Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
9
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
muscle cell and muscle fiber are two terms for the same structure.
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
flat.
Review: Which connective tissue coat surrounds groups of muscle fibers, separating them
into fascicles?
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.
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
band?
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.
Muscle contraction occurs because cross-bridges attach to and walk along the thin filaments at
both
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
crossbridge;
(3) the cross-bridge generates force as it rotates toward the center of the sarcomere (power
stroke);
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.
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.
2
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.
4
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.
SHORT VERSION:
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
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
single
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
arrives
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
detected).
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
units.
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.
pg 314
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.
Various types of exercises can induce changes in the fibers in a skeletal muscle. Endurance-type
(aerobic)
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
filaments.
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.
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
continuous,
rhythmic activity, cardiac muscle depends greatly on aerobic respiration to generate ATP.
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
OBJECTIVE
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
division
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
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.
Summary on pg 324