Human Anatomy 4th Edition Saladin

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Chapter 10: The Muscular System I: Introduction

Chapter Overview
Skeletal muscle is the most abundant tissue in the body. Along with bones, it
provides the basis for movement, and is central to the health and activity of an individual.
The term “muscular system” refers only to skeletal muscle but cardiac and smooth
muscle are also briefly discussed in the chapter. Muscle Types and Functions
• Skeletal Muscle
Skeletal muscle consists of striated cell called muscle fibers or myofibers.
Striations give the muscle a striped appearance under the microscope, and are the result
of overlapping arrangement of proteins. Skeletal muscle produces voluntary movement
by attaching to bone. It is sometimes called voluntary muscle because it is subject to
conscious control.
• Cardiac Muscle
Cardiac muscle is found in the walls of the heart, and is responsible for its
contraction. Like skeletal muscle, cardiac muscle is striated, but it is involuntary. The
cells are called myocytes, cardiomyocytes, or cardiocytes.
• Smooth Muscle
Contractile proteins are not arranged in the same way as in other muscle types,
and there are no striations. The cells are called myocytes and they are short and fusiform.
Smooth muscle is involuntary.

Functions of Muscle
1. Movement is an obvious function of muscle. Skeletal muscle underlies our
ability to transport ourselves by walking bipedally and to perform actions that
range from rolling our eyeballs to picking up a pencil. Communication depends
on facial expressions, speech, and writing (or typing on a computer). All these
actions are possible because of movement of skeletal muscles. Contraction of
smooth and cardiac muscle underlies movement of organs that support digestion
and circulation.
2. A second function is that skeletal muscles maintain stability by resisting the pull
of gravity so we do not fall over, and by holding bones in place.
3. Control of body openings and passages is a role of muscle. Sphincter muscles
in the eyelids, pupil, and mouth control admission of light, food, and drink. Other
sphincters control elimination of waste from the urethra and anus, and the
movement of bile, food, and other materials through tubes in the body.
4. As much as 85% of body heat is produced by skeletal muscle.
5. Muscle plays a role in glycemic control, regulation of blood sugar. A lower
skeletal muscle mass is associated with increased risk of type 2 diabetes mellitus.

Universal Properties of Muscle

Muscle tissues share the following characteristics.
• Excitability
When stimulated by chemical signals, mechanical stretch, or other stimuli, muscle
cells respond with electrical changes across the membrane.
• Conductivity

64
 Electrical excitation is passed along the length of the plasma membrane. The
passage of the electrical signal initiates events that lead to contraction.
• Contractility
Muscle fibers have the ability to shorten when stimulated. This unique quality
means that they can pull on bones or other tissues to create movement.
• Extensibility
Muscle fibers must be able to stretch back to relaxed length between each
contraction.
• Elasticity
When a muscle cell is stretched and then tension is released, it recoils to the
original length.

General Anatomy of Muscles

A skeletal muscle is an organ and consists of more than skeletal muscle tissue. It
also contains connective tissue, nerve tissue, and blood vessels.
Connective Tissues and Fascicles
• Endomysium
The endomysium is a thin sleeve of areolar connective tissue that surrounds each
muscle fiber.
• Perimysium
The perimysium wraps muscle fibers together in bundles called fascicles.
• Epimysium
The epimysium surrounds the entire muscle and extends beyond the muscle as a
tendon that connects the muscle to bone.
• Fascia
Sheets of connective tissue separate neighboring muscles.
1. Fascicles and Muscle Shape
Strength of a muscle and direction of pull are determined partly by orientation of
fascicles. Differences in fascicle orientation form the basis for classification of
muscles into five types.
2. Fusiform Muscles
Fusiform muscles are thick in the middle and tapered at each end. Examples include
biceps brachii and gastrocnemius.
3. Parallel Muscles
These are long, strap-like muscles of uniform width and parallel fascicles. An
example is sartorius.
4. Convergent Muscles
Fan-shaped muscles begin at a broad origin and converge on a narrower insertion.
Examples are pectoralis major.
5. Pennate Muscles
Pennate muscles are feather-shaped. Fascicles insert obliquely on a tendon that runs
the length of the muscle like the shaft of a feather. An example is deltoid.
6. Circular Muscles (Sphincters)
Sphincters form rings around body openings. An example is orbicularis oculi.
Muscle Attachments

65
 Muscle attachments are either direct or indirect. A direct (fleshy) attachment
occurs when collagenous fibers of the epimysium are continuous with the periosteum.
The red muscle tissue seems to emerge directly from the bone. In an indirect attachment,
a tendon emerges from the connective tissue of the muscle and merges into the
periosteum. Other attachments include an aponeurosis, a place where a muscle is
connected to a broad, sheetlike tendon. Muscles of facial expression attach to collagen
fibers of the dermis.
Most skeletal muscles span at least one joint, attaching to different bones at the
origin (relatively stationary end) and insertion (more mobile end). The traditional
definition of a muscle’s origin and insertion does not perfectly correspond to function
because the more stationary end of the muscle may actually be the more mobile end
during certain actions. Still, origins and insertions are useful tools for students when
memorizing muscle attachments.
Functional Groups of Muscles
Skeletal muscles seldom work independently. Rather, they function in groups
whose combined actions produce coordinated movement of a joint. Muscles can be
classified into at least four categories according to their actions. Of course, they may act
a certain way during one action and in another way during different actions at the same
joint.
1. Prime Mover (Agonist)
The prime mover is the muscle that produces the most force during a particular
action. For example, the brachialis is the prime mover during flexion of the elbow.
2. Synergist
A synergist is a muscle that aids the prime mover. Synergists may perform the
same action as the prime mover or may stabilize the joint or modify the direction of
movement so that the action of the prime mover is more coordinated and specific.
3. Antagonist
An antagonist is a muscle that opposes the prime mover. For example, triceps
brachii opposes the movement of biceps brachii.
4. Fixator
A fixator is a muscle that prevents a bone from moving. For example, during
contraction the biceps brachii would pull the scapula laterally if there were not fixator
muscles that attach it to the vertebral column.
Intrinsic and Extrinsic Muscles
Intrinsic muscles have their origin and insertion within a region such as the hand
or foot. Extrinsic muscles have their origin elsewhere but act upon the specific region.
For example, flexion of the fingers depends on both long, extrinsic forearm flexors, and
local, intrinsic hand muscles.
Muscles, Bones, and Levers
Bones act as levers on which muscles exert force. A lever rotates around a fixed
point, the fulcrum. Rotation occurs when effort applied to one point on the lever
overcomes a resistance (load) located at some other point. The effort arm is the part of a
lever from the fulcrum to the point of effort. The resistance arm is the part of a lever
from the fulcrum to the point of resistance. In the body, the bone acts as a lever, the joint
is a fulcrum, and the effort is generated by muscle.

