Download The Autonomic Nervous System

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Introduction (pp. 526–528)
Comparison of the Somatic and Autonomic
Nervous Systems (pp. 526–527)
ANS Divisions (pp. 527–528)
ANS Anatomy (pp. 528–535)
Parasympathetic (Craniosacral) Division
(pp. 529–530)
Sympathetic (Thoracolumbar) Division
(pp. 530–534)
Visceral Reflexes (pp. 534–535)
The Autonomic
Nervous System
ANS Physiology (pp. 535–540)
Neurotransmitters and Receptors
(pp. 535–536)
The Effects of Drugs (p. 536)
Interactions of the Autonomic Divisions
(pp. 536–539)
Control of Autonomic Functioning
(pp. 539–540)
Homeostatic Imbalances of the
ANS (pp. 540–541)
Developmental Aspects of the
ANS (p. 541)
T
he human body is exquisitely sensitive to changes in its internal
environment, and engages in a lifelong struggle to balance competing demands for resources under ever-changing conditions.
Although all body systems contribute, the stability of our internal environment depends largely on the autonomic nervous system (ANS),
the system of motor neurons that innervates smooth and cardiac muscle and glands (Figure 14.1).
At every moment, signals stream from visceral organs into the CNS,
and autonomic nerves make adjustments as necessary to ensure optimal support for body activities. In response to changing conditions, the
ANS shunts blood to “needy” areas, speeds or slows heart rate, adjusts
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Central nervous system (CNS)
Peripheral nervous system (PNS)
Motor (efferent) division
Sensory (afferent)
division
Somatic nervous
system
Autonomic nervous
system (ANS)
Sympathetic
division
Parasympathetic
division
Figure 14.1 Place of the ANS in the structural organization of the nervous system.
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blood pressure and body temperature, and increases or decreases stomach secretions.
Most of this fine-tuning occurs without our awareness or attention. Can you tell when your arteries are constricting or when
your pupils are dilating? Probably not, but if you’ve ever been
stuck in a checkout line, and your full bladder was contracting
as if it had a mind of its own, you’ve been very aware of a visceral
activity. These functions, both those we’re aware of and those
that occur without our awareness or attention, are controlled
by the ANS. Indeed, as the term autonomic (auto = self; nom =
govern) implies, this motor subdivision of the peripheral nervous system has a certain amount of functional independence.
The ANS is also called the involuntary nervous system, which
reflects its subconscious control, or the general visceral motor
system, which indicates the location of most of its effectors.
Introduction
Define autonomic nervous system and explain its
relationship to the peripheral nervous system.
Compare the somatic and autonomic nervous systems
relative to effectors, efferent pathways, and neurotransmitters released.
Compare and contrast the functions of the parasympathetic
and sympathetic divisions.
Comparison of the Somatic
and Autonomic Nervous Systems
In our previous discussions of motor nerves, we have focused
largely on the activity of the somatic nervous system. So, before
describing autonomic nervous system anatomy, we will point
out the major differences between the somatic and autonomic
systems as well as some areas of functional overlap.
Both systems have motor fibers, but the somatic and autonomic nervous systems differ (1) in their effectors, (2) in their
efferent pathways, and (3) to some degree in target organ
responses to their neurotransmitters. Consult Figure 14.2 for a
summary of the differences as we discuss them next.
Effectors
The somatic nervous system stimulates skeletal muscles,
whereas the ANS innervates cardiac and smooth muscle and
glands. Differences in the physiology of the effector organs account for most of the remaining differences between somatic
and autonomic effects on their target organs.
Efferent Pathways and Ganglia
In the somatic nervous system, the motor neuron cell bodies are
in the CNS, and their axons extend in spinal or cranial nerves all
the way to the skeletal muscles they activate. Somatic motor
fibers are typically thick, heavily myelinated group A fibers that
conduct nerve impulses rapidly.
In contrast, the ANS uses a two-neuron chain to its effectors.
The cell body of the first neuron, the preganglionic neuron, resides in the brain or spinal cord. Its axon, the preganglionic
axon, synapses with the second motor neuron, the ganglionic
neuron, in an autonomic ganglion outside the CNS. The axon
of the ganglionic neuron, called the postganglionic axon, extends to the effector organ. If you think about the meanings of
all these terms while referring to Figure 14.2, understanding the
rest of the chapter will be much easier.
Preganglionic axons are lightly myelinated, thin fibers, and
postganglionic axons are even thinner and are unmyelinated.
Consequently, conduction through the autonomic efferent
chain is slower than conduction in the somatic motor system.
For most of their course, many pre- and postganglionic fibers
are incorporated into spinal or cranial nerves.
Keep in mind that autonomic ganglia are motor ganglia, containing the cell bodies of motor neurons. Technically, they are
sites of synapse and information transmission from preganglionic to ganglionic neurons. Also, remember that the somatic
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Chapter 14 The Autonomic Nervous System
Cell bodies in central
nervous system
Neurotransmitter
at effector
Peripheral nervous system
SOMATIC
NERVOUS
SYSTEM
Single neuron from CNS to effector organs
Effector
organs
ACh
Heavily myelinated axon
Effect
+
Skeletal muscle
Stimulatory
Two-neuron chain from CNS to effector organs
NE
SYMPATHETIC
Lightly myelinated
preganglionic axons
Ganglion
ACh
Unmyelinated
postganglionic axon
Acetylcholine (ACh)
+
Epinephrine and
norepinephrine
Adrenal medulla
PARASYMPATHETIC
AUTONOMIC NERVOUS SYSTEM
ACh
Blood vessel
ACh
ACh
Lightly myelinated
preganglionic axon
Unmyelinated
postganglionic
axon
Ganglion
Smooth muscle
(e.g., in gut), glands,
cardiac muscle
Stimulatory
or inhibitory,
depending
on neurotransmitter
and receptors
on effector
organs
Norepinephrine (NE)
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Figure 14.2 Comparison of somatic and autonomic nervous systems.
motor division lacks ganglia entirely. The dorsal root ganglia are
part of the sensory, not the motor, division of the PNS.
Neurotransmitter Effects
All somatic motor neurons release acetylcholine (ACh) at their
synapses with skeletal muscle fibers. The effect is always
excitatory, and if stimulation reaches threshold, the muscle
fibers contract.
Neurotransmitters released onto visceral effector organs by
postganglionic autonomic fibers include norepinephrine (NE)
secreted by most sympathetic fibers, and ACh released by
parasympathetic fibers. Depending on the type of receptors
present on the target organ, the organ’s response may be either
excitation or inhibition (Figure 14.2; see Table 14.2 on p. 536).
Overlap of Somatic and Autonomic Function
Higher brain centers regulate and coordinate both somatic and
autonomic motor activities, and nearly all spinal nerves (and
many cranial nerves) contain both somatic and autonomic
fibers. Moreover, most of the body’s adaptations to changing
internal and external conditions involve both skeletal muscle
activity and enhanced responses of certain visceral organs. For
example, when skeletal muscles are working hard, they need
more oxygen and glucose and so autonomic control mecha-
nisms speed up heart rate and dilate airways to meet these needs
and maintain homeostasis.
The ANS is only one part of our highly integrated nervous system, but according to convention we will consider it an individual
entity and describe its role in isolation in the sections that follow.
ANS Divisions
The two arms of the ANS, the parasympathetic and sympathetic
divisions, generally serve the same visceral organs but cause essentially opposite effects. If one division stimulates certain
smooth muscles to contract or a gland to secrete, the other division inhibits that action. Through this dual innervation, the
two divisions counterbalance each other’s activities to keep
body systems running smoothly. The sympathetic division mobilizes the body during activity, whereas the parasympathetic
arm promotes maintenance functions and conserves body energy. Let’s elaborate on these functional differences by focusing
briefly on extreme situations in which each division is exerting
primary control.
Role of the Parasympathetic Division
The parasympathetic division, sometimes called the “resting
and digesting” system, keeps body energy use as low as possible,
even as it directs vital “housekeeping” activities like digestion
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and elimination of feces and urine. (This explains why it is a
good idea to relax after a heavy meal: so that digestion is not interfered with by sympathetic activity.) Parasympathetic activity
is best illustrated in a person who relaxes after a meal and reads
the newspaper. Blood pressure and heart rate are regulated at
low normal levels, and the gastrointestinal tract is actively digesting food. In the eyes, the pupils are constricted and the
lenses are accommodated for close vision to improve the clarity
of the close-up image.
