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Chapter 4, pages: 134 & table 4.

Chapter 10, pages: 292-308, 311-321

Chapter 4
Muscular tissue consists of elongated cells called muscle fibers or
myocytes that can use ATP to generate force

Based on location and certain

structural and functional features, muscular tissue is classified into
three types: skeletal, cardiac, and smooth

Chapter 10, pages: 292-308, 311-321

10.1 Overview of Muscular Tissue


Explain the structural differences among the three types

of muscular tissue.
Compare the functions and special properties of the three
types of muscular tissue.

Skeletal muscle tissue is primarily

attached to bones; it is striated and voluntary. Cardiac muscle tissue forms the wall of the heart;
it is
striated and involuntary. Smooth muscle tissue is located primarily in internal organs; it is
(smooth) and involuntary.
2. Through contraction and relaxation, muscular tissue performs four important functions:
body movements; stabilizing body positions; moving substances within the body and regulating
volume; and producing heat.
3. Four special properties of muscular tissues are (1) electrical excitability, the property of
to stimuli by producing action potentials; (2) contractility, the ability to generate tension to do
(3) extensibility, the ability to be extended (stretched); and (4) elasticity, the ability to return to
shape after contraction or extension.

10.2 Skeletal Muscle Tissue


Explain the importance of connective tissue components,

blood vessels, and nerves to skeletal muscles.
Describe the microscopic anatomy of a skeletal muscle
Distinguish thick filaments from thin filaments.
Describe the functions of skeletal muscle proteins.

muscle cell and muscle fiber are two terms for the same structure.

Connective Tissue Components

Connective tissue surrounds and protects muscular tissue. The subcutaneous
layer or hypodermis, which separates muscle from skin from muscles, provides a pathway
for blood vessels and nerves to enter and exit muscles, and protects muscles from physical

trauma. Fascia lines the body wall and limbs that surround and support muscles, allows free
movement of muscles, carries nerves and blood vessels, and fills space between muscles.

The adipose tissue of the subcutaneous layer stores most of the bodys triglycerides,
serves as an insulating layer that reduces heat loss, and protects muscles from physical
trauma. Fascia (FASH-e-a _ bandage) is a dense sheet or broad band of irregular
connective tissue that lines the body wall and limbs and supports and surrounds
muscles and other organs of the body. As you will see, fascia holds muscles with similar
functions together (see Figure 11.21). Fascia allows free
movement of muscles; carries nerves, blood vessels, and lymphatic vessels; and fills
spaces between muscles.
Three layers of connective tissue extend from the fascia to protect and strengthen
skeletal muscle (Figure 10.1):
Epimysium (epi- _ upon) is the outer layer, encircling the entire muscle. It consists of
dense irregular connective tissue.
Perimysium (peri- _ around) is also a layer of dense irregular connective tissue, but it
surrounds groups of 10 to 100 or more muscle fibers, separating them into bundles
called fascicles (little bundles). Many fascicles are large enough to be seen with the
naked eye. They give a cut of meat its characteristic grain; if you tear a piece of meat,
it rips apart along the fascicles.
Endomysium (endo- _ within) penetrates the interior of each fascicle and separates
individual muscle fibers from one another. The endomysium is mostly reticular fibers.

All three connective tissue layers may extend beyond the muscle fibers to form a
ropelike tendon. Tendons and aponeuroses are extensions of connective tissue beyond muscle
fibers that attach the muscle
to bone or to other muscle. A tendon is generally ropelike in shape; an aponeurosis is wide and

Review: Which connective tissue coat surrounds groups of muscle fibers, separating them
into fascicles?

Perimysium bundles groups of muscle fibers into fascicles.

Nerve and Blood Supply

Skeletal muscles are well supplied with nerves and blood vessels.
Generally, an artery and one or two veins accompany each nerve that penetrates a
skeletal muscle. Somatic motor neurons provide the nerve impulses that stimulate skeletal
muscle to contract. Microscopic blood vessels called capillaries are plentiful in muscular
tissue. The blood capillaries bring in oxygen and nutrients and remove heat and the
waste products of muscle metabolism.

Microscopic Anatomy of a Skeletal Muscle Fiber

The most important components of a skeletal muscle are the muscle fibers themselves.
The diameter of a mature skeletal muscle fiber ranges from 10 to 100 _m.* The typical
length of a mature skeletal muscle fiber is about 10 cm (4 in.), although some are as
long as 30 cm (12 in.).
Sarcolemma, Transverse Tubules, and Sarcoplasm
The major cells of skeletal muscle tissue are termed skeletal muscle fibers. Each muscle fiber has
or more nuclei because it arises from the fusion of many myoblasts. Satellite cells are myoblasts
persist after birth. The sarcolemma is a muscle fibers plasma membrane; it surrounds the
Transverse tubules are invaginations of the sarcolemma.