66
 The function of a lever is to either produce a gain in speed and distance or force
of motion. A single lever cannot increase both speed and distance on one hand, and force
on the other. There is a trade-off between speed and force relating to the mechanical
advantage of the lever. Mechanical advantage is equal to the length of the effort arm
divided by the length of the resistance arm. If the mechanical advantage is greater than
1.0 the lever produces more force but relatively less speed and distance.
There are three classes of levers. They differ with respect to which component,
the fulcrum (F), the effort (E), or resistance (R) is in the middle.
• First-class Lever
In a first-class lever, the fulcrum (F) is in the middle. (EFR is the pattern.) The
classic mechanical example is a see-saw. An anatomical example is the atlanto-occipital
joint, where the muscles of the neck pull down on the occipital bone (E), the joint in the
middle is the fulcrum (F), and the weight of the head is the resistance (R).
• Second-class Lever
Resistance is in the middle. (ERF is the pattern.) A mechanical example is the
wheelbarrow. Lifting handles (E) makes it pivot on the wheel (F) and lift the load (R) in
the middle. An anatomical example is opening the mandible. Effort (E) is exerted by the
digastric muscle to pull down the mouth, resistance (R) is given by the temporalis
muscle, and the TMJ is the fulcrum (F).
• Third-class Lever
Effort is applied between the fulcrum and resistance. (FER is the pattern.) A
mechanical example is a baseball bat. A right-handed player puts the left hand at the
knob which acts as a fulcrum (F), the right hand produces effort (E). Resistance is
provided by the ball (R). An anatomical example is the elbow joint. The fulcrum (F) is
the joint, the biceps tendon provides the effort (E), and the resistance (R) or load is
whatever is in the hand or simply the combined weight of the forearm and hand.

Microscopic Anatomy of Skeletal Muscle

Ultrastructure of Muscle Fibers
The arrangement of contractile proteins within a muscle fiber underlies the ability
of muscle to contract. Muscle fibers have several unique characteristics that contribute to
their function.
Multiple Nuclei
Muscle fibers are unusual in that they have multiple nuclei pressed against the
plasma membrane. During embryological development, stem cells called myoblasts fuse
to produce each fiber, resulting in multinucleate mature cells.
Sarcolemma
The plasma membrane of muscle fibers, the sarcolemma, has infoldings called
transverse (T) tubules that penetrate the fiber. When the cell is stimulated, the electrical
current is carried to the interior of the cell.
Sarcoplasm
The sarcoplasm (cytoplasm) is full of long protein bundles called myofibrils.
Most other organelles are crowded between them. The sarcoplasm contains an
abundance of glycogen that provides stores of energy, and myoglobin that binds oxygen.
Myofilaments

67
 The myofibrils are composed of myofilaments, string-like proteins. The structure
and interaction between myofilaments provide the key to muscle contraction. There are
three kinds of myofilaments: thick filaments, thin filaments, and elastic filaments.
1. Thick Filaments
Thick filaments are composed of the protein myosin. In a single thick filament,
numerous myosin heads stick outward in a spiral array. On either end, the heads
angle in different directions, and the middle is bare.
2. Thin Filaments
Thin filaments are primarily composed of two intertwined strands of fibrous (F)
actin. A string of subunits is called globular (G) actin. Each G actin has an active
site that can bind to a myosin head. Thin filaments also have tropomyosin, a
protein that blocks the active binding site when the muscle is relaxed so that the
myosin head cannot bind. Troponin is bound to tropomyosin and can bind
calcium.
3. Elastic Filaments
Elastic filaments are made of the protein titin (connectin) and anchor thick
filaments to the Z disc. The elastic filaments help keep the thick and thin
filaments aligned, resist overstretching, and help the cell recoil to resting length
after it has been stretched.
Striations and Sarcomeres
In skeletal and cardiac muscle myosin and actin are organized in a precise pattern
that accounts for striations seen under the microscope. Striations are produced because
dark A bands alternate with lighter I bands. The A band is an area where there are thick
filaments and where thick and thin filaments overlap. The H band contains only thick
filaments, which originate at the M line. The I band is bisected by the Z disc (or Z line).
The Z disc anchors thin filaments and elastic filaments. Each segment of a myofibril
from one Z disc to the next is a sarcomere. The sarcomere is the functional contractile
unit of a muscle fiber.
The Nerve-Muscle Relationship
Skeletal muscle only contracts if it is stimulated by a nerve. If nerve connections
are severed or poisoned, muscle is paralyzed.
Motor Neurons
Skeletal muscle fibers are innervated by somatic motor neurons, whose cell
bodies lie in the spinal cord or brain. Axons, somatic motor fibers, lead to skeletal
muscles. At the distal end, the axon branches to numerous individual muscle fibers.
The Neuromuscular Junction
The functional connection between a nerve fiber and a target cell is a synapse.
The neuromuscular junction is the synapse between a nerve fiber and a muscle fiber.
Each branch of a motor fiber ends at a synaptic knob, nestled in a depression on the
sarcolemma called the motor end plate. The two cells are separated by a tiny gap, the
synaptic cleft.
The synaptic knob contains synaptic vesicles that are filled with acetylcholine
(ACh). When a signal reaches the synaptic knob, some of the vesicles release ACh,
which diffuses across the synaptic cleft and binds to ACh receptors on the sarcolemma.
The receptors initiate events that lead to contraction.
Motor Unit

