993). The
dynein is involved in retrograde transport. These motor pro- motor nerve fibers originate in the anterior horn neurons
teins bind and transport organelles, vesicles, and other mate- of the spinal cord and terminate at the neuromuscular junc-
rials. Some pathogens exploit this process to invade the tions of skeletal muscle. Motor fibers can be large or small
nervous system by entering the distal axon and traveling to with the difference that the large fibers innervate the extra-
the soma by retrograde transport. Examples of such path- fusal fibers of the muscle and the small fibers innervate the
ogens include tetanus toxin, herpes simplex, rabies, and polio intrafusal fibers of the muscle. The sensory nerve fibers
viruses (Saladin, 2010). carry peripheral dendrites of the posterior root ganglion
Anatomy of the Sympathetic Nervous System Chapter 36 497

FIGURE 36.2 Connective tissue coverings of a nerve fiber.

neurons. Sensory fibers can be myelinated or nonmyeli- The cell bodies of the presynaptic SNS neurons are found
nated, and terminate at the periphery, specialized end in the intermediolateral cell columns. The columns are part
organs, or receptors. Cutaneous sensory fibers travel in of the gray matter of the thoracic (T1-T12) and upper
cutaneous nerves; deep sensory fibers travel in the muscular lumbar (L1-L2 or 3) segments of the spinal cord in both
and deep branches or the nerve trunks and terminate in the right and left lateral horns. The IMLs are organized
deeper structures such as muscles, tendons, articular and somatotrophically. Preganglionic axons exiting the spinal
periarticular structures, bone, and connective tissue. Sym- cord enter the white rami communicans to join a network
pathetic nerve fibers are the postganglionic processes of of prevertebral and paravertebral ganglia in the sympathetic
the neurons from the sympathetic trunk. Generally, these trunk or chain. These preganglionic/presynaptic axons are
fibers are nonmyelinated and are destined for blood vessels, relatively short, myelinated, and cholinergic.
glandular structures of the skin, and deeper structures The postsynaptic/postganglionic neurons are located in
(Figure 36.3). the paravertebral and prevertebral ganglia. The paraver-
tebral ganglia form the right and left sympathetic chains
on each side of the vertebral column and extend the length
of the column. The prevertebral ganglia are located in the
SYMPATHETIC NERVOUS SYSTEM plexuses that surround the origins of the main branches
of the abdominal aorta. The postsynaptic neurons are motor
Anatomy
fibers that leave the spinal cord through the anterior roots and
The sympathetic nervous system (SNS) is governed by enter the anterior rami of the spinal nerves T1-L2 or 3.
upper motor neurons (i.e., neurons superior to the The neurons then pass through a white ramus before
spinal cord) derived from the hypothalamus. These fibers entering a ganglia of the sympathetic chain. The postgan-
descend into the brain stem and spinal cord as the hypotha- glionic axons exit the ganglia through the gray rami com-
lamospinal tracts. The hypothalamospinal tracts travel in municans and travel to their target organs primarily along
the white matter posterolateral to the dorsal horn of the blood vessels within the body cavities or into the head.
spinal cord and descend to end on neuronal cell bodies These postganglionic axons are long, unmyelinated, and
within the intermediolateral (IML) cell column in the thor- primarily adrenergic; the exception being the innervation
acolumbar regions of the spine, extending from TI to L2. of the sweat glands, which are cholinergic.
498 PART IV Anatomy of the Spinal Nerves and Cervical Plexus

FIGURE 36.3 Composition and distribution of a sympathetic nerve.

The presynaptic fibers travel via four routes. The first the trunk involved in innervating abdominopelvic viscera)
being an ascent in the sympathetic trunk to synapse with to reach the prevertebral ganglia.
a postsynaptic neuron of a higher paravertebral ganglion. The postsynaptic fibers are destined for distribution
The second being a descent in the sympathetic trunk to within the neck, body wall, and limbs. Some postganglionic
synapse with a postsynaptic neuron of a lower paravertebral fibers may not distribute themselves to the viscera but
region. The third accounts for the presynaptic neurons instead reenter a spinal nerve segment via a gray ramus
entering and immediately synapsing with a postsynaptic communicans to be disseminated to the skin with its
neuron of the paravertebral ganglion at the respected level. smooth muscle (arrector pili), blood vessels, and glands
The fourth course consists of the axon passing through the and the vessels of the extremities and walls of the body
sympathetic trunk without synapsing and continuing cavities. Topographically, the white rami communicans
through an abdominopelvic splanchnic nerve (a branch of are found slightly lateral to the gray ramus communicans
Anatomy of the Sympathetic Nervous System Chapter 36 499