Myofibrils and Sarcoplasmic Reticulum

Each muscle fiber (cell) contains hundreds of myofibrils, the contractile elements of skeletal
Sarcoplasmic reticulum (SR) surrounds each myofibril. Within a myofibril are thin and thick
arranged in compartments called sarcomeres.

Dilated end sacs of the sarcoplasmic reticulum called terminal

cisterns (reservoirs) butt against the T tubule from both sides. A transverse tubule and
the two terminal cisterns on either side of it form a triad (tri- _ three). In a relaxed
muscle fiber, the sarcoplasmic reticulum stores calcium ions (Ca2_). Release of Ca2_ from
the terminal cisterns of the sarcoplasmic reticulum triggers muscle contraction
Which structure shown here releases calcium ions to trigger muscle contraction?

The sarcoplasmic reticulum releases calcium ions to trigger muscle contraction.

Filaments and the Sarcomere

Key Idea: Myofibrils contain two types of filaments: thick filaments and thin filaments.
Within myofibrils are smaller protein structures called filaments or myofilaments. Thin
filaments are 8 nm in diameter and 12 _m long* and composed mostly of the protein
actin, while thick filaments are 16 nm in diameter and 12 _m long and composed
mostly of the protein myosin. Both thin and thick filaments are directly involved in the
contractile process. Overall, there are two thin filaments for every thick filament in the
regions of filament overlap. The filaments inside a myofibril do not extend the entire

length of a muscle fiber. Instead, they are arranged in compartments called

The overlapping of thick and thin filaments produces striations. Darker A bands alternate with
lighter I
bands. Table 10.1 summarizes the components of the sarcomere.

Review: Which of the following is the smallest: muscle fiber, thick filament, or myofibril?
Which is largest?

The following are arranged from smallest to largest: thick filament, myofibril, muscle fiber.

Muscle Proteins
Myofibrils are composed of three types of proteins: contractile, regulatory, and structural. The
contractile proteins are myosin (thick filament) and actin (thin filament). Regulatory proteins are
tropomyosin and troponin, both of which are part of the thin filament. Structural proteins include
titin (links Z disc
to M line and stabilizes thick filament), myomesin (forms M line), nebulin (anchors thin filaments
to Z discs and regulates length of thin filaments during development), and dystrophin (links thin
filaments to sarcolemma). Table 10.2 summarizes the different types of skeletal muscle fiber
proteins. Table 10.3 summarizes the levels of organization within a skeletal muscle.
Which proteins connect into the Z disc? Which proteins are present in the A band? In the I

Actin and titin anchor into the Z disc. A bands contain myosin, actin, troponin, tropomyosin, and
titin; I bands contain actin, troponin, tropomyosin, and titin.

10.3 Contraction and Relaxation

of Skeletal Muscle Fibers

Outline the steps involved in the sliding filament

mechanism of muscle contraction.
Describe how muscle action potentials arise at the
neuromuscular junction.

Muscle contraction occurs because cross-bridges attach to and walk along the thin filaments at
ends of a sarcomere, progressively pulling the thin filaments toward the center of a sarcomere.
As the
thin filaments slide inward, the Z discs come closer together, and the sarcomere shortens.
2. The contraction cycle is the repeating sequence of events that causes sliding of the filaments:
(1) Myosin
ATPase hydrolyzes ATP and becomes energized; (2) the myosin head attaches to actin, forming a
(3) the cross-bridge generates force as it rotates toward the center of the sarcomere (power
and (4) binding of ATP to the myosin head detaches it from actin. The myosin head again
hydrolyzes the
ATP, returns to its original position, and binds to a new site on actin as the cycle continues.
Key idea: During muscle contractions, thin filaments move toward the M line of each sarcomere.

ExcitationContraction Coupling (not important?) Pg 304

The Neuromuscular Junction

The neuromuscular junction (NMJ) is the synapse between a somatic motor neuron and a skeletal
muscle fiber. The NMJ includes the axon terminals and synaptic end bulbs of a motor neuron, plus
adjacent motor end plate of the muscle fiber sarcolemma.