68
 When a nerve signal approaches the end of the axon, it spreads out over all of its
terminal branches and stimulates all the muscle fibers supplied by them. These motor
fibers contract in unison, behaving as a functional unit. One motor neuron plus the fibers
it supplies is a motor unit. Motor fibers of a single motor unit are dispersed throughout
an entire muscle, an arrangement that causes a weak contraction over a wide area. It is
advantageous to have multiple motor units in a muscle because it helps to prevent fatigue.
The muscle as a whole can sustain long-term contraction.
Smaller motor units are found in areas where we need fine motor control. For
example, muscles that cause subtle movements of the eyeball have only 3 to 6 muscle
fibers per nerve fiber. In contrast, the gastrocnemius has about 1000 muscle fibers per
nerve fiber. Large motor units are associated with strength.
The Blood Supply
The muscular system as a whole receives about 1.25 liters of blood per minute at
rest, or about a quarter of the blood pumped by the heart. During exercise, however, the
amount of blood that goes to muscles increases to about 75 percent of cardiac output,
because working muscle has a great demand for glucose and oxygen. Ultimately,
capillaries pierce the endomysium of every muscle fiber in the body.

Relating Structure to Function

Contraction and Relaxation
Contraction and relaxation occurs in four phases.
1. Excitation
Acetylcholine from a motor neuron diffuses across the synaptic cleft and binds to
receptors on the sarcolemma. Receptors are gated sodium/potassium channels that open
as long as acetylcholine is bound to them. The flow of ions through the sodium and
potassium gated channels leads to a change in voltage across the sarcolemma. This sets
off a wave of electrical excitation that spreads along the sarcolemma, down the T tubules,
and to the interior of the cell.
2. Excitation-contraction Coupling
The electrical events are linked to contraction. Electrical signals passing down
the T tubules indirectly open calcium channels in the sarcoplasmic reticulum (SR). The
SR releases calcium to the cytosol. Calcium binds to the troponin on the thin filaments,
which causes the tropomyosin to shift position so that it no longer blocks the active
binding sites for the myosin heads.
3. Contraction
The myosin head, activated by ATP, swings forward to the “cocked” position and
binds to actin. The myosin head then swivels and pulls the actin a short distance, the
“power stroke”. The myosin head releases, binds a new ATP, recocks, and repeats the
process. The myosin heads pull the thin filament smoothly over the thick filament. This
is called the sliding filament theory of contraction. The myofilaments do not shorten.
Rather, they slide over each other. The result is that the thin filaments pull on the Z discs
and the sarcomere shortens. Ultimately, the muscle fiber shortens.
4. Relaxation
Relaxation begins when nerve signals stop arriving at the neuromuscular junction
and the nerve cell stops releasing acetylcholine. Acetylcholinesterase breaks down
remaining ACh. With no electrical stimulation, the SR begins to pump calcium back into

69
the cisternae. Tropomyosin moves back to the resting position and blocks the active
binding sites for myosin. The myosin-actin bridges no longer form and the muscle
relaxes.
Muscle Growth and Atrophy
When people exercise their muscles increase in size. In contrast, after long
periods of inactivity, muscles atrophy. Skeletal muscle fibers are incapable of mitosis.
In response to exercise, muscles grow through hypertrophy, the growth of existing cells.
Exercise leads to an increase in protein myofibrils within fibers. In contrast, muscle
atrophies if it is not used or if there is damage to nerves that supply muscles.
Physiological Classes of Muscle Fibers
Slow oxidative (SO) fibers respond slowly but resist fatigue. They are also called
slow-twitch, red, or Type 1 fibers. These fibers have abundant mitochondria, myoglobin,
and capillaries that give them a deep red color. They produce ATP mostly through
aerobic respiration, which does not generate lactic acid. Lactic acid contributes to
fatigue, so these fibers do not fatigue easily.
Fast glycolytic (FG) fibers are also called fast-twitch, white, or Type II fibers.
They respond quickly but fatigue more quickly than SO fibers because they rely on
anaerobic respiration that produces lactic acid. They have fewer mitochondria and
capillaries and less myoglobin so are relatively pale. They are adapted for quick response
but not endurance.
All muscles are composed of both SO and FG fibers but the proportion of fibers
differs according to the function of the muscle. In addition, individuals may vary in the
proportion of fibers in muscles, as has been demonstrated in elite athletes.

Cardiac and Smooth Muscle

Cardiac Muscle
Cardiac muscle makes up most of the heart. It is striated like skeletal muscle.
Cardiomyocytes, unlike skeletal muscle fibers, have a single nucleus, they are short,
branching cells, and they have specialized junctions called intercalated discs.
Intercalated discs contain electrical gap junctions that allow cells to communicate and act
as a group in a coordinated fashion.
Cardiac muscle fibers are rich in glycogen and myoglobin. Mitochondria are
large and fill about 25 percent of the cell, in contrast to skeletal muscle fibers where
mitochondria only make up about 2 percent. Cardiac muscle is adapted to aerobic
respiration, and is resistant to fatigue, but is vulnerable to disruption of oxygen supply.
Cardiac myocytes are incapable of mitosis so repair of damaged tissue is entirely by
fibrosis (scarring).
Like skeletal muscle cells, calcium is key for muscle contraction. Unlike skeletal
muscle fibers, calcium in cardiocytes comes not only from the SR but also from the
extracellular fluid.
The nervous system does not stimulate contraction of cardiac muscle. Rather,
cardiocytes contract rhythmically on their own. The sinoatrial node normally sets the
beat. The autonomic nervous system moderates heart rate and strength.
Smooth Muscle
Smooth muscle myocytes are fusiform and have a single nucleus. Thick and thin
filaments are not aligned and there are no visible striations or sarcomeres. Thin filaments