Parasympathetic
Sympathetic
Eye
Brain stem
Salivary
glands
Heart
Skin*
Cranial
Cervical
Sympathetic
ganglia
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The sympathetic division is often referred to as the “fight-orflight” system. Its activity is evident when we are excited or find
ourselves in emergency or threatening situations, such as being
frightened by street toughs late at night. A rapidly pounding
heart; deep breathing; dry mouth; cold, sweaty skin; and dilated
eye pupils are sure signs of sympathetic nervous system mobilization. Not as obvious, but equally characteristic, are changes
in brain wave patterns and in the electrical resistance of the skin
(galvanic skin resistance)—events that are recorded during lie
detector examinations.
During any type of vigorous physical activity, the sympathetic division also promotes a number of other adjustments.
Visceral (and sometimes cutaneous) blood vessels are constricted, and blood is shunted to active skeletal muscles and the
vigorously working heart. The bronchioles in the lungs dilate,
increasing ventilation (and ultimately increasing oxygen delivery to body cells), and the liver releases more glucose into the
blood to accommodate the increased energy needs of body cells.
At the same time, temporarily nonessential activities, such as
gastrointestinal tract motility, are damped. If you are running
from a mugger, digesting lunch can wait! It is far more important that your muscles be provided with everything they need to
get you out of danger. In such active situations, the sympathetic
division generates a head of steam that enables the body to cope
with situations that threaten homeostasis. Its function is to provide the optimal conditions for an appropriate response to
some threat, whether that response is to run, to see better, or to
think more clearly.
We have just looked at two extreme situations in which one
or the other branch of the ANS dominates. An easy way to remember the most important roles of the two ANS divisions is to
think of the parasympathetic division as the D division [digestion, defecation, and diuresis (urination)], and the sympathetic
division as the E division (exercise, excitement, emergency, embarrassment). A more detailed summary of the effects of each
division on various organs is presented in Table 14.4 (p. 538).
Remember, however, that while we may find it easy to think
of the two ANS divisions as working in an all-or-none fashion
as described above, this is rarely the case. A dynamic antagonism exists between the divisions, and fine adjustments are
made continuously by both.
Salivary
glands
Lungs
Lungs
Role of the Sympathetic Division
Eye
T1
Heart
Stomach
Stomach
Pancreas
Liver and
gallbladder
Thoracic
Pancreas
Liver
and gallbladder
L1
Adrenal
gland
Lumbar
Bladder
Bladder
Genitals
Genitals
Sacral
Figure 14.3 Overview of the subdivisions of the ANS. The
parasympathetic and sympathetic divisions differ anatomically in
(1) the sites of origin of their nerves, (2) the relative lengths of their
preganglionic and postganglionic fibers, and (3) the locations of
their ganglia (indicated here by synapse sites).
*Although sympathetic innervation to the skin is mapped to the cervical region here, all nerves to the periphery carry postganglionic sympathetic fibers.
2. Which relays instructions from the CNS to muscles more
quickly, the somatic nervous system or the ANS? Explain why.
3. Which branch of the ANS would predominate if you were
lying on the beach enjoying the sun and the sound of the
waves? Which branch would predominate if you were on a
surfboard and a shark appeared within a few feet of you?
For answers, see Appendix G.
ANS Anatomy
For the parasympathetic and sympathetic divisions, describe
the site of CNS origin, locations of ganglia, and general
fiber pathways.
Anatomically, the sympathetic and parasympathetic divisions
differ in
C H E C K Y O U R U N D E R S TA N D I N G
1. Name the three types of effectors of the autonomic nervous
system.
1. Their origin sites. Parasympathetic fibers emerge from the
brain and sacral spinal cord (are craniosacral). Sympa-
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Chapter 14 The Autonomic Nervous System
thetic fibers originate in the thoracolumbar region of the
spinal cord.
2. The relative lengths of their fibers. The parasympathetic
division has long preganglionic and short postganglionic
fibers. The sympathetic division has the opposite condition.
3. The location of their ganglia. Most parasympathetic ganglia are located in the visceral effector organs. Sympathetic
ganglia lie close to the spinal cord.
Ciliary
ganglion
CN III
Eye
Lacrimal
gland
CN VII
CN IX
CN X
Pterygopalatine
ganglion
Nasal
mucosa
Submandibular
ganglion
Note that these and other differences are illustrated in Figure 14.3 and summarized in Table 14.1, p. 534.
We begin our detailed exploration of the ANS with the
anatomically simpler parasympathetic division.
Submandibular
and sublingual
glands
Otic ganglion
Parotid gland
Parasympathetic (Craniosacral) Division
Heart
The parasympathetic division is also called the craniosacral
division because its preganglionic fibers spring from opposite
ends of the CNS—the brain stem and the sacral region of the
spinal cord (Figure 14.4). The preganglionic axons extend from
the CNS nearly all the way to the structures to be innervated.
There the axons synapse with ganglionic neurons located in
terminal ganglia that lie very close to or within the target organs. Very short postganglionic axons issue from the terminal
ganglia and synapse with effector cells in their immediate area.
Cardiac and
pulmonary
plexuses
Lung
Liver and
gallbladder
Cranial Outflow
Celiac
plexus
Preganglionic fibers run in the oculomotor, facial, glossopharyngeal, and vagus cranial nerves. Their cell bodies lie in associated
motor cranial-nerve nuclei in the brain stem (see Figures 12.15
and 12.16). We describe the precise locations of the neurons of
the cranial parasympathetics next.
1. Oculomotor nerves (III). The parasympathetic fibers of the
oculomotor nerves innervate smooth muscles in the eyes
that cause the pupils to constrict and the lenses to bulge—
actions needed to focus on close objects. The preganglionic
axons found in the oculomotor nerves issue from the
accessory oculomotor (Edinger-Westphal) nuclei in the midbrain. The cell bodies of the ganglionic neurons are in the
ciliary ganglia within the eye orbits (see Table 13.2, p. 496).
2. Facial nerves (VII). The parasympathetic fibers of the facial
nerves stimulate many large glands in the head. Fibers that
activate the nasal glands and the lacrimal glands of the
eyes originate in the lacrimal nuclei of the pons. The preganglionic fibers synapse with ganglionic neurons in the
pterygopalatine ganglia (tereh-go-palah-tı-n) just posterior to the maxillae. The preganglionic neurons that
stimulate the submandibular and sublingual salivary
glands originate in the superior salivatory nuclei of the
pons and synapse with ganglionic neurons in the
submandibular ganglia, deep to the mandibular angles
(see Table 13.2, p. 498).
3. Glossopharyngeal nerves (IX). The parasympathetics in
the glossopharyngeal nerves originate in the inferior salivatory nuclei of the medulla and synapse in the otic ganglia,
located just inferior to the foramen ovale of the skull. The
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Stomach
Pancreas
S2
Large
intestine
S4
Small
intestine
Pelvic
splanchnic
nerves
Inferior
hypogastric
plexus
Rectum
Urinary
bladder
and ureters
Genitalia (penis, clitoris, and vagina)
Preganglionic
Postganglionic
CN
Cranial nerve
Figure 14.4 Parasympathetic division of the ANS.
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postganglionic fibers course to and activate the parotid
salivary glands anterior to the ears (see Table 13.2, p. 500).
Cranial nerves III, VII, and IX supply the entire parasympathetic innervation of the head; however, only the
preganglionic fibers lie within these three pairs of cranial
nerves—postganglionic fibers do not. Many of the postganglionic fibers “hitch a ride” with branches of the trigeminal
nerve (V), taking advantage of its wide distribution, while
others travel independently to their destinations.
4. Vagus nerves (X). The remaining and major portion of the
parasympathetic cranial outflow is via the vagus (X)
nerves. Between them, the two vagus nerves account for
about 90% of all preganglionic parasympathetic fibers in
the body. They provide fibers to the neck and to nerve
plexuses (interweaving networks of nerves) that serve virtually every organ in the thoracic and abdominal cavities.
The vagal nerve fibers (preganglionic axons) arise mostly
from the dorsal motor nuclei of the medulla and synapse in
terminal ganglia usually located in the walls of the target
organ. Most terminal ganglia are not individually named.
Instead they are collectively called intramural ganglia, literally, “ganglia within the walls.”
As the vagus nerves pass into the thorax, they send
branches to the cardiac plexuses supplying fibers to the
heart that slow heart rate, the pulmonary plexuses serving the lungs and bronchi, and the esophageal plexuses
(ĕ-sofah-jeal) supplying the esophagus.