and morphologically these former rami are generally corresponding nerve segment, entering one of the branches
larger in size due to the preponderance of myelinated fibers. of spinal nerves (Figure 36.4).
Additionally, white ramus communicans will carry
visceral afferent fibers from the body cavities. Moreover,
in the upper lumbar region the white rami usually ascend
Ganglia
obliquely from a given ganglion, because they typically The sympathetic nervous system is comprised of sympa-
originate from a nerve from the next more superior thetic chain ganglia, paravertebral ganglia, and prevertebral
level. Gray rami tend to run horizontally or travel ganglia. The sympathetic chain is found at the base of the
inferiorly only slightly from a ganglion to their skull and extends to the lowest part of the vertebral column

FIGURE 36.4 Sympathetic divisions of ANS.


500 PART IV Anatomy of the Spinal Nerves and Cervical Plexus

at the coccyx. The sympathetic chain travels posteriorly to calcium ions exciting the cells to release their contents—
the carotid sheath in the neck. In the thorax, the ganglia lies either acetylcholine or norepinephrine. In the preganglionic
anterior to the costovertebral junction. In the abdomen, the neurons, acetylcholine is synthesized from acetyl-CoA and
ganglia passes posterior to the medial arcuate ligament and choline by the enzyme choline acetylesterase. It is then
runs anteriorly on the anterolateral aspect of the vertebral stored in vesicles. Once secreted into a tissue, the acetyl-
bodies. They are then found posterior to the common iliac choline performs its function before splitting into an acetate
vessels and commonly have four ganglia bilaterally, as does ion and choline. This reaction is catalyzed by the enzyme
the lumbar segment of this chain. These ganglia may vary in acetylcholinesterase. The choline formed is then trans-
size from 6 mm to 6 cm. The most constant and usually ported back to the terminal nerve, ending where it is used
largest of these ganglia is the ganglion located on the second again to synthesize acetylcholine.
lumbar vertebra. The lumbar segments of the sympathetic In the sympathetic postganglioninc nerve endings, nor-
chains pass between the psoas major and inferior vena cava epinephrine synthesis begins in the axoplasm and is termi-
on the right side and between the psoas major and aorta on nated in the secretory vesicles of the terminal nerve fibers.
the left side. The sacral part of the sympathetic chain pro- Tyrosine is hydroxylated into the substance dopa, which is
vides two to three branches (sacral splanchnics) to the then decarboxylated into dopamine. Dopamine is then
inferior hypogastric plexus, to be discussed later. Other transported into the secretory vesicles, where it is hydrox-
branches of the lumbosacral sympathetic chain include vas- ylated into norepinephrine. In the adrenal medulla, norepi-
cular, osseous, and articular twigs. The cervical, thoracic, nephrine is further methylated into epinephrine. After
lumbar, and sacral portions of the sympathetic chain consist secretion of the norepinephrine by the nerve endings, there
of 3, 12, 4, and 4-5 ganglia, respectively. are three mechanisms in which it is removed. The first
The paravertebral ganglia of the sympathetic chain run mechanism consists of the reuptake of norepinephrine into
on either side of the vertebral bodies and include the cer- the adrenergic nerve endings by active transport. The
vical ganglia, thoracic ganglia, lumbar ganglia, and pelvic second mechanism consists of diffusion of the norepi-
ganglia. The most rostral ganglion is the superior cervical nephrine into the surrounding bodily fluids and blood.
ganglion, which arises from fusion of C1 to C4. The next The third mechanism consists of the destruction of small
two ganglia are the middle cervical ganglion from C5 amounts of the norepinephrine by tissue enzymes such as
and C6, and the inferior cervical ganglion from C7 and monoamine oxidase. The norepinephrine secreted from
C8. The prevertebral ganglia includes celiac ganglia, aorti- the terminal nerve endings usually lasts a few seconds,
corenal ganglia, superior mesenteric ganglia, and the but the norepinephrine and epinephrine secreted by the
inferior mesenteric ganglia. These ganglia are the plexuses adrenal medulla is active until diffused into a tissue and
that surround the origins of the main branches of the metabolized by the liver.
abdominal aorta. The chromaffin cells of the adrenal The acetylcholine, norepinephrine, and epinephrine
medulla are an exception for the sympathetic ganglia, as secreted by the nerve endings must bind to a receptor in
the presynaptic neurons do not synapse in a ganglia before order to stimulate an effector organ. Sodium or calcium
traveling to the adrenal medulla. The paravertebral and pre- channels are opened to depolarize and excite a cell, while
vertebral ganglia provide postganglionic neurons for inner- others cause potassium channels to open and inhibit the cell.
vations of the target organs. Other receptor actions include altering the expression of
enzymes or intracellular chemicals such as adenylyl cyclase
and cyclic adenosine monophosphate. Two principal types
Physiology
of acetylcholine receptors are the muscarinic and nicotinic
The sympathetic and parasympathetic fibers secrete acetyl- receptors. The muscarinic receptors are found on all
choline or norepinephrine. Those that secrete acetylcholine effector cells stimulated by the postganglionic neurons.
are considered cholinergic and those that secrete norepi- The nicotinic receptors are found in the autonomic ganglia
nephrine are termed adrenergic. All preganglionic neurons at the synapses between the preganglionic and postgan-
are cholinergic. The acetylcholine or acetylcholine-like glionic neurons. There are two types of adrenergic
substances excite the sympathetic or parasympathetic post- receptors, alpha and beta receptors. Both receptors are
ganglionic neurons. The parasympathetic postganglionic further divided into subsets such as beta1 and beta2
neurons are cholinergic. The sympathetic postganglionic receptors. Norepinephrine excites mainly alpha receptors
neurons are adrenergic with the exception of the sympa- but also affects beta receptors to a lesser extent. Epi-
thetic nerve fibers to the sweat glands, piloerector muscles nephrine excites both types of receptors equally. Alpha
of the hair, and a few blood vessels. receptors are known to function for vasoconstriction, iris
In the nerve endings, when an action potential is stimu- dilation, intestinal relaxation, intestinal sphincter con-
lated and spreads to the terminal fibers, depolarization traction, pilomotor contraction, and bladder sphincter
increases the permeability of the fiber membranes to contraction. Beta receptors, on the other hand, affect
Anatomy of the Sympathetic Nervous System Chapter 36 501