70
attach to dense bodies, masses of protein scattered through the sarcoplasm, and on the
inner face of the sarcolemma. There are no T tubules.
Like skeletal and cardiac muscle, calcium is required for contraction but most of it
comes from the extracellular environment. During relaxation, calcium is pumped out of
the cell. The autonomic nervous system supplies nervous stimulus for contraction.
Smooth muscle, unlike either skeletal or cardiac muscle, is capable of mitosis;
therefore, injured smooth muscle regenerates.
Multi-unit smooth muscle is found in the largest arteries and pulmonary air
passages. Terminal branches of autonomic nerve fibers synapse with individual
myocytes to form a motor unit. Each motor unit contracts independently of others.
Single-unit smooth muscle is more widespread, occurring in most blood vessels,
digestive, respiratory, and reproductive tracts. Here, the nerve fibers pass through the
tissue and exhibit swellings called varicosities where they release neurotransmitters. In
this way, multiple muscle cells are nonselectively stimulated in the vicinity of the
varicosity. Muscle cells are also electrically coupled at gap junctions so that they
stimulate each other. The result is that a large number of cells contract as a single unit.
Smooth muscle contracts and relaxes slowly in response to nerve stimulation,
chemicals, and stretch. Metabolism is mostly aerobic. Smooth muscle is resistant to
fatigue because of its relatively low energy requirements compared to skeletal and
cardiac muscle.

Developmental and Clinical Perspectives

Embryonic Development of Muscle
Most muscle arises from embryonic mesoderm. The mesoderm of the trunk
forms systematically arranged blocks called somites. Somites divide into dermatomes,
sclerotomes, and myotomes. Myotomes form the muscles of the trunk. In week 4,
mesodermal cells of the myotomes become myoblasts and begin to divide. Some
myoblasts fuse to form multinucleate myotubes. By week 9, muscle groups are present
and in position relative to the skeleton, and muscle fibers begin to contract in response to
nervous stimulation. By week 17, fetal muscle contractions are strong enough to be felt
by the mother.
Some myoblasts persist after birth as satellite cells. They contribute to new
muscle growth in childhood and may regenerate a limited amount of damaged skeletal
muscle in adults.
Cardiac muscle develops in association with the heart tube. Mesenchymal cells
near the heart tube differentiate into myoblasts that divide mitotically, similar to skeletal
muscle. The cells do not fuse but join together and develop intercalated discs. At week
3, the heart begins beating. Mitosis in cardiac myocytes continues after birth until about
age 9.
Smooth muscle develops from myoblasts associated with the embryonic gut,
blood vessels, and other organs. The myoblasts do not fuse, but in single-unit smooth
muscle cells become interconnected through gap junctions.
The Aging Muscular System
As people age, muscle performance declines. Muscle mass and strength peak in
the 20’s. By age 80, most people have half as much strength and endurance as they did
when they were young. Underlying reasons are that aged muscle fibers have fewer

71
myofibrils and are weaker and smaller. Sarcomeres become increasingly disorganized,
the mitochondria shrink and have fewer oxidative enzymes, and the muscle has less ATP,
less glycogen, and less myoglobin. It fatigues quickly. There is reduced blood
circulation. Changes in the nervous system contribute to decreased endurance and slower
response. In addition, overall body composition changes as lean body mass declines and
fat mass increases.
Diseases of the Muscular System
Muscle diseases are called myopathies and include muscular dystrophy and
myasthenia gravis.
Duchenne muscular dystrophy is the most common of a group of hereditary
diseases that lead to degeneration of skeletal muscles, loss of strength, and death. It
usually manifests in males because it is sex-linked. A defective gene for the protein
dystrophin leads to a weak sarcolemma and death of muscle fibers. Children appear
normal until they begin to walk but eventually muscles atrophy and patients are usually
confined to a wheelchair by adolescence. Afflicted individuals rarely live beyond age 20.
Myasthenia gravis is an autoimmune disease that is most prevalent in women
between the ages of 20 and 40. Antibodies attack the neuromuscular junctions and
trigger destruction of acetylcholine receptors. The muscle fibers become less and less
sensitive to acetylcholine. Symptoms include drooping eyelids, difficulty swallowing,
weakness in the limbs, and poor physical endurance. Some people die quickly due to
respiratory failure, while others live normal life spans.

Key Concepts
• Skeletal Muscle
Voluntary, striated muscle that usually attaches to bones.
• Muscle Fibers
Skeletal muscle cells. Also called myofibers.
• Striations
Alternating dark and light bands that can be seen with aid of a microscope.
• Cardiac Muscle
Muscle that makes up the wall of the heart. It is involuntary and striated.
• Cardiocytes
Cardiac muscle cells. Also called myocytes.
• Smooth Muscle
Involuntary muscle associated with blood vessels and organs. Cells are short and
fusiform. Cells are also called myocytes.
• Excitability
Ability to respond rapidly to stimuli by changing electrical charges across the plasma
membrane. A key property of muscle cells.
• Contractility
Ability to shorten substantially when stimulated.
• Endomysium
Connective tissue that surrounds each muscle fiber.
• Perimysium
Connective tissue that surrounds fascicles, bundles of muscle fibers.
• Epimysium