When the main trunks of the vagus nerves reach the
esophagus, their fibers intermingle, forming the anterior
and posterior vagal trunks, each containing fibers from
both vagus nerves. These vagal trunks then “ride” the
esophagus down to the abdominal cavity. There they send
fibers through the large abdominal aortic plexus [formed
by a number of smaller plexuses (e.g., celiac, superior
mesenteric, and hypogastric) that run along the aorta]
before giving off branches to the abdominal viscera. The
vagus nerves innervate the liver, gallbladder, stomach,
small intestine, kidneys, pancreas, and the proximal half of
the large intestine.
Sacral Outflow
The rest of the large intestine and the pelvic organs are served by
the sacral outflow, which arises from neurons located in the lateral gray matter of spinal cord segments S2–S4. Axons of these
neurons run in the ventral roots of the spinal nerves to the ventral
rami and then branch off to form the pelvic splanchnic nerves,
which pass through the inferior hypogastric (pelvic) plexus in
the pelvic floor (Figure 14.4). Some preganglionic fibers synapse
with ganglia in this plexus, but most synapse in intramural ganglia in the walls of the following organs: distal half of the large intestine, urinary bladder, ureters, and reproductive organs.
Sympathetic (Thoracolumbar) Division
The sympathetic division is anatomically more complex than
the parasympathetic division, partly because it innervates more
organs. It supplies not only the visceral organs in the internal
body cavities but also all visceral structures in the superficial
(somatic) part of the body. This sounds impossible, but there is
an explanation—some glands and smooth muscle structures in
the soma (sweat glands and the hair-raising arrector pili muscles of the skin) require autonomic innervation and are served
only by sympathetic fibers. In addition, all arteries and veins (be
they deep or superficial) have smooth muscle in their walls that
is innervated by sympathetic fibers. But we will explain these
matters later—let us get on with the anatomy of the sympathetic division.
All preganglionic fibers of the sympathetic division arise
from cell bodies of preganglionic neurons in spinal cord segments T1 through L2 (Figure 14.3). For this reason, the sympathetic division is also referred to as the thoracolumbar division
(thorah-ko-lumbar). The presence of numerous preganglionic sympathetic neurons in the gray matter of the spinal
cord produces the lateral horns—the so-called visceral motor
zones (see Figures 12.31b, p. 469, and 12.32, p. 470). The lateral
horns are just posterolateral to the ventral horns that house
somatic motor neurons. (Parasympathetic preganglionic neurons in the sacral cord are far less abundant than the comparable
sympathetic neurons in the thoracolumbar regions, and lateral
horns are absent in the sacral region of the spinal cord. This is a
major anatomical difference between the two divisions.)
After leaving the cord via the ventral root, preganglionic
sympathetic fibers pass through a white ramus communicans
[plural: rami communicantes (kom-munı̆-kantēz)] to enter
an adjoining sympathetic trunk ganglion forming part of the
sympathetic trunk (or sympathetic chain, Figure 14.5). Looking
like strands of glistening white beads, the sympathetic trunks
flank each side of the vertebral column. The sympathetic trunk
ganglia are also called chain ganglia or paravertebral (“near the
vertebrae”) ganglia.
Although the sympathetic trunks extend from neck to
pelvis, sympathetic fibers arise only from the thoracic and lumbar cord segments, as shown in Figure 14.3. The ganglia vary in
size, position, and number, but typically there are 23 in each
sympathetic trunk—3 cervical, 11 thoracic, 4 lumbar, 4 sacral,
and 1 coccygeal.
Once a preganglionic axon reaches a trunk ganglion, one of
three things can happen to the axon, as shown by the three pathways in Figure 14.5b:
1
The axon can synapse with a ganglionic neuron in the same
trunk ganglion.
2 The axon can ascend or descend the sympathetic trunk to
synapse in another trunk ganglion. (These fibers running
from one ganglion to another connect the ganglia together
to form the sympathetic trunk.)
3 The axon can pass through the trunk ganglion and emerge
from the sympathetic trunk without synapsing.
Preganglionic fibers following the third pathway help form
several splanchnic nerves (splanknik) that synapse in collateral,
or prevertebral, ganglia located anterior to the vertebral column. Unlike sympathetic trunk ganglia, the collateral ganglia
are neither paired nor segmentally arranged and occur only in
the abdomen and pelvis.
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Chapter 14 The Autonomic Nervous System
Spinal cord
Dorsal root
Dorsal root
Dorsal root ganglion
Ventral root
Lateral horn
(visceral
motor zone)
Dorsal ramus of
spinal nerve
Rib
Ventral ramus of
spinal nerve
Sympathetic
trunk ganglion
Gray ramus
communicans
Sympathetic
trunk
Ventral ramus
of spinal nerve
Ventral root
White ramus
communicans
Sympathetic
trunk ganglion
Sympathetic trunk
1 Synapse at the same level
Gray ramus
communicans
White ramus
communicans
Thoracic
splanchnic nerves
(a) Location of the sympathetic trunk
Skin (arrector
pili muscles
and sweat
glands)
To effector
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Blood vessels
2 Synapse at a higher or lower level
Figure 14.5 Sympathetic trunks and pathways. (a) Diagram
of the right sympathetic trunk in the posterior thorax, along the
side of the vertebral column. (b) Synapses between preganglionic
and ganglionic sympathetic neurons can occur at three different
locations—either in a sympathetic trunk ganglion at the same or
a different level, or in a collateral ganglion.
Splanchnic nerve
Collateral ganglion
(such as the celiac)
Target organ
in abdomen
(e.g., intestine)
3 Synapse in a distant collateral
ganglion anterior to the vertebral
column
(b) Three pathways of sympathetic innervation
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Regardless of where the synapse occurs, all sympathetic ganglia are close to the spinal cord, and their postganglionic fibers
are typically much longer than their preganglionic fibers. Recall
that the opposite condition exists in the parasympathetic division, an important anatomical distinction.
Pathways with Synapses in Trunk Ganglia
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When synapses are made in sympathetic trunk ganglia, the
postganglionic axons enter the ventral (or dorsal) ramus of the
adjoining spinal nerves by way of communicating branches
called gray rami communicantes (Figure 14.5). From there
they travel via branches of the rami to their effectors, including
sweat glands and arrector pili muscles of the skin. Anywhere
along their path, the postganglionic axons may transfer over to
nearby blood vessels and innervate the vascular smooth muscle
all the way to their final branches.
Notice that the naming of the rami communicantes as white
or gray reflects their appearance, revealing whether or not their
fibers are myelinated (and has no relationship to the white and
gray matter of the CNS). Preganglionic fibers composing the
white rami are myelinated. Postganglionic axons forming the
gray rami are not.
The white rami, which carry preganglionic axons to the sympathetic trunks, are found only in the T1–L2 cord segments, regions of sympathetic outflow. However, gray rami carrying
postganglionic fibers headed for the periphery issue from every
trunk ganglion from the cervical to the sacral region, allowing
sympathetic output to reach all parts of the body. Note that rami
communicantes are associated only with the sympathetic division
and never carry parasympathetic fibers.
Pathways to the Head Sympathetic preganglionic fibers serving the head emerge from spinal cord segments T1–T4 and ascend the sympathetic trunk to synapse with ganglionic neurons
in the superior cervical ganglion (Figure 14.6). This ganglion
contributes sympathetic fibers that run in several cranial nerves
and with the upper three or four cervical spinal nerves. Besides
serving the skin and blood vessels of the head, its fibers stimulate the dilator muscles of the irises of the eyes, inhibit the nasal
and salivary glands (the reason your mouth goes dry when you
are scared), and innervate the smooth (tarsal) muscle that lifts
the upper eyelid. The superior cervical ganglion also sends direct branches to the heart.
Sympathetic preganglionic fibers
innervating the thoracic organs originate at T1–T6. From there
the preganglionic fibers run to synapse in the cervical trunk
ganglia. Postganglionic fibers emerging from the middle and
inferior cervical ganglia enter cervical nerves C4–C8 (Figure 14.6). Some of these fibers innervate the heart via the
cardiac plexus, and some innervate the thyroid gland, but most
serve the skin. Additionally, some T1–T6 preganglionic fibers
synapse in the nearest trunk ganglion, and the postganglionic
fibers pass directly to the organ served. Fibers to the heart, aorta,
lungs, and esophagus take this direct route. Along the way, they
run into the plexuses associated with those organs.
Pathways to the Thorax
Pathways with Synapses in Collateral Ganglia
Most of the preganglionic fibers from T5 down synapse in collateral ganglia, and so most of these fibers enter and leave the sympathetic trunks without synapsing. They form several nerves
called splanchnic nerves, including the thoracic splanchnic
nerves (greater, lesser, and least) and the lumbar and sacral
splanchnic nerves.