A synapse is a region where communication occurs between two neurons, or between

a neuron and a target cellin this case, between a somatic motor neuron and a muscle

fiber. At most synapses a small gap, called the synaptic cleft, separates the two cells.
Because the cells do not physically touch, the action potential cannot jump the gap
from one cell to another. Instead, the first cell communicates with the second by
releasing a chemical messenger called a neurotransmitter.
A nerve impulse (nerve action potential) elicits a muscle action
potential in the following way (Figure 10.9c):
1 Release of acetylcholine. Arrival of the nerve impulse at the synaptic end bulbs
stimulates voltage-gated channels to open. Because calcium ions are more
concentrated in the extracellular fluid, Ca2_ flows inward through the open channels. The
entering Ca2_ in turn stimulates the synaptic vesicles to undergo exocytosis. During
exocytosis, the synaptic vesicles fuse with the motor neurons plasma membrane,
liberating ACh into the synaptic cleft. The ACh then diffuses across the synaptic cleft
between the motor neuron and the motor end plate.
Activation of ACh receptors. Binding of two molecules of ACh to the receptor on the
motor end plate opens an ion channel in the ACh receptor. Once the channel is open,
small cations, most importantly Na_, can flow across the membrane.

Production of muscle action potential. The inflow of Na_ (down its electrochemical
gradient) makes the inside of the muscle fiber more positively charged. This change in
the membrane potential triggers a muscle action potential. Each nerve impulse
normally elicits one muscle action potential. The muscle action potential then
propagates along the sarcolemma into the system of T tubules. This causes the
sarcoplasmic reticulum to release its stored Ca 2_ into the sarcoplasm and the muscle
fiber subsequently contracts.

Termination of ACh activity. The effect of ACh binding lasts only briefly because ACh
is rapidly broken down by an enzyme called acetylcholinesterase (AChE). This
enzyme is attached to collagen fibers in the extracellular matrix of the synaptic cleft.
AChE breaks down
ACh into acetyl and choline, products that cannot activate the ACh receptor.

When a nerve impulse reaches the synaptic end bulbs of a somatic motor neuron, it triggers
exocytosis of the synaptic vesicles, which releases acetylcholine (ACh). ACh diffuses across the
synaptic cleft and binds to ACh receptors, initiating a muscle action potential.
Acetylcholinesterase then quickly breaks down ACh into its component parts.
Figure 10.10 on pg 308 also summarizes it.

Pg 311-321

10.5 Control of Muscle Tension


Describe the structure and function of a motor unit, and

define motor unit recruitment.
Explain the phases of a twitch contraction
Describe how frequency of stimulation affects muscle
tension, and how muscle tone is produced.
Distinguish between isotonic and isometric contractions

The number of impulses per second is the frequency ofstimulation.

A motor unit consists of a somatic motor neuron plus all of the skeletal muscle fibers it
Whole muscles that control precise movements consist of many small motor units
(Groups of motor units often work together to coordinate the contractions of a single muscle;
all of the motor units within a muscle are considered a motor pool.)
The process in which the number of active motor units increases is called motor unit
(process of increasing the number of active motor units).
Question: What is the effect of the size of a motor unit on its
strength of contraction? (Assume that each muscle fiber
can generate about the same amount of tension.)

Motor units having many muscle fibers are capable of more

forceful contractions than those having only a few fibers.

Twitch Contraction
A record of a contraction is called a myogram. It consists of a latent period, a contraction period,
and a relaxation period.
A twitch contraction is a brief contraction of all muscle fibers in a motor unit in response to a
action potential.

Twitches of skeletal muscle fibers last anywhere from 20 to 200 msec. This is very long
compared to the brief 12 msec* that a muscle action potential lasts.
The delay, which lasts about 2 msec, is termed the latent period. During the latent
period, the muscle action potential sweeps over the sarcolemma and calcium ions are
released from the sarcoplasmic reticulum. The second phase, the contraction period,
lasts 10100 msec. During this time, Ca2_ binds to troponin, myosin-binding sites on
actin are exposed, and cross-bridges form. Peak tension develops in the muscle fiber.
During the third phase, the relaxation period, also lasting 10100 msec, Ca2_ is
actively transported back into the sarcoplasmic reticulum.

During the latent period, the muscle action potential sweeps over the sarcolemma and calcium
ions are released from the sarcoplasmic reticulum.

Frequency of Stimulation
Wave summation is the increased strength of a contraction that occurs when a second stimulus
before the muscle fiber has relaxed completely following a previous stimulus.
Repeated stimuli can produce unfused (incomplete) tetanus, a sustained muscle contraction
with partial relaxation between stimuli. More rapidly repeating stimuli produce fused
(complete) tetanus, a sustained
contraction without partial relaxation between stimuli (in which individual twitches cannot be


Review: Would the peak force of the second contraction in part b be larger or smaller if
the second stimulus were applied a few milliseconds later?

