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Chapter 08
Muscular System
HOLE’S ESSENTIALS
OF HUMAN ANATOMY
& PHYSIOLOGY

Charles J. Welsh

© 2021 McGraw Hill. All rights reserved. Authorized only for instructor use in the classroom.
No reproduction or further distribution permitted without the prior written consent of McGraw Hill.
 8.1: Introduction to the Muscular System

Muscles are organs that generate force to cause all types of

movement

Examples of muscle actions are walking, breathing, pumping

blood, and moving food in the digestive tract

The 3 types of muscle tissue in the body are skeletal, smooth, and
cardiac muscle

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 8.2: Structure of a Skeletal Muscle

Body contains over 600 skeletal muscles

Connective tissue coverings over muscles:
• Layers of dense connective tissue, called fascia, surround and
separate each muscle
• This connective tissue extends beyond the ends of the muscle,
and gives rise to tendons that are fused to the periosteum of
bones
• Sometimes muscles are connected to each other by broad
sheets of connective tissue called aponeuroses

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 Connective Tissue Coverings

Fascia blends with the epimysium, the layer of connective tissue

around each skeletal muscle
The perimysium extends inward from the epimysium; it surrounds
bundles of skeletal muscle fibers, called fascicles, within each
muscle
Each muscle cell (fiber) is covered by a connective tissue layer
called endomysium

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 A Skeletal Muscle and Its Connective Tissues

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 Skeletal Muscle Fibers

Each muscle fiber is a single, long, cylindrical muscle cell

Fibers respond to stimulation by contracting
Cell membrane of a muscle fiber is the sarcolemma
Cytoplasm of a muscle cell is the sarcoplasm; it contains many
mitochondria and nuclei
Sarcoplasm contains parallel myofibrils, which are active in muscle
contraction:
• Thick filaments in myofibrils consist of the protein myosin
• Thin filaments in myofibrils are mainly composed of the protein actin,
but also contain troponin and tropomyosin
• The organization of these filaments produces bands called striations

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 The Sarcomeres of Myofibrils

Myofibrils are made up of many units called sarcomeres, joined end-to-

end
A sarcomere extends from one Z line to the next
Striations consist of an alternating pattern of light and dark bands
I bands (light bands) are made up of actin filaments, which are anchored
to the Z lines
A bands (dark bands) are made up of overlapping thick and thin
filaments
In the center of the A band is the H zone, which consists of myosin
filaments only
The M line, in the center of the H zone, consists of proteins that hold the
myosin filaments in place

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 Structure of a Skeletal Muscle Fiber

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 The Structure of a Sarcomere

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 The Sarcoplasm of a Skeletal Muscle Fiber

Beneath the sarcolemma of a muscle fiber lies a network of

membranous channels, called the sarcoplasmic reticulum (SR),
which is the endoplasmic reticulum of a muscle cell
The SR is associated with transverse (T) tubules, invaginations of
the sarcolemma
Each T tubule lies between 2 cisternae of the sarcoplasmic
reticulum; T tubules are open to the outside of the muscle fiber
The sarcoplasmic reticulum and transverse tubules activate the
muscle contraction mechanism when the fiber is stimulated

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 The SR and T Tubules of the Sarcoplasm

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 Neuromuscular Junction 1

Skeletal muscle fibers contract only when stimulated by a motor

neuron
Each skeletal muscle fiber (cell) is functionally (not physically)
connected to the axon of a motor neuron, creating a synapse
The neuron communicates with the muscle fiber by way of
chemicals called neurotransmitters, which are released at the
synapse
Neuromuscular junction: a synapse between a motor neuron and
a muscle fiber that it regulates

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 Neuromuscular Junction 2

The cytoplasm of the distal end of the motor neuron contains numerous
mitochondria and synaptic vesicles storing neurotransmitters

The muscle fiber membrane in this area contains a specialized region

called the motor end plate, in which the sarcolemma is tightly folded

The motor end plate contains specific receptors for the neurotransmitter

When an electrical impulse reaches the end of the axon of a motor

neuron, synaptic vesicles release neurotransmitter into the synaptic
cleft, the gap between the membranes of the neuron and muscle fiber

The neurotransmitters diffuse across the cleft, bind to the motor end
plate, and stimulate the muscle fiber to contract

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 Neuromuscular Junction 3

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 8.3: Skeletal Muscle Contraction

Muscle contraction involves several events, that result in the

shortening of sarcomeres, and the pulling of the muscle against its
attachments
The pulling force is exerted by the binding of myosin molecules to
actin molecules
The shortening of a muscle fiber results from an increase in the
overlap between actin and myosin filaments, as they slide past
each other
Shortening of muscle fibers results in shortening of the entire
muscle, which then pulls on the attached body part to cause
movement

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 Role of Myosin and Actin 1

Myosin consists of two twisted strands, with globular heads

projected outward along the strands
A group of myosin molecules forms a thick filament
Actin is a globular protein arranged in twisted filaments (a double
helix), containing myosin binding sites
Troponin and tropomyosin are 2 proteins associated with the
surface of the actin molecules; together, these 3 proteins form the
thin filaments

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 Thin and Thick Filaments

Access the text alternative for slide images.