The splanchnic nerves contribute to a number of interweaving nerve plexuses known collectively as the abdominal aortic
plexus, which clings to the surface of the abdominal aorta. This
complex plexus contains several ganglia that together serve the
abdominopelvic viscera (splanchni = viscera). From superior to
inferior, the most important of these ganglia (and related subplexuses) are the celiac, superior mesenteric, and inferior
mesenteric, named for the arteries with which they most closely
associate (Figure 14.6). Postganglionic fibers issuing from these
ganglia generally travel to their target organs in the company of
the arteries serving these organs.
Sympathetic innervation of the
abdomen is via preganglionic fibers from T5 to L2, which travel
in the thoracic splanchnic nerves to synapse mainly at the celiac
and superior mesenteric ganglia. Postganglionic fibers issuing
from these ganglia serve the stomach, intestines (except the distal half of the large intestine), liver, spleen, and kidneys.
Pathways to the Abdomen
Pathways to the Pelvis Preganglionic fibers innervating the
pelvis originate from T10 to L2 and then descend in the sympathetic trunk to the lumbar and sacral trunk ganglia. Some fibers
synapse there and the postganglionic fibers run in lumbar and
sacral splanchnic nerves to plexuses on the lower aorta and in
the pelvis. Other preganglionic fibers pass directly to these autonomic plexuses and synapse in collateral ganglia, such as the
inferior mesenteric ganglion. Postganglionic fibers proceed
from these plexuses to the pelvic organs (the urinary bladder
and reproductive organs) and also the distal half of the large
intestine. For the most part, sympathetic fibers inhibit the
activity of the muscles and glands in these visceral organs.
Pathways with Synapses in the Adrenal Medulla
Some fibers traveling in the thoracic splanchnic nerves pass
through the celiac ganglion without synapsing and terminate by
synapsing with the hormone-producing medullary cells of the
adrenal gland (Figure 14.6). When stimulated by preganglionic
fibers, the medullary cells secrete NE and epinephrine (also
called noradrenaline and adrenaline, respectively) into the
blood, producing the excitatory effects we have all felt as a
“surge of adrenaline.” Embryologically, sympathetic ganglia and
the adrenal medulla arise from the same tissue. For this reason,
the adrenal medulla is sometimes viewed as a “misplaced” sympathetic ganglion, and its hormone-releasing cells, although
lacking nerve processes, are considered equivalent to ganglionic
sympathetic neurons.
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Chapter 14 The Autonomic Nervous System
Eye
Lacrimal gland
Nasal mucosa
Pons
Sympathetic trunk
(chain) ganglia
Blood vessels;
skin (arrector pili
muscles and
sweat glands)
Superior
cervical
ganglion
Salivary glands
Middle
cervical
ganglion
Inferior
cervical
ganglion
T1
Heart
Cardiac and
pulmonary
plexuses
Lung
Greater splanchnic nerve
Lesser splanchnic nerve
Liver and
gallbladder
Celiac ganglion
L2
Stomach
White rami
communicantes
Superior
mesenteric
ganglion
Spleen
Adrenal medulla
Kidney
Sacral
splanchnic
nerves
Lumbar
splanchnic nerves
Small
intestine
Inferior
mesenteric
ganglion
Large
intestine
Rectum
Preganglionic
Postganglionic
Genitalia (uterus, vagina, and
penis) and urinary bladder
Figure 14.6 Sympathetic division of the ANS. Sympathetic innervation to peripheral
structures (blood vessels, glands, and arrector pili muscles) occurs in all areas but is shown only
in the cervical area.
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TABLE 14.1
Anatomical and Physiological Differences Between
the Parasympathetic and Sympathetic Divisions
CHARACTERISTIC
PARASYMPATHETIC
SYMPATHETIC
Origin
Craniosacral outflow: brain stem nuclei of cranial nerves III, VII, IX, and X;
spinal cord segments S2–S4.
Thoracolumbar outflow: lateral horns of gray matter of spinal cord
segments T1–L2.
Location of ganglia
Ganglia (terminal ganglia) are within the visceral organ (intramural) or
close to the organ served.
Ganglia are within a few centimeters of CNS: alongside vertebral
column (sympathetic trunk ganglia) and anterior to vertebral
column (collateral, or prevertebral, ganglia).
Relative length of preand postganglionic fibers
Long preganglionic; short postganglionic.
Short preganglionic; long postganglionic.
Rami communicantes
None.
Gray and white rami communicantes. White rami contain myelinated preganglionic fibers; gray contain unmyelinated postganglionic fibers.
Degree of branching of
preganglionic fibers
Minimal.
Extensive.
Functional role
Maintenance functions; conserves
and stores energy; “rest and digest.”
Prepares body for activity; “fight-or-flight.”
Neurotransmitters
All preganglionic and postganglionic
fibers release ACh (are cholinergic
fibers).
All preganglionic fibers release ACh. Most postganglionic fibers
release norepinephrine (are adrenergic fibers); postganglionic fibers
serving sweat glands and some blood vessels of skeletal muscles
release ACh. Neurotransmitter activity is augmented by release of
adrenal medullary hormones (norepinephrine and epinephrine).
Stimulus
Visceral Reflexes
14
1 Sensory receptor
Dorsal root ganglion
in viscera
2 Visceral sensory
neuron
3 Integration center
•May be preganglionic
neuron (as shown)
•May be a dorsal horn
interneuron
•May be within walls
of gastrointestinal
tract
Spinal cord
Autonomic ganglion
4 Efferent pathway
(two-neuron chain)
•Preganglionic neuron
•Ganglionic neuron
5 Visceral effector
Response
Figure 14.7 Visceral reflexes. Visceral reflex arcs have the same
five elements as somatic reflex arcs. The visceral afferent (sensory)
fibers are found both in spinal nerves (as depicted here) and in autonomic nerves.
Because most anatomists consider the ANS to be a visceral
motor system, the presence of sensory fibers (mostly visceral
pain afferents) is often overlooked. However, visceral sensory
neurons, which send information concerning chemical
changes, stretch, and irritation of the viscera, are the first link
in autonomic reflexes. Visceral reflex arcs have essentially the
same components as somatic reflex arcs—receptor, sensory
neuron, integration center, motor neuron, effector—except that
a visceral reflex arc has two neurons in its motor component
(Figure 14.7; compare with Figure 13.14).
Nearly all the sympathetic and parasympathetic fibers we
have described so far are accompanied by afferent fibers conducting sensory impulses from glands or muscles. This means
that peripheral processes of visceral sensory neurons are found
in cranial nerves VII, IX, and X, splanchnic nerves, and the sympathetic trunk, as well as in spinal nerves.
Like sensory neurons serving somatic structures (skeletal
muscles and skin), the cell bodies of visceral sensory neurons
are located either in the sensory ganglia of associated cranial
nerves or in dorsal root ganglia of the spinal cord. Visceral sensory neurons are also found in sympathetic ganglia where preganglionic neurons synapse.
Furthermore, complete three-neuron reflex arcs (with sensory
neurons, interneurons, and motor neurons) exist entirely within
the walls of the gastrointestinal tract. Neurons composing these
reflex arcs make up the enteric nervous system, which plays an important role in controlling gastrointestinal tract activity. We will
discuss the enteric nervous system in more detail in Chapter 23.
The fact that visceral pain afferents travel along the same
pathways as somatic pain fibers helps explain the phenomenon
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of referred pain, in which pain stimuli arising in the viscera are
perceived as somatic in origin. For example, a heart attack may
produce a sensation of pain that radiates to the superior thoracic wall and along the medial aspect of the left arm. Because
the same spinal segments (T1–T5) innervate both the heart and
the regions to which pain signals from heart tissue are referred,
the brain interprets most such inputs as coming from the more
common somatic pathway. Cutaneous areas to which visceral
pain is commonly referred are shown in Figure 14.8.
C H E C K Y O U R U N D E R S TA N D I N G
535
Heart
Lungs and
diaphragm
Liver
Gallbladder
Appendix
4. State whether each of the following is a characteristic of
the sympathetic or parasympathetic nervous system: short
preganglionic fibers; origin from thoracolumbar region of
spinal cord; terminal ganglia; collateral ganglia, innervates
adrenal medulla.
5. How does a visceral reflex differ from a somatic reflex?
Heart
Liver
Stomach
Pancreas
Small intestine
Ovaries
Colon
Kidneys
Urinary
bladder
For answers, see Appendix G.