TABLE 36.1 Sympathetic Nervous System Receptor Subtypes and Functions

Target SNS Receptor Function


Cardiac output β1, β2 Increases
SA node: heart rate β1, β2 Increases
Atria: contractility β1, β2 Increases
AV node: conduction and β1 Increases
automaticity
Ventricle: contractility, β1, β2 Increases
automaticity
Vascular smooth muscle α1, β2 Contraction with α and relaxation with β
Veins α1, α2, β2 Constriction with α and dilation with β
Bronchioles-smooth muscles β2 (major), α1 Relaxation with β

Pupil dilator muscle α1 Dilation


Ciliary muscle β2 Relaxation
Salivary glands β, α1 β, stimulates viscous secretions; α, stimulates postassium secretions
Lacrimal glands β Stimulates protein secretions
Liver α1 , β2 Glycogenolysis, gluconeogenesis
Adipose cells β1 Lipolysis

Gastrointestinal sphincters α1 , α2 , β2 Contraction


Gastrointestinal motility α1 , α2 , β2 Decreases
Pancreas α2 Decreases insulin secretion from beta cells, increases glucagon secretion
from alpha cells
Adrenal medulla Nicotinic Secretes epinephrine and norepinephrine
Detrusor muscle of bladder β2 Relaxation
Internal urethral sphincter α1 Contraction
Uterus α1 , β2 Contraction with α and relaxation with β
Genitalia α1 Contraction

Sweat glands Muscuranic Stimulates


(major), α1

Errector pili α1 Stimulates contraction

vasodilation, cardioacceleration, increased myocardial dilates pupils, dilates bronchioles, releases adrenaline and
strength, bronchodilation, intestinal relaxation, uterus stored energy in the form of glycogen, increases sweating,
relaxation, glycogenolysis, lipolysis, and bladder wall decreases blood flow to the skin, decreases peristalsis, and
relaxation (Table 36.1). causes dryness in the mouth.