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 Role of Myosin and Actin 2

According to the sliding filament model of muscle contraction, during muscle

contraction, a myosin head attaches to a binding site on the actin filament,
forming a cross-bridge
This binding causes the head to bend, pulling on the actin filament, and moving
it toward the center of the sarcomere
The head then releases, and attaches to the next binding site on the actin,
pulling this site toward the center
As this occurs again and again, the filaments increase their overlap, and the
sarcomere shortens from both ends
When many sarcomeres shorten at the same time, the muscle fiber shortens
Energy from the conversion of ATP to ADP is provided to the cross-bridges by
the enzyme ATPase; ATP breakdown causes the heads to return to the “cocked”
position, ready to bind to another actin binding site

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 The Sliding Filament Model

1. Relaxed muscle

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 Sarcomere Shortening in Muscle Contraction

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 Stimulus for Contraction 1

Acetylcholine is the neurotransmitter for skeletal muscle fiber contraction

at the neuromuscular junctions
Acetylcholine is produced in the motor neuron, and stored in the synaptic
vesicles at the distal end of the neuron
Acetylcholine is released into the synaptic cleft in response to an impulse in
the motor neuron; it then stimulates the muscle fiber
Upon receipt of the muscle impulse, the sarcoplasmic reticulum releases its
stored calcium to the cytosol of the muscle fiber
The high concentration of calcium in the sarcoplasm interacts with the
troponin and tropomyosin molecules, which move aside, exposing the
myosin binding sites on the actin filaments
Cross-bridges now form, and pull on the actin filaments, using the energy of
ATP; this causes the sarcomere to shorten

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 Stimulus for Contraction 2

The contraction continues as long as the nerve impulse continues

After the nerve impulse stops, these events lead to relaxation of
the muscle:
• The enzyme acetylcholinesterase, in the motor end plate,
rapidly decomposes the acetylcholine
• Calcium is returned to the sarcoplasmic reticulum, using ATP as
an energy source
• ATP now binds to the myosin heads, and the linkages between
myosin and actin are broken
The actin returns to its original position and the muscle relaxes

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 Major Events of
TABLE 8.1 Major Events of Muscle Contraction and
Muscle Contraction Relaxation
Muscle Fiber Contraction
1. An impulse travels down a motor neuron axon.
2. The motor neuron releases the neurotransmitter
acetylcholine (ACh).
3. ACh binds to ACh receptors in the muscle fiber
membrane.
4. The sarcolemma is stimulated. An impulse travels over
the surface of the muscle fiber and deep into the fiber
through the transverse tubules.
5. The impulse reaches the sarcoplasmic reticulum, and
calcium channels open.
6. Calcium ions diffuse from the sarcoplasmic reticulum
into the cytosol and bind to troponin molecules.
7. Tropomyosin molecules move and expose specific sites
on actin where myosin heads can bind.
8. Cross-bridges form, linking thin and thick filaments.
9. Thin filaments are pulled toward the center of the
sarcomere by pulling of the cross-bridges.
10.The muscle fiber exerts a pulling force on its
attachments as a contraction occurs.

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 Major Events of Muscle Relaxation

TABLE 8.1 Major Events of Muscle Contraction and

Relaxation
Muscle Fiber Relaxation
1. Acetylcholinesterase decomposes acetylcholine, and
the muscle fiber membrane is no longer stimulated.
2. Calcium ions are actively transported into the
sarcoplasmic reticulum.
3. ATP breaks cross-bridge linkages between actin and
myosin filaments without breakdown of the ATP itself.
4. Breakdown of ATP “cocks” the myosin heads.
5. Troponin and tropomyosin molecules block the
interaction between myosin and actin filaments.
6. The muscle fiber remains relaxed, yet ready, until
stimulated again.

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 Energy Sources for Contraction
Energy for muscle fiber contraction comes from molecules of ATP; this chemical is in
limited supply, and so must be regenerated
The initial source of energy for muscle contraction is ATP that is stored in the muscle
Creatine phosphate is present to initially regenerate ATP from ADP and phosphate, as it
also contains high energy bonds
Whenever the supply of ATP is sufficient, the enzyme creatine phosphokinase promotes
the synthesis of creatine phosphate
As ATP decomposes, the energy from creatine phosphate can be transferred to ADP
molecules, converting them back to ATP
Creatine phosphate is rapidly used up too, and as its supply declines, the cell must rely
on cellular respiration to generate ATP

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 Oxygen Supply and Cellular Respiration