Ureters
ANS Physiology
Define cholinergic and adrenergic fibers, and list the
different types of their receptors.
Describe the clinical importance of drugs that mimic or
inhibit adrenergic or cholinergic effects.
State the effects of the parasympathetic and sympathetic
divisions on the following organs: heart, blood vessels, gastrointestinal tract, lungs, adrenal medulla, and external
genitalia.
Describe autonomic nervous system controls.
Neurotransmitters and Receptors
The major neurotransmitters released by ANS neurons are
acetylcholine (ACh) and norepinephrine (NE). ACh, the same
neurotransmitter secreted by somatic motor neurons, is released
by (1) all ANS preganglionic axons and (2) all parasympathetic
postganglionic axons at synapses with their effectors. AChreleasing fibers are called cholinergic fibers (kolin-erjik).
In contrast, most sympathetic postganglionic axons release
NE and are classified as adrenergic fibers (adren-erjik). Some
exceptions are sympathetic postganglionic fibers innervating
sweat glands and some blood vessels in skeletal muscles. These
fibers secrete ACh.
Unfortunately for memorization purposes, the effects of
ACh and NE on their effectors are not consistently either excitation or inhibition. Why not? The response of visceral effectors
to these neurotransmitters depends not only on the neurotransmitter but also on the receptor to which it attaches. The two or
more kinds of receptors for each autonomic neurotransmitter
allow it to exert different effects (activation or inhibition) at different body targets. Table 14.2 provides a comprehensive summary of the receptor types that we introduce next.
Figure 14.8 Map of referred pain. This map shows the anterior
skin areas to which pain is referred from certain visceral organs.
Cholinergic Receptors
The two types of receptors that bind ACh are named for drugs
that bind to them and mimic acetylcholine’s effects. The first
of these receptors identified were the nicotinic receptors
(niko-tinik), which respond to nicotine. A mushroom poison,
muscarine (muskah-rin), activates a different set of ACh receptors, named muscarinic receptors. All ACh receptors are either
nicotinic or muscarinic.
Nicotinic receptors are found on (1) the sarcolemma of
skeletal muscle cells at neuromuscular junctions (which, as you
will recall, are somatic and not autonomic targets), (2) all ganglionic neurons, both sympathetic and parasympathetic, and
(3) the hormone-producing cells of the adrenal medulla. The
effect of ACh binding to nicotinic receptors anywhere is always
stimulatory. Just as at the sarcolemma of skeletal muscle (examined in Chapter 9), ACh binding to any nicotinic receptor directly opens ion channels, depolarizing the postsynaptic cell.
Muscarinic receptors occur on all effector cells stimulated by
postganglionic cholinergic fibers—that is, all parasympathetic
target organs and a few sympathetic targets, such as eccrine
sweat glands and some blood vessels of skeletal muscles. The effect of ACh binding to muscarinic receptors can be either inhibitory or stimulatory, depending on the subclass of
muscarinic receptor found on the target organ. For example,
binding of ACh to cardiac muscle receptors slows heart activity,
whereas ACh binding to receptors on smooth muscle of the gastrointestinal tract increases its motility.
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TABLE 14.2
Cholinergic and Adrenergic Receptors
MAJOR LOCATIONS *
EFFECT OF BINDING
Nicotinic
All ganglionic neurons; adrenal medullary
cells (also neuromuscular junctions of skeletal
muscle)
Excitation
Muscarinic
All parasympathetic target organs
Excitation in most cases; inhibition of cardiac
muscle
NEUROTRANSMITTER
RECEPTOR TYPE
Acetylcholine
Cholinergic
Limited sympathetic targets:
Norepinephrine (and
epinephrine released
by adrenal medulla)
Adrenergic
1
■
Eccrine sweat glands
Activation
■
Blood vessels in skeletal muscles
Vasodilation (may not occur in humans)
Heart predominantly, but also kidneys and
adipose tissue
Increases heart rate and strength; stimulates
renin release by kidneys
2
Lungs and most other sympathetic target
organs; abundant on blood vessels serving the
heart, liver and skeletal muscle
Effects mostly inhibitory; dilates blood vessels
and bronchioles; relaxes smooth muscle walls
of digestive and urinary visceral organs;
relaxes uterus
3
Adipose tissue
Stimulates lipolysis by fat cells
1
Most importantly blood vessels serving the
skin, mucosae, abdominal viscera, kidneys,
and salivary glands; also, virtually all sympathetic target organs except heart
Constricts blood vessels and visceral organ
sphincters; dilates pupils of the eyes
2
Membrane of adrenergic axon terminals;
pancreas; blood platelets
Inhibits NE release from adrenergic terminals;
inhibits insulin secretion by pancreas; promotes
blood clotting
14
* Note that all of these receptor subtypes are also found in the CNS.
Adrenergic Receptors
There are also two major classes of adrenergic (NE-binding) receptors: alpha () and beta (). These receptors are further divided into subclasses (1 and 2; 1, 2, and 3). Organs that
respond to NE (or to epinephrine) have one or more of these
receptor subtypes.
NE or epinephrine can have either excitatory or inhibitory effects on target organs depending on which subclass of receptor
predominates in that organ. For example, binding of NE to the 1
receptors of cardiac muscle prods the heart into more vigorous
activity, whereas epinephrine binding to 2 receptors in bronchiole smooth muscle causes it to relax, dilating the bronchiole.
The Effects of Drugs
Knowing the locations of the cholinergic and adrenergic receptor subtypes allows specific drugs to be prescribed to obtain the
desired inhibitory or stimulatory effects on selected target
organs. For example, atropine is an anticholinergic drug that
blocks muscarinic ACh receptors. It is routinely administered
before surgery to prevent salivation and to dry up respiratory
system secretions. Ophthalmologists also use it to dilate the
pupils for eye examination. The anticholinesterase drug
neostigmine inhibits the enzyme acetylcholinesterase, preventing enzymatic breakdown of ACh and allowing it to accumulate
in synapses. This drug is used to treat myasthenia gravis, a con-
dition in which skeletal muscle activity is impaired for lack of
ACh stimulation.
As we described in Chapter 11, NE is one of our “feeling
good” neurotransmitters, and drugs that prolong the activity of
NE on the postsynaptic membrane help to relieve depression.
Hundreds of over-the-counter drugs used to treat colds, coughs,
allergies, and nasal congestion contain sympathomimetics
(phenylephrine and others), sympathetic-mimicking drugs that
stimulate -adrenergic receptors.
Much pharmaceutical research is directed toward finding
drugs that affect only one subclass of receptor without upsetting
the whole adrenergic or cholinergic system. An important
breakthrough was finding drugs that mainly activate 2 receptors. People with asthma use such 2 activators to dilate their
lung bronchioles without activating 1 receptors, which would
increase their heart rate. Selected drug classes that influence
ANS activity are summarized in Table 14.3.
Interactions of the Autonomic Divisions
As we mentioned earlier, most visceral organs receive dual
innervation. Normally, both ANS divisions are partially active,
producing a dynamic antagonism that allows visceral activity to
be precisely controlled. However, one division or the other usually exerts the predominant effects in given circumstances, and
in a few cases, the two divisions actually cooperate with each
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TABLE 14.3
537
Selected Drug Classes That Influence the Activity of the Autonomic Nervous System
DRUG CLASS
RECEPTOR BOUND
EFFECTS
EXAMPLE
CLINICAL USE
Nicotinic agents (little
therapeutic value, but
important because of
presence of nicotine in
tobacco)
Nicotinic ACh receptors on all ganglionic
neurons and in CNS
Typically stimulation of
sympathetic effects; blood
pressure increases
Nicotine
Used in smoking cessation
products
Parasympathomimetic
agents (muscarinic
agents)
Muscarinic ACh
receptors
Mimic effects of ACh,
enhance parasympathetic
effects
Pilocarpine
Glaucoma (opens aqueous
humor drainage pores)
Bethanechol
Difficulty urinating (increases
bladder contraction)
Neostigmine
Myasthenia gravis, (increases
availability of ACh)
Sarin
Used as chemical warfare
agent (similar to widely used
insecticides)
Albuterol (Ventolin)
Asthma (dilates bronchioles
by binding to 2 receptors)
Phenylephrine
Colds (nasal decongestant,
binds to 1 receptors)
Propranolol
Hypertension (member of
a class of drugs called betablockers that decrease heart
rate and blood pressure)
Acetylcholinesterase
inhibitors
Sympathomimetic
agents
Sympatholytic agents
None; bind to the
enzyme (AChE) that
degrades ACh
Adrenergic receptors
Adrenergic receptors
Indirect effect at all ACh
receptors; prolong the
effect of ACh
Enhance sympathetic
activity by increasing NE
release or binding to
adrenergic receptors
Decrease sympathetic
activity by blocking
adrenergic receptors or
inhibiting NE release
other. Table 14.4 contains an organ-by-organ summary of the
differing effects of the two divisions.