Metabolic
Function The sympathetic nervous system also affects metabolism.
The SNS is known for its fight-or-flight response. The SNS When stimulated, the SNS releases glucose from the liver,
arouses the body in response to stress and increased increases blood glucose concentration, and increases glyco-
demands. Generally, the sympathetic division increases genolysis in the liver and muscles. Sympathetic stimulation
heart rate and blood pressure, increases respiration and also increases muscle strength, basal metabolic rate, and
blood flow to the muscles, dilates blood vessels to the heart, mental activity.
502 PART IV Anatomy of the Spinal Nerves and Cervical Plexus

Cardiac and Vascular Pupils


The sympathetic division is known to increase the heart The central regulation of pupillary action lies in the dorsal
rate and blood pressure, with differences in blood flow midbrain and Edinger-Westphal nucleus. The afferent
depending on the body part. Any increase in blood pathway is located along the optic nerve. The efferent
pressure and cardiac output increases the activity of the pathways consist of preganglionic innervations of the pupil
afferent pathway, reflexively inhibiting the sympathetic from the C8-T2 spinal cord, which run through the superior
system and activating the parasympathetic system. Any cervical ganglion. Postganglionic fibers destined for
decrease in the blood pressure or cardiac output decreases pupillary action extend along the carotid artery to the cav-
the afferent activity, reflexively increasing the sympa- ernous sinus and then enter the orbit via the fifth cranial
thetic system and increasing the blood pressure. The nerve. The result is pupillary dilation.
afferent pathway includes the arterial baroreceptors
located in the carotid sinus, aortic arch, and thoracic
arteries. The baroreceptors respond to changes in the blood Glands
pressure and conduct signals through the glossophar-
Sympathetic stimulation of the alimentary glands causes a
yngeal and vagus nerves. The cardiac mechanoreceptors
secretion of concentrated enzymes and mucus. It also
are sensitive to mechanical deformations in the cardiac
affects the blood vessels that supply the glands through
chambers, thus increasing afferent activity and conducting
vasoconstriction. When the sympathetic nerves terminating
signals through the vagus nerve. The pulmonary stretch
on the sweat glands are stimulated, large quantities of sweat
receptors are sensitive to lung volumes, thus affecting
are secreted. The salivary glands, nasal and lacrimal glands,
the afferent activity and conducting signals through the
as well as the majority of the gastrointestinal glands, are
vagus nerve. The efferent pathway consists of the postgan-
stimulated by the parasympathetic nervous system and
glionic fibers innervating the atria, ventricles, and cor-
inhibited or relaxed to an extent by the sympathetic nervous
onary arteries. Stimulation causes an increase in heart
system.
rate and increases myocardial contractility and coronary
vasodilation. Postganglionic sympathetic fibers inner-
vate the vasculature from plexuses on proximal vessels
or from the somatic nerve. Most systemic blood vessels, Genitourinary
especially the ones located in the abdominal viscera and The genitourinary system’s afferent activity is along the
skin of the limbs, are constricted by the sympathetic fibers. autonomic and somatic pathways. The sympathetic inner-
Alpha adrenergic innervation is responsible for vasocon- vation stems from the T11-L2 or 3 spinal segments, the
striction, while beta adrenergic innervation is responsible inferior mesenteric and superior hypogastric ganglia, and
for vasodilation. the hypogastric nerves. The nerves are responsible for
uterine contraction in females, ejaculation in males, bladder
wall inhibition, destrusor and trigone muscle contraction,
Thermoregulation and urethral muscle contraction. The SNS does not control
the external sphincter, which is innervated by the pudendal
Central regulation of thermoregulation is located in the pre-
nerve and is under somatic, not autonomic, control.
optic and anterior hypothalamus where a set point is estab-
lished by the thermosensitive neurons. A body temperature
below the set point causes autonomic reflexes to generate
heat by shivering and reduce heat loss via cutaneous vaso- Gastrointestinal
constriction and piloerection. When body temperature The spinal centers, nucleus of the tractus solaris, and
exceeds the set point, a stimulation of sweating and cuta- nucleus ambiguus control gastrointestinal activity. The
neous vasodilation occurs to increase the flow of blood to afferent pathway synapses locally or in the ganglia and
the surface of the body from which heat is lost. The afferent spinal cord. The efferent pathways are located in the enteric
activity originates from thermosensitive neurons that are nervous system, which consists of nerves and plexuses
not only located in the hypothalamus but also in the skin, embedded in the walls of the gastrointestinal tract and inte-
abdominal viscera, spinal cord, and brain stem. The efferent grated into local circuits for secretion, absorption, peri-
pathway is controlled by the SNS and PNS. The SNS also stalsis, and sphincter coordination. The sympathetic
innervates the sweat glands to regulate evaporative heat innervation from the thoracolumbar segments stimu-
loss. The SNS fibers cause vasoconstriction of the cuta- lates the esophageal sphincters, decreases peristalsis, and
neous vasculature to reduce heat loss. The accompanying relaxes the internal rectal sphincter. Again, the external
piloerection reduces the surface area and reduces con- sphincters are innervated by the pudendal nerve and under
vective heat loss. somatic control.
Anatomy of the Sympathetic Nervous System Chapter 36 503