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 Connective tissue that surrounds entire muscle. It extends beyond muscle as the
tendon, connecting it to the periosteum.
• Origin
Muscle attachment at the relatively stationary end.
• Insertion
Muscle attachment at the more mobile end.
• Prime Mover (Agonist)
The muscle that produces most of the force during a particular action at a joint.
• Synergist
A muscle that aids the prime mover.
• Antagonist
Muscle that opposes the prime mover.
• Fixator
Muscle that prevents a bone from moving.
• Intrinsic and Extrinsic Muscles
Intrinsic muscles are completely contained within a certain region. Extrinsic muscles
act on a specific region but have origins elsewhere.
• Lever
Elongated, rigid object that rotates around a fixed fulcrum. In the body, long bones
are levers.
• First-class Lever
Has the fulcrum in the middle. Effort applied by muscles on one side of the fulcrum
overcomes resistance (load) on the opposite side. Example is the atlanto-occipital
joint.
• Second-class Lever
Has resistance in middle. Force applied to one side overcomes resistance in the
middle and causes a joint at the opposite end to pivot. Example is the mandible where
force is supplied by the digastric, resistance is tension of the temporalis, and the
fulcrum is the TMJ.
• Third-class Lever
Has effort in the middle. Fulcrum is at one end, effort supplied by muscles in the
middle, and resistance is at end opposite the fulcrum. Example is the elbow joint.
• Sarcolemma
Plamsa membrane of muscle cells.
• Transverse (T) Tubules
Infoldings of the sarcolemma that carry an electrical current from the surface of the
cell to the interior.
• Sarcoplasm
Cytoplasm of muscle cells.
• Myofibrils
Organelles in sarcoplasm that consist of long protein bundles.
• Sarcoplasmic Reticulum (SR)
Smooth endoplasmic reticulum of a muscle fiber that has dilated sacs called terminal
cisternae.
• Myofilaments

73
 Protein microfilaments that are found in myofibrils.
• Thick Filaments
Compared of myosin. Interact with thin filaments to cause muscle contraction.
• Thin Filaments
Composed mainly of actin. Also contain troponin and tropomyosin.
• Sarcomere
Segment of a myofibril that extends from one Z disc to the next. The functional
contractile unit of the muscle fiber.
• Somatic Motor Neurons
Innervate skeletal muscle cells. Axons that lead to muscle cells are called somatic
motor fibers.
• Neuromuscular Junction
A synapse between a nerve fiber and a muscle cell.
• Acetylcholine (ACh)
Neurotransmitter released by synaptic vesicles at the end of the somatic motor fibers.
Binds to ACh receptors on sarcolemma and initiates events that lead to contraction.
• Motor Unit
One nerve fiber and all the muscle fibers it innervates.
• Excitation-contraction Coupling
The connection between excitation (chemical and electrical events) and contraction
(mechanical movement of the fiber).
• Slow Oxidative (SO) Fibers
Slow twitch fibers are well adapted to aerobic respiration due to numerous
mitochondria and resist fatigue. Have a long twitch.
• Fast Glycolytic (FG) Fibers
Fast twitch fibers rely on anaerobic respiration and fatigue relatively easily. They
respond quickly and have fast twitches.
• Myoblasts
Cells that give rise to muscle fibers.
• Duchenne Muscular Dystrophy
Incurable, sex-linked hereditary disorder in which skeletal muscles degenerate, lose
strength, and atrophy. Due to a defective gene for the protein dystrophin.
• Myasthenia Gravis
Autoimmune disorder where antibodies attack the neuromuscular junction and trigger
the destruction of ACh receptors.

Learning Strategies/Teaching Tips

• Demonstrate the relevant images and animations from Anatomy & Physiology
Revealed in lectures and labs.
• Terminology associated with the microscopic anatomy of muscle fibers can be
overwhelming. Encourage students to draw pictures of a skeletal muscle fiber, then a
sarcomere, and label the components. This simple exercise seems to help many
students understand the relationships among the structures.

74
• Ask students to list the steps of muscle contraction from the release of ACh from the
motor neuron to the shortening of the sarcomere. Making a flow-chart outlining the
steps effectively organizes and reinforces the material.
• To help students understand lever classes, have them identify the point of resistance,
effort, and fulcrum using the letters “R”, “E”, and “F” for different joints. (For
example, a first-class lever is “RFE”.) Using their knowledge, they should be able to
classify the talocrural joint.
• Brainstorm in lecture or lab about the similarities and differences among skeletal
muscle, cardiac muscle and smooth muscle. Have students make charts the compare
and contrast the three types.

Additional Reading
Vogel, Steven. 2002. Prime Mover: A Natural History of Muscle. W. W. Norton &
Company.