Antagonistic Interactions
Antagonistic effects, described earlier, are most clearly seen on
the activity of the heart, respiratory system, and gastrointestinal
organs. In a fight-or-flight situation, the sympathetic division
increases heart rate, dilates airways, and inhibits digestion and
elimination. When the emergency is over, the parasympathetic
division restores heart rate and airway diameter to resting levels
and then attends to processes that refuel your body cells and discard wastes.
Sympathetic and Parasympathetic Tone
We have described the parasympathetic division as the “resting
and digesting” division, but the sympathetic division is the major
actor in controlling blood pressure, even at rest. With few exceptions, the vascular system is entirely innervated by sympathetic
fibers that keep the blood vessels in a continual state of partial
constriction called sympathetic, or vasomotor, tone. When a
higher blood pressure is needed to maintain blood flow, the sympathetic fibers fire more rapidly, causing blood vessels to constrict and blood pressure to rise. When blood pressure is to be
decreased, sympathetic fibers fire less rapidly and the vessels dilate. Alpha-blockers, drugs that interfere with the activity of these
vasomotor fibers, are sometimes used to treat hypertension.
During circulatory shock (inadequate blood delivery to body
tissues), or when more blood is needed to meet the increased
needs of working skeletal muscles, blood vessels serving the skin
and abdominal viscera are strongly constricted. This blood
“shunting” helps maintain circulation to vital organs or enhance blood delivery to skeletal muscles.
On the other hand, parasympathetic effects normally dominate the heart and the smooth muscle of digestive and urinary
tract organs. These organs exhibit parasympathetic tone. The
parasympathetic division slows the heart and dictates the normal activity levels of the digestive and urinary tracts. However,
the sympathetic division can override these parasympathetic
effects during times of stress. Drugs that block parasympathetic
responses increase heart rate and cause fecal and urinary retention. Except for the adrenal glands and sweat glands of the skin,
most glands are activated by parasympathetic fibers.
Cooperative Effects
The best example of cooperative ANS effects is seen in controls
of the external genitalia. Parasympathetic stimulation causes
vasodilation of blood vessels in the external genitalia and is responsible for erection of the male penis or female clitoris during
sexual excitement. (This may explain why sexual performance is
sometimes impaired when people are anxious or upset and the
sympathetic division is in charge.) Sympathetic stimulation
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TABLE 14.4
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Effects of the Parasympathetic and Sympathetic Divisions on Various Organs
TARGET ORGAN OR SYSTEM
PARASYMPATHETIC EFFECTS
SYMPATHETIC EFFECTS
Eye (iris)
Stimulates sphincter pupillae muscles; constricts
pupils
Stimulates dilator pupillae muscles; dilates pupils
Eye (ciliary muscle)
Stimulates muscle, which results in bulging of the
lens for close vision
Weakly inhibits muscle, which results in flattening
of the lens for far vision
Glands (nasal, lacrimal,
gastric, pancreas)
Stimulates secretory activity
Inhibits secretory activity; causes vasoconstriction
of blood vessels supplying the glands
Salivary glands
Stimulates secretion of watery saliva
Stimulates secretion of thick, viscous saliva
Sweat glands
No effect (no innervation)
Stimulates copious sweating (cholinergic fibers)
Adrenal medulla
No effect (no innervation)
Stimulates medulla cells to secrete epinephrine
and norepinephrine
Arrector pili muscles attached
to hair follicles
No effect (no innervation)
Stimulates contraction (erects hairs and produces
“goosebumps”)
Heart (muscle)
Decreases rate; slows heart
Increases rate and force of heartbeat
Heart (coronary blood vessels)
No effect (no innervation)
Causes vasodilation*
Urinary bladder/urethra
Causes contraction of smooth muscle of bladder
wall; relaxes urethral sphincter; promotes voiding
Causes relaxation of smooth muscle of bladder
wall; constricts urethral sphincter; inhibits voiding
Lungs
Constricts bronchioles
Dilates bronchioles*
Digestive tract organs
Increases motility (peristalsis) and amount of secretion by digestive organs; relaxes sphincters to
allow movement of foodstuffs along tract
Decreases activity of glands and muscles of digestive system and constricts sphincters (e.g., anal
sphincter)
Liver
Increases glucose uptake from blood
Stimulates release of glucose to blood*
Gallbladder
Excites (gallbladder contracts to expel bile)
Inhibits (gallbladder is relaxed)
Kidney
No effect (no innervation)
Promotes renin release; causes vasoconstriction;
decreases urine output
Penis
Causes erection (vasodilation)
Causes ejaculation
Vagina/clitoris
Causes erection (vasodilation) of clitoris; increases
vaginal lubrication
Causes contraction of vagina
Blood vessels
Little or no effect
Constricts most vessels and increases blood pressure; constricts vessels of abdominal viscera and
skin to divert blood to muscles, brain, and heart
when necessary; NE constricts most vessels; epinephrine dilates vessels of the skeletal muscles
during exercise*
Blood coagulation
No effect (no innervation)
Increases coagulation*
Cellular metabolism
No effect (no innervation)
Increases metabolic rate*
Adipose tissue
No effect (no innervation)
Stimulates lipolysis (fat breakdown)
*
Effects are mediated by epinephrine release into the bloodstream from the adrenal medulla.
then causes ejaculation of semen by the penis or reflex contractions of a female’s vagina.
H O M E O S TAT I C I M B A L A N C E
Autonomic neuropathy (damage to autonomic nerves) is a
common complication of diabetes mellitus. One of the earliest
and most troubling symptoms is sexual dysfunction, with up to
75% of male diabetics experiencing erectile dysfunction. Women,
on the other hand, often experience reduced vaginal lubrication. Other frequent manifestations of autonomic neuropathy
include dizziness after standing suddenly (poor blood pressure
control), urinary incontinence, sluggish eye pupil reactions, and
impaired sweating. Just how the elevated blood glucose levels in
diabetics damage nerves is still a mystery. ■
Unique Roles of the Sympathetic Division
The adrenal medulla, sweat glands and arrector pili muscles of
the skin, the kidneys, and most blood vessels receive only sympathetic fibers. It is easy to remember that the sympathetic system innervates these structures because most of us sweat under
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stress, our scalp “prickles” during fear, and our blood pressure
skyrockets (from widespread vasoconstriction) when we get excited. We have already described how sympathetic control of
blood vessels regulates blood pressure and shunting of blood in
the vascular system. We will now consider several other
uniquely sympathetic functions.
Thermoregulatory Responses to Heat The sympathetic division mediates reflexes that regulate body temperature. For example, applying heat to the skin causes reflexive dilation of
blood vessels in that area. When systemic body temperature is
elevated, sympathetic nerves cause the skin’s blood vessels to dilate, allowing the skin to become flushed with warm blood, and
activate the sweat glands to help cool the body. When body temperature falls, skin blood vessels are constricted, and, as a result,
blood is restricted to deeper, more vital organs.
Sympathetic impulses
stimulate the kidneys to release renin, a hormone that promotes
an increase in blood pressure. We describe this renin-angiotensin
mechanism in Chapter 25.
Release of Renin from the Kidneys
Through both direct neural stimulation
and release of adrenal medullary hormones, the sympathetic division promotes a number of metabolic effects not reversed by
parasympathetic activity. It (1) increases the metabolic rate of
body cells; (2) raises blood glucose levels; and (3) mobilizes fats
for use as fuels. The medullary hormones also cause skeletal
muscle to contract more strongly and quickly.
As a side effect, muscle spindles are stimulated more often
and, consequently, nerve impulses traveling to the muscles occur more synchronously. These neural bursts, which put muscle
contractions on a “hair trigger,” are great if you have to make a
quick jump or run, but they can be embarrassing or even disabling to the nervous musician or surgeon.
Metabolic Effects
Localized Versus Diffuse Effects
In the parasympathetic division, one preganglionic neuron
synapses with one (or at most a few) ganglionic neurons. Additionally, all parasympathetic fibers release ACh, which is quickly
destroyed (hydrolyzed) by acetylcholinesterase. Consequently,
the parasympathetic division exerts short-lived, highly localized
control over its effectors.