Adrenal Medulla Sympathetic Nervous System Thorax


The adrenal medulla synthesizes norepinephrine, The sympathetic division in the thoracic cavity contains
dopamine, and epinephrine (adrenaline). Chromaffin cells cranial nerves (the vagus nerve), cervical ganglia, and tho-
are responsible for the synthesis and excretion of these racic ganglia. The vagus nerve sends fibers to the lungs,
hormones into the bloodstream. The cells respond to heart, and esophagus. Cardiopulmonary splanchnic nerves
stress, emotions, and fight-or-flight stimuli. Notable are also involved in the innervations of these three organs.
effects of these hormones include increased heart rate The atria, ventricles, and coronary arteries of the heart are
and blood pressure, vascular constriction in the skin and innervated by fibers from the cervical ganglia, superior,
gastrointestinal tract, bronchiole and capillary dilation, middle, and inferior cardiac nerves, as well as the thoracic
and an increase in metabolism. ganglia at the levels of T1-T4. The thoracolumbar sympa-
thetic fibers from the dorsolateral region of the anterior
column of gray matter pass with the anterior roots of the
SYMPATHETIC NERVOUS SYSTEM thoracic column and the upper 2-3 lumbar spinal nerves.
DIVISIONS These fibers terminate in sympathetic trunk ganglia and
plexuses before coursing to their destined organs. The car-
Sympathetic Nervous System Head diopulmonary splanchnic nerves from the paravertebral
The two major sources of sympathetic nerves in the head ganglia of C4-T4 travel to the heart and lungs.
are the sympathetic chain ganglia and the cranial nerves.
Those that perform functions in the head and the dilator Sympathetic Nervous System Abdomen
muscle of the iris have their cell bodies in the superior cer-
vical ganglia. These fibers to the head pass from the gan- The SNS division of abdominal innervations consists of the
glion by means of a cephalic arterial branch to form abdominopelvic splanchnic nerves from the thoracic and
periarterial plexuses of nerves, which follow the branches abdominal sympathetic trunks, prevertebral sympathetic
of carotid arteries or they pass to cranial nerves. The ganglia, and the abdominal aortic plexus and its extensions
cranial sympathetic fibers include fibers in the oculo- including the periarterial plexuses. The nerve plexuses are
motor, facial, glossopharyngeal, and vagus nerves. The mixed with PNS fibers and visceral efferent fibers. The pre-
sympathetic efferent fibers of the oculomotor nerve ter- synaptic sympathetic fibers arise from cell bodies in IML
minate with sympathetic motor neurons, also known as from T5-L2 or 3 and pass through the anterior roots to
postganglionic fibers, which proceed to the eyeball. They the anterior rami and white communicating branches of
supply the ciliaris muscle and sphincter muscle. No sym- the thoracic and upper lumbar spinal nerves to reach the
pathetic afferent fibers are connected with this nerve. The sympathetic trunks. The fibers pass through the paraver-
sympathetic efferent fibers of the facial nerve travel to the tebral ganglia of the trunks without synapsing to enter the
salivary glands, submaxillary, and sublingual glands. abdominopelvic splanchnic nerves, which convey fibers
Other fibers function as vasodilator and secretory fibers to the prevertebral ganglia of the abdominal cavity. The
to the mucus membrane of the nose, soft palate, tonsils, abdominopelvic splanchnic nerves include the lower tho-
uvula, roof of the mouth, gums, upper lips, parotid, and racic splanchnic nerves (greater, lesser, and least), and
orbital glands. A few sympathetic afferent fibers con- the lumbar splanchnic nerves. The main source of presyn-
nected to the facial nerve have been reported. The glosso- aptic SNS fibers is the lower thoracic splanchnic nerves.
pharyngeal nerve has sympathetic afferent fibers that pass The greater (T5-T9 or T10), lesser (T10-T11), and the least
to the parotid glands, the lower gums, mucus membrane (T12) travel to the celiac, superior, mesenteric, and aorti-
and glands of the tongue, and the floor of the mouth. corenal sympathetic ganglia, respectively. The lumbar
The preganglionic sympathetic efferent fibers of the vagus splanchnic nerves arise from sympathetic trunks, which
nerve end in the sympathetic ganglia near the organs sup- then pass to the intermesenteric, inferior mesenteric, and
plied by the vagus nerve. These organs include the heart, superior hypogastric plexuses. The primary function of
esophagus, stomach, small intestine, and the greater part of these nerves is vasoconstriction and to inhibit peristalsis.
the large intestine. The postganglionic efferent fibers
from the associated ganglia terminate on the smooth Sympathetic Nervous System Splanchnic
muscles of these organs. Some fibers are distributed
Nerves
to the bronchial tree, gallbladder, and bile ducts. In
addition, the vagus nerve provides secretory fibers to the The splanchnic nerves convey visceral efferent and afferent
stomach and pancreas. The afferent fibers of the vagus fibers to and from the viscera of the body cavities. The car-
nerve are also distributed to similar organs that the efferent diopulmonary splanchnic nerves from C4-T5 are destined
fibers terminate on. for the viscera of the thoracic cavity: the heart, larynx,
504 PART IV Anatomy of the Spinal Nerves and Cervical Plexus