75
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drought settled over the hot, depleted Nation. By August, many of
the captured Cherokees had succumbed to sickness or even death.
Removal itself began during the autumn. A few early contingents had
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weather changed. Winter stalked the doomed procession with the
tenacity of a bloodhound. By the time the Cherokees crossed the
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The tragedy would be recorded in history as the “Trail of Tears.”
Along the route, old Whitepath died. The wife of John Ross gave her
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knew.” But disaster was not the final conqueror. For out of that
cruelty came sacrifice; out of that death came rebirth.
The improbable
source of that
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warriors to action. In Smithsonian Institution
the quick, sudden
tangle that followed, Tsiskwa-kaluya, or Bird Chopper,
at least one soldier was son of Yonahguskah, the
was killed. Tsali and famous Cherokee chief and
his small band fled spokesman who stayed with the
across the river high small group in the Great Smoky
into the Great Mountains.
Smoky Mountains.
They hid in a massive rock shelter at the head of Deep Creek.
Located on top of a steep cliff, the actual camping place lay in the
midst of extensive, thick laurel and hemlock “roughs.” Several
hundred other Cherokees escaped from the soldiers or the
stockades and found similar hiding places on the rugged, overgrown
sides of the Smokies. Most of them lasted out the winter, subsisting
at a near-starvation level on roots, herbs, nuts, and small game.
Confronted with such determination and the likelihood of a
prolonged, wearisome mission of search and arrest in the rugged
mountains, General Scott offered a compromise. If Tsali and his
small party would come down and give themselves up for
punishment, the rest of the Cherokee fugitives would be allowed to
stay in the mountains until a solution could be reached by all sides.
Scott sent W. H. Thomas, a white man who had grown up with the
Cherokees, into the Smokies to present the terms. Thomas found
Tsali, who silently listened and decided on his own accord to
accompany Thomas out of the mountains. Early in the year of 1839,
Tsali and his brother and his eldest son were shot by a firing squad.
The youngest son, Wasituna (for Washington) was left to take word
of the deaths back to the Cherokees who remained in their hills.
They had held onto their homeland in the Great Smoky Mountains.
By nothing more than the thin grip of desperate determination, they
had held on, and they would remain. Reinforced by General Scott’s
promise, scattered friends in the East, and Thomas’ political
negotiations with Washington and North Carolina, the Cherokee
remnant soon became the Eastern Band. Their homeland would now
be known as the Qualla Reservation. So the Cherokees East, along
with the white pioneers of the Great Smokies, turned together to
brace the mountainous challenge of the 19th century.
 Aaron Swaniger was an individualist who occasionally
stayed in Cades Cove. To some “mountaineers” he
was a “hillbilly.”
Edouard E. Exline
 From Pioneer to Mountaineer
While events of the early 19th century in the surrounding southland
and the nation were moving inexorably toward conflict on bloody
battlefields to decide issues which could not or would not be
resolved in the political arena, people in the Great Smokies were
pursuing their struggle to survive and adapt to their stern and
splendid surroundings.
The early explorers, the long hunters, the initial homesteaders, the
trailblazers and the groundbreakers—these had forever set a human
seal upon the wilderness. Now it was the time of pioneer becoming
mountaineer. Henceforth, as new settlers or curious travelers or
specialized seekers in a dozen fields made their way into the
mountains, they would find someone already there to welcome them.
That “someone” was becoming known by terms which might
alternately serve as a source of description, derision, or definition.
Highlander. Hillbilly. Mountaineer. The least offensive word was
“highlander,” with its overtones of the misty Scottish landscape and
fierce clan loyalties from which many of the Smokies’ family lines
had recently descended. “Mountaineer” varied. Used to denote the
proud individualism that characterized many of the stalwart men and
women whose roots held deep and fast in this isolated place,
“mountaineer” was a strong, acceptable name. But turned into a
catchword for some picturesque, inadequate character who divided
his time between the homemade dulcimer and the home-run
distillery, “mountaineer” was suspect. “Hillbilly” came to verge on
insult, as it conjured up cartoons of lanky, sub-human creatures who
were quick to feud, slow to work, and often indifferent to the
“progress” by which helpful visitors would like to transform mountain
lives and attitudes.
Of course, the trouble with any single word that tried to summarize
these complex and distinctive lives was its limited ability to convey
more than a stereotype or a single facet. Yet the 19th and early 20th
centuries saw the rise and wide adoption of such terms, with an
accompanying unease—sometimes outrage—on the part of those
described. This tension has persisted into the present day, for
Southern Appalachian natives often have felt they have been
misunderstood, or exploited, by the curious outlanders.
 Cherokee veterans of Thomas’ Legion attending a
Confederate reunion in the early 1900s in New Orleans
include (front from left) Young Deer, unidentified man,
Pheasant, Chief David Reed, (back from left) Dickey
Driver, Lt. Col. W. W. Stringfield, Lt. Suatie Owl, and
Jim Keg. Stringfield was a white officer in the legion
which participated with varying degrees of success in
several skirmishes in the Smokies and, perhaps more
importantly, which helped build the Oconaluftee
Turnpike across the mountains.
National Park Service
The visitors indeed were curious—curious about mountain people
but also about topography, altitudes, plants, wildlife, and the rich
variety of natural resources abounding throughout these hills.
Naturalists and botanists followed the lead of Frenchman André
Michaux and Philadelphian William Bartram, who had come
collecting plants in the Southern mountains during the previous
century. It was Michaux who had told mountain herb-gatherers about
ginseng’s commercial value, and Bartram who had discovered and
described the showy flame azalea brightening the spring woods.
Among 19th-century arrivals, S. B. Buckley wrote the earliest
comprehensive botanical report of the Great Smokies. He marveled
at that scenery, “surpassing anything we remember to have seen
among the White Mountains of New Hampshire,” and at the variety
of flora. “Here,” he wrote in the mid-1800s, “is a strange admixture of
Northern and Southern species of plants, while there are quite a
number which have been found in no other section of the world.”
Later naturalists would share his enthusiasm and enlarge on his
studies.
Journalists came. One was a reporter named Charles Lanman,
secretary to Daniel Webster, who rode through the hills in 1848 and
wrote a book called Letters from the Alleghany Mountains. Through
his descriptive adventures readers had a glimpse into a region more
remote to their experience than many foreign countries. If the
Smokies were described by him as one large upthrust, perhaps that
was because he saw the range through a purple haze. He wrote at
one point:
“This mountain is the loftiest of a large brotherhood which lies
crowded together between North Carolina and Tennessee. Its height
cannot be less than five thousand feet above the level of the sea ...
and all I can say of its panorama is that I can conceive of nothing
more grand and imposing.”
Lanman was only the first of many writers who would come seeking
high adventure and good copy, but his lack of exactitude about the
physical features of the mountains was soon to be remedied by
another group of visitors. Some scientists could not be content with
hunters’ yarns and the poetic prose of journalists; they wanted
precise facts and figures by which both native and stranger could
better appreciate the landscape.
One of these was Thomas Lanier Clingman, whose career included
being a U.S. senator and a Confederate general as well as a
scientist. A contemporary historian described him as being arrogant,