In contrast, preganglionic sympathetic axons branch profusely as they enter the sympathetic trunk, and they synapse
with ganglionic neurons at several levels. As a result, when the
sympathetic division is activated, it responds in a diffuse and
highly interconnected way. Indeed, the literal translation of
sympathetic (sym = together; pathos = feeling) relates to the
bodywide mobilization this division provokes. Nevertheless,
parts of the sympathetic nervous system can be activated individually. For example, just because your eye pupils dilate in dim
light doesn’t necessarily mean that your heart rate speeds up.
Effects produced by sympathetic activation are much longerlasting than parasympathetic effects. In contrast to the
parasympathetic system’s ACh, NE is inactivated more slowly
because it must be taken back up into the presynaptic ending
539
Communication at
subconscious level
Cerebral cortex
(frontal lobe)
Limbic system
(emotional input)
Hypothalamus
Overall integration
of ANS, the boss
Brain stem
(reticular formation, etc.)
Regulation of pupil size,
respiration, heart, blood
pressure, swallowing, etc.
Spinal cord
Urination, defecation,
erection, and ejaculation
reflexes
Figure 14.9 Levels of ANS control. The hypothalamus stands at
the top of the control hierarchy as the integrator of ANS activity, but
it is influenced by subconscious cerebral inputs via limbic system
connections.
before being hydrolyzed or stored. More importantly, NE and
epinephrine are secreted into the blood by adrenal medullary
cells when the sympathetic division is mobilized. Although epinephrine is more potent at increasing heart rate and raising
blood glucose levels and metabolic rate, these hormones have
essentially the same effects as NE released by sympathetic neurons. In fact, circulating adrenal medullary hormones produce
25–50% of all the sympathetic effects acting on the body at a
given time. These effects continue for several minutes until the
hormones are destroyed by the liver.
In short, sympathetic nerve impulses act only briefly, but the
hormonal effects they provoke linger. The widespread and prolonged effect of sympathetic activation helps explain why we need
time to “come down” after an extremely stressful experience.
Control of Autonomic Functioning
Although the ANS is not usually considered to be under voluntary control, its activity is regulated by CNS controls in the
spinal cord, brain stem, hypothalamus, and cerebral cortex
(Figure 14.9). In general, the hypothalamus is the integrative
center at the top of the ANS control hierarchy. From there, orders flow to lower and lower CNS centers for execution. Although the cerebral cortex may modify the workings of the
ANS, it does so at the subconscious level and by acting through
limbic system structures on hypothalamic centers.
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Brain Stem and Spinal Cord Controls
The hypothalamus is the “boss,” but the brain stem reticular formation appears to exert the most direct influence over autonomic functions (see Figure 12.16 on p. 448). For example,
certain motor centers in the ventrolateral medulla (cardiac and
vasomotor centers) reflexively regulate heart rate and blood vessel diameter. Other medullary regions oversee gastrointestinal
activities. Most sensory impulses involved in eliciting these autonomic reflexes reach the brain stem via vagus nerve afferents.
Although not considered part of the ANS, the medulla and pons
also contain respiratory centers that mediate involuntary control of respiration and receive inputs from the hypothalamus.
Midbrain centers (oculomotor nuclei) control the muscles concerned with pupil diameter and lens focus.
Defecation and micturition reflexes that promote emptying
of the rectum and urinary bladder are integrated at the spinal
cord level but are subject to conscious inhibition. We will describe all of these autonomic reflexes in later chapters in relation
to the organ systems they serve.
Hypothalamic Controls
14
As we noted, the hypothalamus is the main integration center
of the autonomic nervous system. In general, anterior hypothalamic regions direct parasympathetic functions, and posterior areas direct sympathetic functions. These centers exert
their effects both directly and via relays through the reticular
formation, which in turn influences the preganglionic motor
neurons in the brain stem and spinal cord (Figure 14.9). The
hypothalamus contains centers that coordinate heart activity,
blood pressure, body temperature, water balance, and endocrine activity. It also contains centers that mediate various
emotional states (rage, pleasure) and biological drives (thirst,
hunger, sex).
The hypothalamus also mediates our reactions to fear via its
associations with the amygdala and the periaqueductal gray
matter. Emotional responses of the limbic system of the cerebrum to danger and stress signal the hypothalamus to activate
the sympathetic system to fight-or-flight status. In this way, the
hypothalamus serves as the keystone of the emotional and visceral brain, and through its centers emotions influence ANS
functioning and behavior.
Cortical Controls
It was originally believed that the ANS is not subject to voluntary controls. However, we have all had occasions when remembering a frightening event made our heart race (sympathetic
response) or just the thought of a favorite food, pecan pie for
example, made our mouth water (a parasympathetic response).
These inputs converge on the hypothalamus through its connections to the limbic lobe.
Additionally, studies have shown that voluntary cortical control of visceral activities is possible—a capability untapped by
most people.
During
biofeedback training, subjects are connected to monitoring
Influence of Biofeedback on Autonomic Function
devices that provide an awareness of what is happening in their
body. This awareness is called biofeedback. The devices detect
and amplify changes in physiological processes such as heart
rate and blood pressure, and these data are “fed back” in the
form of flashing lights or audible tones. Subjects are asked to try
to alter or control some “involuntary” function by concentrating on calming, pleasant thoughts. The monitor allows them to
identify changes in the desired direction, so they can recognize
the feelings associated with these changes and learn to produce
the changes at will.
Biofeedback techniques have been successful in helping individuals plagued by migraine headaches. They are also used by
cardiac patients to manage stress and reduce their risk of heart
attack. However, biofeedback training is time-consuming and
often frustrating, and the training equipment is expensive and
difficult to use.
C H E C K Y O U R U N D E R S TA N D I N G
6. Name the division of the ANS that does each of the following: increases digestive activity; increases blood pressure;
dilates bronchioles; decreases heart rate; stimulates the
adrenal medulla to release its hormones; causes ejaculation.
7. Would you find nicotinic receptors on skeletal muscle?
Smooth muscle? Eccrine sweat glands? The adrenal medulla?
CNS neurons?
8. Which part of the brain is the main integration center of the
ANS? Which part exerts the most direct influence over autonomic functions?
For answers, see Appendix G.
Homeostatic Imbalances
of the ANS
Explain the relationship of some types of hypertension,
Raynaud’s disease, and autonomic dysreflexia to disorders
of autonomic functioning.
The ANS is involved in nearly every important process that
goes on in the body, so it is not surprising that abnormalities
of autonomic functioning can have far-reaching effects and
threaten life itself. Most autonomic disorders reflect exaggerated or deficient controls of smooth muscle activity. Of these,
the most devastating involve blood vessels and include conditions such as hypertension, Raynaud’s disease, and autonomic
dysreflexia.
Hypertension, or high blood pressure, may result from an
overactive sympathetic vasoconstrictor response promoted by
continuous high levels of stress. Hypertension is always serious
because it increases the workload on the heart, which may precipitate heart disease, and increases the wear and tear on artery
walls. Stress-induced hypertension can be treated with adrenergic receptor–blocking drugs.
Raynaud’s disease is characterized by intermittent attacks
causing the skin of the fingers and toes to become pale, then
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Chapter 14 The Autonomic Nervous System
cyanotic and painful. Commonly provoked by exposure to
cold or emotional stress, it is an exaggerated vasoconstriction
response. The severity of this disease ranges from merely uncomfortable to such severe blood vessel constriction that ischemia and gangrene (tissue death) results. Vasodilators (for
example, adrenergic blockers) usually suffice, but to treat very
severe cases, preganglionic sympathetic fibers serving the affected regions are severed (a procedure called sympathectomy).
The involved vessels then dilate, reestablishing adequate blood
delivery to the region.
Autonomic dysreflexia is a life-threatening condition involving uncontrolled activation of autonomic neurons. It occurs in
a majority of individuals with quadriplegia and in others with
spinal cord injuries above the T6 level, usually in the first year
after injury. The usual trigger is a painful stimulus to the skin
or overfilling of a visceral organ, such as the urinary bladder.
Arterial blood pressure skyrockets to life-threatening levels,
which may rupture a blood vessel in the brain, precipitating
stroke. This may be accompanied by a headache, flushed face,
sweating above the level of the injury, and cold, clammy skin
below. The precise mechanism of autonomic dysreflexia is not
yet clear.
C H E C K Y O U R U N D E R S TA N D I N G
9. Jackson works long, stress-filled shifts as an air traffic
controller at a busy airport. His doctor has prescribed a betablocker. Why might his doctor have done this? What does a
beta-blocker do?
For answers, see Appendix G.
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sensory neurons) (see Figure 12.1, 3 ). Neural crest cells reach
their ultimate destinations by migrating along growing axons.