trachea, bronchi, and lungs. The presynaptic sympathetic Vasodilator fibers are distributed to the same organs as well
fibers involved in the innervations of the viscera of the as the external genitalia. Inhibitory fibers also terminate on
abdominopelvic cavity (stomach and intestines) pass to external genitalia. The afferent sacral sympathetic fibers
the prevertebral ganglia through abdominopelvic splanchnic follow a similar distribution by conduction impulses from
nerves. Regarding the sympathetic innervation in the the pelvic viscera to the second, third, and fourth sacral
abdomen, preganglionic splanchnic nerves from the thoracic nerves, where they originate in the spinal ganglia.
sympathetic chains descend through the diaphragm to
synapse in preaortic (prevertebral) ganglia. The greater
splanchnic (visceral) nerve, derived from T5 to T9 seg-
SYMPATHETIC NERVOUS SYSTEM INJURY
ments, terminates on the celiac ganglia; the lesser splanchnic Axons respond to injury in three ways: segmental demyelin-
nerve, derived from T10-T11 segments, terminates on the ation, Wallerian degeneration, and axonal degeneration. Seg-
celiac or aorticorenal ganglia; and the least (lowest) mental demyelination and Wallerian degeneration are most
splanchnic (if present), derived from T12, terminates on commonly due to traumatic nerve injury, whereas axonal
the aorticorenal ganglia. A fourth or accessory splanchnic degeneration is more characteristically seen in metabolic
nerve may be found in 4% of cases with a course and termi- and toxic nerve disorders. Segmental demyelination occurs
nation similar to the least splanchnic nerve. The preaortic when a focal segmental nerve is subjected to a mild com-
plexus formed by the aforementioned ganglia and nerves pressive or tractional force, with nerve segments distal and
(derived from both sympathetic fibers and vagal fibers, both proximal to the injury functioning normally. Conduction
visceral sensory and visceral motor) progress inferiorly across the injured segments becomes impaired as the myelin
anterior to the descending abdominal aorta. Preganglionic sheath becomes damaged. Focal demyelination of axons
branches of the sympathetic chain that exit directly without within a fascicle affects some of the fibers causing an asyn-
synapsing are known as the lumbar and sacral splanchnics. chronous conduction, meaning impulses reach their desti-
These branches contribute to the formation of the inter- nation after a delay. More severe compression may lead to
mesenteric plexus, inferior mesenteric plexus, and superior blockage of conduction across the nerve segment resulting
hypogastric plexus, which is the distal prolongation of the in a demyelinating conduction block. Wallerian degeneration
preaortic plexus inferior to the bifurcation of the descending follows second- to fifth-degree injury. The Wallerian deg-
abdominal aorta. Some of the lumbar splanchnics, especially neration is axon and myelin sheath degeneration distal to
the more inferior branches, have a propensity to travel pos- the transaction. Distal segment changes begin with disruption
terior to the common iliac arteries. It should be emphasized of retrograde and anterograde flow of signals within the
that the term splanchnic (referring to the viscera) does not axon. During the disruption, calcium and sodium ions influx
imply whether a nerve is pre- or postganglionic. Although the plasma membrane and initiate apoptosis of the cell body.
the sympathetic splanchnics of the abdomen and pelvis Axonal injury also recruits leukocytes and cytokines to cause
are indeed preganglionic, the splanchnics superior to T5 that changes in the surrounding cells. This entire process is part of
travel primarily to the heart and lungs are postganglionic. the Wallerian degeneration. Macrophages are recruited to the
All presynaptic fibers in the abdominopelvic splanchnic injury site to phagocytose axon and myelin debris. If regen-
nerves, except those involved in innervating the suprarenal eration does not take place proximal to the site of injury,
glands, synapse in prevertebral ganglia. Some presynaptic degeneration takes place. In the proximal segment, the extent
sympathetic fibers pass through the celiac prevertebral of injury determines the extent of apoptosis of the cell body
ganglia without synapsing, directly terminating on cells of or axonal breakdown. Nerve regeneration, opposite of Wal-
the medulla of the suprarenal gland. The suprarenal med- lerian degeneration and demyelination, involves prolifer-
ullary cells (chromaffin cells) function as a postsynaptic ation of Schwann cells and remyelination. Typically,
neuron that does not release neurotransmitter substance onto remyelination occurs after first-degree injuries while col-
the cells of a specific effector organ, but releases it into the lateral axon sprouting and proximal-to-distal nerve regener-
bloodstream to circulate throughout the body. ation occur after second- to fifth-degree lesions. If only a few
of the axons supplying a muscle or organ are damaged, the
intact motor axons reinnervate the denervated muscle fiber
Sympathetic Nervous System Sacral
through a process known as collateral sprouting. These nerve
The sympathetic efferent fibers of the sacral region stem sprouts originate from the nodes of Ranvier or the nerve ter-
from the second, third, and fourth sacral nerves. These pre- minals. The factors that influence the extent and quality of
ganglionic fibers are collected together in the pelvic nerve regeneration includes axon branching, the integrity of the
and proceed to the hypogastric or pelvic plexuses. From endoneurium, the maturation of regenerated sympathetic
here, the postganglionic fibers are distributed to the pelvic nerve fibers, the pattern of innervations and recovery, and
viscera. Motor fibers pass to smooth muscle of organs such compensatory mechanisms (Daroff, Fenichel, Jankovic, &
as the descending colon, rectum, anus, and bladder. Mazziotta, 2012).
Anatomy of the Sympathetic Nervous System Chapter 36 505