aggressive, with “more than common ability” but limited scientific
knowledge, whose chief service lay in arousing public curiosity in the
mountains, mineralogy, and geography. He became involved in a
scholarly feud with Dr. Elisha Mitchell, a transplanted Connecticut
professor at the University of North Carolina, over which peak
constituted the highest point east of the Mississippi River. While they
were trying to settle the question, Mitchell was killed in an accidental
fall on the slopes of the North Carolina pinnacle which later was
given his name. Clingman’s name came to grace the mountain he
had explored and measured: 2,025-meter (6,643-foot)-high
Clingmans Dome on the crest of the Great Smokies, only 13 meters
(43 feet) lower than the lofty Mt. Mitchell.
The most fascinated and impressive visitor during these years of the
mid-19th century came from another mountain terrain. Arnold Guyot,
remembered today by the peak at the eastern end of the Great
Smokies which bears his name, was born in Switzerland in 1807. His
studies in physical geography had won him distinction throughout
Europe before he came to America and accepted a chair at
Princeton University in 1854. Paul Fink, a historian of the Great
Smoky Mountains, has said that although forerunners of Guyot
glimpsed segments of the Smokies and described certain details,
“it remained for this man of foreign birth to penetrate these
mountains, spend months among them, measure their heights for
the first time, and have drawn under his own direction the first map
we have showing the range in detail.”
Clingman secured for his friend Guyot a local guide named Robert
Collins. The mountain man and the professor struggled through the
roughest laurel “hells” and up the steepest slopes, measuring,
calibrating, and finally naming many of the unknown heights. Guyot’s
journals combined precision and poetry, and they related the
awesome Smokies to the human scene in ways that had not been
previously possible. From that point on, natives and visitors alike
could both know and appreciate more of this green homeland. But,
as Paul Fink has pointed out, “With Guyot’s labors the early
explorations of the Smokies ceased.”
Why? What happened to cut off so abruptly the increasing flow of
visitors to this virgin country? The happening was war.
The Great Smokies country, with its upland farms and small home
crafts, was not in the mainstream of the decisive struggle between a
plantation South and an industrial North. Nor was it in the
mainstream of the violent action that convulsed its surrounding
region. There had been slaves in some of the more prosperous
mountain households, but few citizens in the Great Smokies area
would have waged war either to defend or abolish the peculiar
institution.
What some did resist was being conscripted by either side.
“Scouting” became a well-used word defining a new experience in
the Smokies. It applied to anyone hiding out in the hills to escape
going into the Confederate or the Union army. Secretly supplied with
food and clothes by their families and sympathetic friends, such
“scouts” could hold out for years against the searches of outlander
officials. Sometimes they did in fact become scouts, guiding escaped
captives from Andersonville and other Confederate prisons through
the mountains toward northern territory, and those fleeing from
Yankee prisons toward their southern homes.
Many of the mountain people, of course, followed the example of
their neighbors throughout the region and put on the formal uniform
of blue or gray. There were sharp divisions within counties, towns,
and families in the choice between state and nation. Perhaps no
single section of the United States was as bitterly torn in its
allegiance.
Tennessee and North Carolina had long held strong Union
sentiments; but when Lincoln called for troops in the aftermath of the
firing on Fort Sumter, the two states officially rallied to the
Confederate cause. North Carolinians, who had been notably
reluctant to leave the Union and who bristled at the injustices of “a
rich man’s war and a poor man’s fight,” nonetheless sent more men
to the Confederacy than any other state. Many of these were
western North Carolinians, following the leadership of their own
Zebulon (Zeb) Baird Vance, born in Buncombe County and
occupying the governor’s chair in Raleigh during the war. Yet the fact
that adjoining East Tennessee was overwhelmingly Union—and sent
more men into the Federal forces than some of the New England
states—affected the North Carolinians as well. With the two states’
actual secession from the Union, numerous mountain pockets in
effect seceded from their states and chose to remain loyal to the
Union. Thus the little rebellion inside the larger revolt compounded
the agonizing conflict of war and made every cove and community
and hearthside a potential battleground.
And no matter which army the men marched with, their
characteristics remained surprisingly intact. The historian of one
North Carolina Confederate regiment described some of the soldiers
from Haywood County:
“These mountain men had always been accustomed to
independence of thought and freedom of action, and having elected
for their company officers their neighbors and companions, they had
no idea of surrendering more of their personal liberty than should be
necessary to make them effective soldiers. Obedient while on duty
 and independent
while off duty, this
spirit to a marked
degree they
retained to the close
of the war.”
The experience of
Radford Gatlin
concentrated in a
single episode both
the sharp divisions
and the ironies of
war in the
mountains.
Gatlinburg, now a
commercial and
flourishing tourist
mecca at the edge
of the park, bears
the name of a man
who was driven out
of that town
because of his
unpopular stand
during the war. The
sturdily built,
enterprising, and
somewhat arrogant
Charles S. Grossman Gatlin was not a
man to conceal his
Mountain women and girls had to beliefs. With his wife
be proficient at making many and a slave woman
things, for there weren’t many—if he had come from
any—stores nearby. Over the years, North Carolina by
Hazel Bell and many another way of Jefferson
woman spent hours and hours County, Tennessee,
churning butter. to the community
 known as White
Oak Flats and had established a successful general store and a less
successful church: the New Hampshire Baptist Gatlinites. When Dick
Reagan was appointed postmaster for a new postal service to be
established in White Oak Flats in 1860, the office was located amidst
the axes, guns, coffee, sugar, and bells of Gatlin’s store, and
Reagan renamed the post office, and therefore the town, after his
good friend the storekeeper.
But when war came and Radford Gatlin not only supported the
Confederacy but made heated speeches in its favor, the strongly
Unionist villagers turned against him. After being beaten by a band
of masked men, Gatlin abandoned his claim to thousands of
hectares that now lie within the park and departed forever from the
place that was to perpetuate his name if not his memory.
The war’s severest hardships followed in the wake of the outliers, or
the bushwhackers. These scavengers favored no cause. As the war
dragged on, they ambushed and raided, stealing meat from the
smokehouse, corn from the crib, and farm animals from barn and
pasture. Scarcity and want became commonplace throughout the
mountains. In North Carolina’s Madison County, a group of citizens
broke into a warehouse and laid claim to a valuable commodity, salt.
Economic want enflamed political emotions. In Tennessee’s Sevier
County, controversial “Parson” Brownlow, Methodist circuit-rider
turned newspaper editor turned politician, sought refuge in the
shadow of the Smokies with Unionist sympathizers when Knoxville
came under Confederate control.
A well-known army unit operated in the Smokies: Col. William
Thomas’ Confederate 69th-N.C., known as Thomas’ Legion of
Indians and Highlanders. “Little Will,” as he was affectionately called,
had become the effective spokesman in Washington and at the state
level for the eastern remnant of the Cherokee. When the Civil War
came and he chose to stay with the South, the Cherokees chose to
stay with him. For a while, they secured mineral supplies for the
Confederacy, including alum and saltpeter for gunpowder. The
Legion guarded Alum Cave in the Smokies. Under Thomas’
direction, his unit also worked on the Oconaluftee Turnpike.
 Page 80: Aunt Celia Ownby cards, or straightens, wool
fibers that have already been washed.