Forming ganglia receive axons from preganglionic neurons in
the spinal cord or brain and send their axons to synapse with
their effector cells in the body periphery. This process depends
upon the presence of nerve growth factor, and is guided by a
number of signaling chemicals similar to those acting in the
CNS.
During youth, impairments of ANS function are usually due
to injuries to the spinal cord or autonomic nerves. In old age the
efficiency of the ANS begins to decline. At least part of the problem appears to be due to structural changes (bloating) of some
preganglionic axon terminals, which become congested with
neurofilaments.
Many elderly people complain of constipation (a result of reduced gastrointestinal tract motility), and of dry eyes and frequent eye infections (both a result of a diminished ability to
form tears). Additionally, when they stand up they may have
fainting episodes due to orthostatic hypotension (ortho =
straight; stat = standing), a form of low blood pressure that occurs because the aging pressure receptors respond less to changes
in blood pressure following changes in position, and because of
slowed responses by aging sympathetic vasoconstrictor centers.
These problems are distressing, but not usually life threatening,
and most can be managed by lifestyle changes or artificial aids.
For example, changing position slowly gives the sympathetic
nervous system time to adjust the blood pressure, and eye drops
(artificial tears) are available for the dry-eye problem.
C H E C K Y O U R U N D E R S TA N D I N G
10. Which embryonic structure gives rise to both the autonomic
ganglia and the adrenal medulla?
Developmental Aspects
of the ANS
Describe some effects of aging on the autonomic nervous
system.
ANS preganglionic neurons derive from the embryonic neural
tube, as do somatic motor neurons. ANS structures in the
PNS—ganglionic neurons, the adrenal medulla, and all autonomic ganglia—derive from the neural crest (along with all
For answers, see Appendix G.
In this chapter, we have described the structure and function
of the ANS, one arm of the motor division of the peripheral
nervous system. Because virtually every organ system still to be
considered depends on autonomic controls, you will be hearing
more about the ANS in chapters that follow. Now that we have
explored most of the nervous system, this is a good time to examine how it interacts with the rest of the body as summarized
in the Making Connections feature, pp. 542–543.
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Homeostatic Interrelationships Between the Nervous
System and Other Body Systems
Endocrine System
■
■
Sympathetic division of the ANS activates the adrenal medulla;
hypothalamus helps regulate the activity of the anterior pituitary
gland and produces the two posterior pituitary hormones
Hormones influence neuronal metabolism
Cardiovascular System
■
■
ANS helps regulate heart rate and blood pressure
Cardiovascular system provides blood containing oxygen and
nutrients to the nervous system and carries away wastes
Lymphatic System/Immunity
■
■
Nerves innervate lymphoid organs; the brain plays a role in regulating immune function
Lymphatic vessels carry away leaked tissue fluids from tissues
surrounding nervous system structures; immune elements protect all body organs from pathogens (CNS has additional
mechanisms as well)
Respiratory System
■
■
14
Nervous system initiates and regulates respiratory rhythm and
depth
Respiratory system provides life-sustaining oxygen; disposes of
carbon dioxide
Digestive System
■
■
Integumentary System
■
■
Sympathetic division of the ANS regulates sweat glands and
blood vessels of skin (therefore heat loss/retention)
Skin serves as heat loss surface
ANS (particularly the parasympathetic division) regulates digestive motility and glandular activity
Digestive system provides nutrients needed for neuronal health
Urinary System
■
■
ANS regulates bladder emptying and renal blood pressure
Kidneys help to dispose of metabolic wastes and maintain
proper electrolyte composition and pH of blood for neural functioning
Skeletal System
■
■
Nerves innervate bones and joints, providing for pain and joint
sense
Bones serve as depot for calcium needed for neural function;
skeletal system protects CNS structures
Muscular System
■
■
Somatic division of nervous system activates skeletal muscles;
maintains muscle health
Skeletal muscles are the effectors of the somatic division
542
Reproductive System
■
■
ANS regulates sexual erection and ejaculation in males; erection of the clitoris in females
Testosterone causes masculinization of the brain and underlies
sex drive and aggressive behavior
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The Nervous System and Interrelationships with the
Muscular, Respiratory, and Digestive Systems
The nervous system pokes its figurative nose into the activities of
virtually every organ system of the body. Hence, trying to choose
the most significant interactions comes pretty close to a lesson in
futility. Which is more important: digestion, elimination of wastes,
or motility? Your answer probably depends on whether you are
hungry or need to make a “pit stop” at the bathroom when you
read this. Since neural interactions with several other systems are
thoroughly plumbed in chapters to come, here we will look at the
all-important interactions between the nervous system and the
muscular, respiratory, and digestive systems.
Respiratory System
Muscular System
Digestive System
Most simply said, the muscular system would cease to function without the nervous system. Unlike visceral or cardiac muscles, both
of which have other controlling systems, somatic motor fibers are
it for skeletal muscle activation and regulation. Somatic nerve
fibers “tell” skeletal muscles not only when to contract but also how
strongly. Additionally, as the nervous system makes its initial synapses with skeletal muscle fibers, it determines their fate as fast or
slow fibers, which forever after affects our potential for muscle speed
and endurance. The interactions of the various brain regions (basal
nuclei, cerebellum, premotor cortex, etc.) and inputs of stretch
receptors also determine our grace—how smooth and coordinated
we are. Nonetheless, keep in mind that as long as the skeletal muscle effector cells are healthy, they help determine the viability of the
neurons synapsing with them. The relationship is truly synergistic.
Another system that depends entirely on the nervous system for
its function is the respiratory system, which continuously refreshes
the blood with oxygen and unloads the carbon dioxide waste to
the sea of air that surrounds us. Neural centers in the medulla and
pons both initiate and maintain the tidelike rhythm of air flushing
into and out of our lungs by activating skeletal muscles that change
the volume (thus the gas pressure) within the lungs. Peripheral receptors supply information about gas concentrations, lung stretch,
and skeletal muscle activity to these CNS respiratory centers.
Although the digestive system responds to many different types of
controls—for example, hormones, local pH, and irritating chemicals—the parasympathetic nervous system is crucial to its normal
functioning. Without the parasympathetic inputs, sympathetic neural activity, which inhibits normal digestion, would be unopposed.
So important are parasympathetic controls that some of the parasympathetic neurons are actually located in the walls of the digestive organs, in what are called intrinsic plexuses. Thus, even if all
extrinsic controls are severed, the intrinsic mechanisms can still
maintain this crucial body function. The role the digestive system
plays for the nervous system is the same one it offers to all body
systems—it sees that ingested foodstuff gets digested and loaded
into the blood for cell use.
14
Nervous System
Case study: On arrival at Holyoke Hospital, Jimmy Chin, a
10-year-old boy, is immobilized on a rigid stretcher so that he is
unable to move his head or trunk. The paramedics report that
when they found him some 50 feet from the bus, he was awake
and alert, but crying and complaining that he couldn’t “get up to
find his mom” and that he had a “wicked headache.” He has severe bruises on his upper back and head, and lacerations of his
back and scalp. His blood pressure is low, body temperature is below normal, lower limbs are paralyzed, and he is insensitive to painful stimuli below the nipples. Although still alert on arrival, Jimmy
soon begins to drift in and out of unconsciousness.
Jimmy is immediately scheduled for a CT scan, and an operating room is reserved.
Relative to Jimmy’s condition:
1. Why were his head and torso immobilized for transport to the
hospital?
2. What do his worsening neurological signs (drowsiness, incoherence, etc.) probably indicate? (Relate this to the type of surgery
that will be performed.)
3. Assuming that Jimmy’s sensory and motor deficits are due to a
spinal cord injury, at what level do you expect to find a spinal
cord lesion?
4. Two days after his surgery, Jimmy is alert and his MRI scan shows
no residual brain injury, but pronounced swelling and damage
to the spinal cord at T4. On physical examination, Jimmy shows
no reflex activity below the level of the spinal cord injury. His blood
pressure is still low. Why are there no reflexes in his lower limbs
and abdomen?
5. Over the next few days, his reflexes return in his lower limbs and
become exaggerated. He is incontinent. Why is Jimmy hyperreflexive and incontinent?
On one occasion, Jimmy complains of a massive headache and
his blood pressure is way above normal. On examination, he is
sweating intensely above the nipples but has cold, clammy skin below the nipples and his heart rate is very slow.
6. What is this condition called and what precipitates it?
7. How does Jimmy’s excessively high blood pressure put him
at risk?
(Answers in Appendix G)
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