10.14 If the second stimulus were applied a little later, the second contraction would be smaller
than the one illustrated in part b.

Muscle Tone
Even at rest, a skeletal muscle exhibits muscle tone (tonos _ tension), a small amount
of tautness or tension in the muscle due to weak, involuntary contractions of its motor
When the motor neurons serving a skeletal muscle are damaged or cut, the muscle
becomes flacid a state of limpness in which muscle tone is lost. To sustain muscle tone,
small groups of motor units are alternatively active and inactive in a constantly shifting
pattern. Muscle tone keeps skeletal muscles firm, but it does not result in a force strong
enough to produce movement. For example, when you are awake, the muscles in the
back of the neck are in normal tonic contraction; they keep the head upright and
prevent it from slumping forward on the chest.

Isotonic and Isometric Contractions

Continuous involuntary activation of a small number of motor units produces muscle tone, which
isessential for maintaining posture.
8. In a concentric isotonic contraction, the muscle shortens to produce movement and to reduce
the angle
at a joint. During an eccentric isotonic contraction, the muscle lengthens.
9. Isometric contractions, in which tension is generated without muscle changing its length, are
because they stabilize some joints as others are moved.
What type of contraction occurs in your neck muscles while you are walking?

10.15 Holding your head upright without movement involves mainly

isometric contractions.

pg 314

10.6 Types of Skeletal Muscle


Compare the structure and function of the three types of

skeletal muscle fibers.

Slow Oxidative Fibers

Slow oxidative (SO) fibers appear dark red because they contain large amounts of
myoglobin and many blood capillaries. Because they have many large mitochondria, SO
fibers generate ATP mainly by aerobic respiration, which is why they are called oxidative
fibers. These fibers are said to be slow because the ATPase in the myosin heads
hydrolyzes ATP relatively slowly and the contraction cycle proceeds at a slower pace
than in fast fibers. As a result, SO fibers have a slow speed of contraction. Their twitch
contractions last from 100 to 200 msec, and they take longer to reach peak tension.
However, slow fibers are very resistant to fatigue and are capable of prolonged,
sustained contractions for many hours. These slow-twitch, fatigue-resistant fibers are
adapted for maintaining posture and for aerobic, endurance-type activities such as
running a marathon.

Fast OxidativeGlycolytic Fibers

Fast oxidativeglycolytic (FOG) fibers are typically the largest fibers. Like slow
oxidative fibers, they contain large amounts of myoglobin and many blood capillaries.
Thus, they also have a dark red appearance. FOG fibers can generate considerable ATP
by aerobic respiration, which gives them a moderately high resistance to fatigue.
Because their intracellular glycogen level is high, they also generate ATP by anaerobic
glycolysis. FOG fibers are fast because the ATPase in their myosin heads hydrolyzes
ATP three to five times faster than the myosin ATPase in SO fibers, which makes their
speed of contraction faster. Thus, twitches of FOG fibers reach peak tension more
quickly than those of SO fibers but are briefer in durationless than 100 msec. FOG
fibers contribute to activities such as walking and sprinting.

Fast Glycolytic Fibers

Fast glycolytic (FG) fibers have low myoglobin content, relatively few blood
capillaries, and few mitochondria, and appear white in color. They contain large
amounts of glycogen and generate ATP mainly by glycolysis. Due to their ability to
hydrolyze ATP rapidly, FG fibers contract strongly and quickly. These fast-twitch fibers
are adapted for intense anaerobic movements of short duration, such as weight lifting
or throwing a ball, but they fatigue quickly. Strength training programs that engage a
person in activities requiring great strength for
short times increase the size, strength, and glycogen content of fast glycolytic fibers.
The FG fibers of a weight lifter may be 50% larger than those of a sedentary person or
an endurance athlete because of increased synthesis of muscle proteins. The overall
result is muscle enlargement due to hypertrophy of the FG fibers.
The motor units of a muscle are recruited in the following order: first SO fibers, then FOG fibers,
finally FG fibers

10.7 Exercise and Skeletal

Muscle Tissue

Describe the effects of exercise on different types of skeletal muscle fibers.

Various types of exercises can induce changes in the fibers in a skeletal muscle. Endurance-type
exercises cause a gradual transformation of some fast glycolytic (FG) fibers into fast oxidative
glycolytic (FOG) fibers.
2. Exercises that require great strength for short periods produce an increase in the size and
strength of fast
glycolytic (FG) fibers. The increase in size is due to increased synthesis of thick and thin
Activities such as weightlifting rely more on anaerobic production of ATP through glycolysis. Such
anaerobic training activities stimulate synthesis of muscle proteins and result, over time, in increased
muscle size (muscle hypertrophy). Muscular hypertrophy is due to increased production of myofibrils,
mitochondria, sarcoplasmic reticulum, and other organelles.