Glycolysis is the first phase of cellular respiration; it is anaerobic

and occurs in the cytoplasm; yields 2 ATP per molecule of glucose
Aerobic respiration it is aerobic (requires oxygen) and occurs in
the mitochondria; yields 28 ATP per molecule of glucose
Hemoglobin in red blood cells carries oxygen to muscle tissue
The pigment myoglobin stores oxygen in muscle tissue for aerobic
respiration; this increases oxygen availability

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 Oxygen Debt 1
During rest or moderate activity, there is enough oxygen available to
support aerobic respiration. Oxygen deficiency may develop during 1-2
minutes of strenuous exercise
In this case, pyruvic acid forms, and then reacts to form lactic acid,
which accumulates as an end product of anaerobic respiration in the
form of lactate.
Lactate diffuses out of muscle cells and is carried in the bloodstream to
the liver. Using ATP, liver cells convert lactate back into glucose

During strenuous exercise, oxygen is used to produce ATP for muscle

contraction, rather than for converting lactate back to glucose
As lactate builds up, an oxygen debt develops; this must be repaid

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 Oxygen Debt 2

Oxygen debt refers to the amount of oxygen that liver cells require
to convert the accumulated lactate back into glucose, plus the
amount that muscle cells need to resynthesize ATP and creatine
phosphate to their original concentrations
Oxygen debt is also called excess post-exercise oxygen
consumption
Repaying the oxygen debt may take several hours
Physical training helps to increase a muscle’s capacity to improve
energy production

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 Heat Production and Muscle Fatigue

Less than half of energy released in reactions of cellular respiration is used to form
ATP; the rest becomes heat

This heat is carried by the blood to other tissues, and helps maintain body
temperature

When a muscle loses its ability to contract during strenuous exercise, it is referred
to as fatigue

Muscle fatigue may arise from electrolyte imbalances and decreased ATP levels

A decrease in pH due to lactic acid accumulation may have a role in muscle fatigue

A muscle cramp, a sustained, painful, involuntary contraction, is thought to occur

due to changes in the extracellular fluid around the muscle fibers, leading to
uncontrolled muscle fiber stimulation by its motor neurons

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 Exercise and Muscle Use
Skeletal muscles respond to changes in activity level
• Hypertrophy: enlargement of a muscle due to repeated exercise
• Atrophy: decrease in muscle size and strength, due to disuse
Type of exercise determines responses of the muscle:
• Low intensity exercise causes slow fibers to increase mitochondria
and capillaries; they become more fatigue-resistant, but maintain size
and strength
• Forceful exercise causes fast fibers to increase numbers of actin and
myosin filaments, which enlarges fibers and entire muscle; this allows
for stronger contractions.
Number of skeletal muscle fibers does not change with hypertrophy or
atrophy

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 8.4: Muscular Responses

Threshold Stimulus: the minimum strength of stimulus required to

generate a impulse through the muscle fiber, release calcium ions,
activate cross-bridges, and contract the muscle
In the body, one motor neuron impulse releases sufficient
acetylcholine (ACh) in the neuromuscular junction to bring a
muscle fiber to its threshold
The response of a single muscle fiber to a single impulse is referred
to as a twitch; a twitch consists of a cycle of contraction and
relaxation
When a muscle fiber contracts, it contracts to its full extent, with
each twitch generating the same force; this is called the all-or-
none response

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 Myogram of a Single Muscle Twitch

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 Summation

A muscle fiber receiving a series of stimuli of increasing frequency

reaches a point when it is unable to relax completely, and the
force of individual twitches combine by the process of summation
Summation allows for a greater force of contraction than a single
twitch can generate
When exposed to higher frequency of stimulation, relaxation time
becomes very short; this is called partial tetany
If the frequency of stimulation is very high, and the sustained
contraction lacks any relaxation, it is called a complete tetanic
contraction
Partial tetany occurs in the body, but complete tetany can only be
accomplished in a lab

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 Myograms of Twitches, Summation, & Tetany

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 Recruitment of Motor Units

A motor neuron and the muscle fibers it controls make up a motor

unit; when stimulated, the muscle fibers of the motor unit
contract all at once
An increase in the number of activated motor units within a
muscle at higher intensities of stimulation is called motor unit
recruitment
Recruitment causes an increase in the strength of a contraction
A muscle achieves maximum tension when all of its motor units
have been recruited

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 Portions of Two Motor Units

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 Sustained Contractions and Muscle Tone

Summation and recruitment together can produce a sustained

contraction of increasing strength
Sustained contraction of muscles allow for performance of daily
activities
Muscle tone is a continuous state of sustained contraction of a few
motor units at a time within a muscle, even when at rest
Muscle tone is important for maintenance of posture

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 Types of Contractions
Muscles do not have to shorten to generate force
Types of muscle contractions:
• Isotonic (“same tension”) contraction:
• Involves shortening of muscle
• Associated with movement, such as lifting a weight
• As muscle shortens, tension remains the same
• Isometric (“same measurement”) contraction:
• Involves force generation without shortening
• Force is used to resist overstretching and oppose gravity
• Example: holding a weight in one position
Most movements are a combination of both types of contraction

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