Edouard E. Exline
Page 81: Hettie, Martha, and Louisa Walker run cotton
through a gin built by their father, John. He made the
rollers out of hickory and the rest out of oak. Three
people were required to operate the gin: one to feed
the cotton into it and one on each end to turn each of
the rollers. The ginned cotton fell into a white oak
basket, also made by John Walker.
Charles S. Grossman
Wash day was a laborious one of lifting large buckets
of water and stirring steaming kettles of dirty clothes.

Maurice Sullivan
Over another fire, Mrs. Kate Duckett and daughter
Tennie of Coopers Creek make hard soap. Mrs.
Duckett stirs the lard with a wooden paddle as Tennie
fans the fire with a hawk wing before dipping into the
kettle with a gourd scoop. It was a five-hour process.
In December 1863, after Gen. Ambrose E. Burnside had secured
Knoxville for the Union, Col. William J. Palmer and the 15th
Pennsylvania Cavalry attacked Thomas’ camp near Gatlinburg. After
a short battle, Thomas and his troops retreated across the
mountains into North Carolina. One month later, Confederate Gen.
Robert P. Vance decided to remedy the situation in the mountains.
With 375 cavalry, 100 infantry, and artillery, he marched from
Asheville, joined Thomas and 150 Indian troops in Quallatown, and
crossed the Smokies during a bitterly cold spell. While Thomas
remained in Gatlinburg, Vance proceeded toward Newport, camped
on Cosby Creek deep in the Smokies, and was surprised there by
none other than Colonel Palmer and his 15th-Pa. In the resulting
rout, General Vance was captured along with about everything else:
men, horses, medical supplies, food, ammunition. In February,
Thomas and his Legion were engaged once more, in the Great
Smoky Mountains near the mouth of Deep Creek. The result was
another defeat, this time by the 14th Illinois Cavalry under Maj.
Francis Davidson.
Thomas and his Legion did not win mighty military victories for the
Confederacy; Governor Vance even accused Thomas’ command of
being “a favorite resort for deserters.” But it appears that this strange
little mountain force did act as a deterrent against wholesale raids in
the Smokies by Federal sympathizers, and to some extent, raids by
marauding bushwhackers. As for “Little Will” himself, mental disorder
in later years brought him his own personal civil war and its losing
battles. He died in a North Carolina hospital.
An equally well-known force in the Great Smokies during the war
was a band of Union raiders led by Col. George W. Kirk. One
contemporary called him “Kirk of Laurel,” referring to a remote
watershed in Madison County where the colonel often camped.
Kirk’s most effective march into the Smokies came near the close of
the war, in the early spring of 1865. With 400 cavalry and 200
infantry he entered the mountains through East Tennessee’s Cocke
County, via Mt. Sterling, and marched into Cataloochee. Turning
aside a Confederate company there, he went on to Waynesville,
then proceeded to Soco Valley and back across the Smokies.
Kirk raided, released Federal prisoners, skirmished with home
guards, and scattered general fear throughout the mountains. In fact,
his main achievement for his cause lay in diverting Confederate
troops and keeping them scattered on the home front rather than
mobilized on the battlefields where they were desperately needed.
Try as they might, the Confederates could not find enough of the
“silver-greys” or the “seed-corn”—as those too old and too young for
regular service were called—to totally protect their homeland from
Kirk’s men, or from renegade bushwhackers who had no cause but
plunder and pillage.
As the Civil War drew toward its final convulsion, the mountain area
engaged in a more familiar struggle for survival. Food was scarce,
soda and salt almost non-existent. Women leached lye from wood
ashes and made soda. There was no substitute for salt; when
available, it cost a precious dollar a pound. Bitter enmities divided
families, communities, and counties. Life had never been easy in the
mountains; now it was rigorously difficult. And the people in this land
of “make do or do without” learned new ways to make do. Continuing
old habits and traditions up their isolated coves and along their steep
hillsides, they created a life that was distinctive, rugged, and adapted
to its natural surroundings.
One historian, John Preston Arthur, has described the mountain
woman’s day as follows:
“Long before the pallid dawn came sifting in through chink and
window they were up and about. As there were no matches in those
days, the housewife ‘unkivered’ the coals which had been smothered
in ashes the night before to be kept ‘alive’ till morning, and with
‘kindling’ in one hand and a live coal held on the tines of a steel fork
or between iron tongs in the other, she blew and blew and blew till
the splinters caught fire. Then the fire was started and the water
brought from the spring, poured into the ‘kittle,’ and while it was
heating the chickens were fed, and cows milked, the children
dressed, the bread made, the bacon fried and then coffee was made
and breakfast was ready. That over and the dishes washed and put
away, the spinning wheel, the loom or the reel were the next to have
attention, meanwhile keeping a sharp lookout for the children,