Nerve injury can be due to trauma, neoplasms, infection, axonotmesis is comparable to Sunderland’s second-degree
inflammation, hereditary, or is iatrogenic. A nerve injury is injury. Also, neurotmesis under Seddon is comparable to
defined by the mechanism of the injury, the degree of Sunderland’s third-, fourth-, and fifth-degree injuries
injury, and the nerve components affected by the injury. (Sunderland, 1990). MacKinnon introduced a sixth-degree
Nerve injury can be classified by either the Seddon or Sun- injury, which is also considered the combination or mixed
derland classification systems to determine interventions. type of nerve injury (Hebert-Blouin & Spinner, 2011).
The Seddon classification includes the terms neuropraxia,
axonotmesis, and neurotmesis for conduction block, loss of
continuity of axons, and loss of nerve trunk continuity. The
second method of classification is the Sunderland classifi-
Trophic Changes
cation, based on the histologic structure of the nerve trunk. Following a nerve injury, changes to the surrounding tissues
This classification recognizes five degrees or types of nerve may evolve but they are uncommon. Primarily after dener-
damage in increasing severity. The first degree of nerve vation, a loss of sweating and alterations in the blood flow,
injury is a temporary conduction block lesion in which primarily vasodilation, takes place. Most importantly, the
axonal continuity is preserved. This may involve segmental extent of damage and trophic changes determines the suscep-
demyelination. The myelin is injured but a complete and tibility of the affected area to future injury. Trophic changes
fast recovery is expected. A second-degree injury includes include those that take place to the texture of the skin, nails,
injury to both the myelin and axon. The continuity of the and hair; color of the skin; temperature of the skin; and change
endoneurial sheath of the nerve fibers is preserved. Wal- in amount and consistency of subcutaneous tissues. Dener-
lerian degeneration occurs below the lesion and later the vated tissue can be extremely susceptible to cold and injury,
axon affected is replaced by regeneration and function is leading to further injury such as ulcerations. These changes
fully restored. The expected recovery is good but slow. typically are located at the periphery of the limb (hands
Third-degree injury consists of continuous fasciculi and and feet). Factors that influence these changes include the
nerve fibers that are lost. Hence, the myelin, axon, and particular nerve injury, nature of the injury, level of the nerve
endoneurium are injured. The damage can be localized or lesion, duration of denervation, and extent of vasomotor dis-
irregularly distributed along the length of the nerve. The turbances. Trophic changes occur more frequently after com-
internal structure of the fascicule becomes disorganized plete nerve injuries affecting the brachial plexus, median,
with the disintegration of axons undergoing Wallerian ulnar, sciatic, medial popliteal, and tibial nerves. With more
degeneration, the loss of endoneurium, and hemorrhage, extensive vasomotor paralysis, longer time periods of dener-
edema, inflammation or fibrosis from the surrounding vation, and severe proximal neuronal changes, the trophic
tissues. In these injuries, axon regeneration becomes dif- changes increase. The order of appearance of trophic changes
ficult as the fibrosis can block axons and delay the growth is variable, but usually progresses from secondary trauma due
or divert the course of the axons. This can cause a fusiform to insensitivity during reinnervation, dry and warm skin, cold
swelling of the fasciculi. Also, the loss of continuity of the and cyanotic skin, changes in texture and consistency of the
nerve fibers is no longer confined to the endoneurium; the skin, causalgia, and finally severe trauma.
axons can become misdirected during regeneration.
Recovery following a third-degree injury is often slow
and variable. A fourth-degree injury includes damage to
Tests to Determine Injury
the myelin sheath, axon, endoneurium, and perineurium.
Thus, the fascicular structure of the nerve has been Clinical evaluation of nerve injuries typically begins with
destroyed with preservation of the nerve trunk continuity history, and the focus should remain on type of symptom,
by disorganized tissue. Regenerating axons are no longer onset, progression, and location, as well as information
confined to the endoneurium or fasciculi, allowing fibrous about potential causes. Physical and neurological exami-
tissue to replace axon regeneration and arrest the growth. A nation to define the type of injury includes testing of sen-
fourth-degree injury requires excision of the damaged sation (using pinprick and light touch for small fibers and
segment and nerve repair, with no expected recovery vibration for large fibers), proprioception, motor strength,
without medical intervention. A fifth-degree injury is com- and deep tendon reflexes.
posed of injury to the myelin sheath, axon, endoneurium, Cranial nerve, autonomic function, motor weakness,
perineurium, and epineurium. A loss of continuity of the and reflexes are also evaluated. Peripheral nervous system
nerve trunk is the core of this type of injury. No recovery disorders are suspected in patients with generalized or
is expected without medical intervention. Other types of diffuse weakness but no sensory deficits. Clues that a
injuries include mixed or combination and partial injuries. peripheral nervous system disorder may be the cause of
Seddon classification of neuropraxia is similar to Sunder- generalized weakness include patterns of generalized
land’s first-degree injury. Seddon’s classification of weakness with a specific cause, deficits in stocking-glove
506 PART IV Anatomy of the Spinal Nerves and Cervical Plexus