10.8 Cardiac Muscle Tissue


Describe the main structural and functional characteristics

of cardiac muscle tissue

1. Cardiac muscle is found only in the heart. Cardiac muscle fibers have the same arrangement
of actin
and myosin and the same bands, zones, and Z discs as skeletal muscle fibers. The fibers connect
to one
another through intercalated discs, which contain both desmosomes and gap junctions.
2. Cardiac muscle tissue remains contracted 10 to 15 times longer than skeletal muscle tissue
due to prolonged
delivery of Ca2_ into the sarcoplasm.
3. Cardiac muscle tissue contracts when stimulated by its own autorhythmic fibers. Due to its
rhythmic activity, cardiac muscle depends greatly on aerobic respiration to generate ATP.

10.9 Smooth Muscle Tissue


Describe the main structural and functional characteristics

of smooth muscle tissue.

10.9 Smooth Muscle Tissue

1. Smooth muscle is nonstriated and involuntary.
2. Smooth muscle fibers contain intermediate filaments and dense bodies; the function of dense
bodies is
similar to that of the Z discs in striated muscle.
3. Visceral (single-unit) smooth muscle is found in the walls of hollow viscera and of small blood
Many fibers form a network that contracts in unison.
4. Multiunit smooth muscle is found in large blood vessels, large airways to the lungs, arrector pili
and the eye, where it adjusts pupil diameter and lens focus. The fibers operate independently
than in unison.

Physiology of Smooth Muscle

5. The duration of contraction and relaxation of smooth muscle is longer than in skeletal muscle
since it
takes longer for Ca2_ to reach the filaments.
6. Smooth muscle fibers contract in response to nerve impulses, hormones, and local factors.
7. Smooth muscle fibers can stretch considerably and still maintain their contractile function.

Visceral smooth muscle fibers connect to one another by gap junctions and contract as a single unit.
Multiunit smooth muscle fibers lack gap junctions and contract independently.
Which type of smooth muscle is more like cardiac muscle than skeletal muscle, with
respect to both its structure and function?

10.16 Visceral smooth muscle is more like cardiac muscle; both contain gap junctions, which
allow action potentials to spread from each cell to its neighbors.

10.10 Regeneration of
Muscular Tissue

Explain how muscle fibers regenerate.

Skeletal muscle fibers cannot divide and have limited powers of regeneration; cardiac muscle
fibers can
regenerate under limited circumstances; and smooth muscle fibers have the best capacity for
and regeneration.

Because mature skeletal muscle fibers have lost the ability to undergo cell division,
growth of skeletal muscle after birth is due mainly to hypertrophy, the enlargement of
existing cells, rather than to hyperplasia, an increase in the number of fibers. Satellite
cells divide slowly and fuse with existing fibers to assist both in muscle growth and in
repair of damaged fibers. Thus, skeletal muscle tissue can regenerate only to a limited
extent. certain smooth
Muscle fibers, such as those in the uterus, retain their capacity for division and thus can
grow by hyperplasia. Also, new smooth muscle fibers can arise from cells called
pericytes, stem cells found in association with blood capillaries and small veins. Smooth
muscle fibers can also proliferate in certain pathological conditions

Which type of muscular tissue has the highest capacity

for regeneration?

smooth muscle tissue has considerably greater powers of regeneration. Such powers
are still limited when compared with other tissues, such as epithelium.
Table 10.5 summarizes the major characteristics of the three
types of muscular tissue.

10.17 The myotome of a somite differentiates into skeletal muscle.

10.5 The I bands and H zones disappear during muscle contraction; the lengths of the thin and
thick filaments do not change.
10.6 If ATP were not available, the cross-bridges would not be able to detach from actin. The
muscles would remain in a state of rigidity, as occurs in rigor mortis.
10.8 Why is tension maximal at a sarcomere length of 2.2 _m?

Summary on pg 324

Terms mentioned in class:

When a muscle fiber receives enough stimulation to contract, it temporarily loses its
excitability and cannot respond for a time. The period of lost excitability, called the
refractory period, is a characteristic of all muscle and nerve cells. The duration of the
refractory period varies with the muscle involved. Skeletal muscle has a short refractory
period of about 5 msec; cardiac muscle has a longer refractory period of about 300