distributions, fasciculations, hypotonia, muscle atrophy, Daroff, R.B., Fenichel, G.M., Jankovic, J., Mazziotta, J.C., 2012. Bradley’s
hyporeflexia, and progressive chronic weakness. neurology in clinical practice, 6th ed. Elsevier Saunders, Philadelphia.
The most common nerve testing to evaluate peripheral Hare, K., Hinsey, J.C., 1942. The autonomic nervous system. Annual
Review of Physiology 4, 407–444.
nerve injuries is electromyography (EMG). However,
Hebert-Blouin, M.N., Spinner, R.J., 2011. Nerve injury and repair. In:
EMG focuses on large myelinated fibers, hiding pure small
Cameron, J.L, Cameron, A.M. (Eds), Current surgical therapy,
fiber neuropathies as associated with normal findings. 10th ed. Elsevier Saunders, Philadelphia.
Other sympathetic testing includes, but is not limited to, Kuznetsov, A.V., 2011. Analytical solution of equations describing slow
induction of sweating (autonomic testing), colorimetric axonal transport based on the stop-and-go hypothesis. Central
tests, electrical skin resistance tests, skin response tests, European Journal of Physics 9 (3), 662–673.
Valsalva maneuvers, blood pressure response tests, body Oztas, E., 2003. Neuronal tracing. Neuroanatomy 2, 2–5.
fluid and tissue analysis, cerebrospinal fluid analysis, and Saladin, K., 2010. Anatomy and physiology: The unity of form and
thermography. function, 6th ed. McGraw-Hill, New York.
Schwartz, J.H., 1979. Axonal transport: Components, mechanisms, and
specificity. Annual Review of Neuroscience 2, 467–504.
REFERENCES Shields, J., 1993. Functional anatomy of the autonomic nervous system.
Brown, A., 2003. Axonal transport of membranous and nonmembranous Journal of Clinical Neurophysiology 10 (2), 2–13.
cargoes: A unified perspective. The Journal of Cell Biology 160 (6), Sunderland, S.S., 1990. The anatomy and physiology of nerve injury.
817–821. Muscle & Nerve 13, 771–784.

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