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Chapter 8

Histology and
Physiology of
Muscles

Skeletal Muscle Fibers
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

Functions of the Muscular System
1. Body movement (Skeletal Muscle)
2. Maintenance of posture (Skeletal
Muscle)
3. Respiration (Skeletal Muscle)
4. Production of body heat (Skeletal
Muscle)
5. Communication (Skeletal Muscle)
6. Constriction of organs and vessels
(Smooth Muscle)
7. Heartbeat (Cardiac Muscle)

Functional Characteristics of Muscle

1. Contractility: the ability to shorten
forcibly
2. Excitability: the ability to receive and
respond to stimuli
3. Extensibility: the ability to be stretched
or extended
4. Elasticity: the ability to recoil and
resume the original resting length

Types of Muscle Tissue
• The three types of muscle tissue are
skeletal, smooth, and cardiac
• These types differ in
– Structure
– Location
– Function
– Means of activation

• Each muscle is a discrete organ
composed of muscle tissue, blood vessels,
nerve fibers, and connective tissue

Types of Muscle Tissue
• Skeletal muscles are responsible for most body
movements
– Maintain posture, stabilize joints, and generate heat

• Smooth muscle is found in the walls of hollow
organs and tubes, and moves substances through
them
– Helps maintain blood pressure
– Squeezes or propels substances (i.e., food, feces) through
organs

• Cardiac muscle is found in the heart and pumps
blood throughout the body

Tab. 8.1

Skeletal Muscle Structure • Skeletal muscle cells are elongated and are often called skeletal muscle fibers • Each skeletal muscle cell contains several nuclei located around the periphery of the fiber near the plasma membrane • Fibers appear striated due to the actin and myosin myofilaments • A single fiber can extend from one end of a muscle to the other • Contracts rapidly but tires easily • Is controlled voluntarily (i. by conscious control) ..e.

which separate and compartmentalize individual muscles or groups of muscles are: – Epimysium: an overcoat of dense collagenous connective tissue that surrounds the entire muscle – Perimysium: fibrous connective tissue that surrounds groups of muscle fibers called fascicles (bundles) – Endomysium: fine sheath of connective tissue composed of reticular fibers surrounding each muscle fiber .Skeletal Muscle Structure • Fascia is a general term for connective tissue sheets • The three muscular fascia.

8.Skeletal Muscle Structure • The connective tissue of muscle provides a pathway for blood vessels and nerves to reach muscle fibers Fig.1 .

which connect muscle to bone Fig.Skeletal Muscle Structure • The connective tissue of muscle blends with other connective tissue based structures. such as tendons.2 . 8.

mys. and sarco: prefixes used to refer to muscle – Muscle contraction depends on two kinds of myofilaments: actin and myosin • Myofibrils are densely packed. rod-like contractile elements • They make up most of the muscle volume .Skeletal Muscle Structure • Muscle Fibers – Terminology • Sarcolemma: muscle cell plasma membrane • Sarcoplasm: cytoplasm of a muscle cell • Myo.

2 .Fig. 8.

8.Skeletal Muscle Structure • Actin (thin) myofilaments consist of two helical polymer strands of F actin (composed of G actin). tropomyosin.2 . and troponin • The G actin contains the active sites to which myosin heads attach during contraction • Tropomyosin and troponin are regulatory subunits bound to actin Fig.

which enables the head to move – A rod • A cross-bridge is formed when a myosin head binds to the active site on G actin Fig. which breaks down ATP – A hinge region. 8.2 .Skeletal Muscle Structure • Myosin (thick) myofilaments consist of myosin molecules • Each myosin molecule has – A head with an ATPase.

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Skeletal Muscle Structure • Sarcomeres – The smallest contractile unit of a muscle – Sarcomeres are bound by Z disks that hold actin myofilaments – Six actin myofilaments surround a myosin myofilament – Myofibrils appear striated because of A bands and I bands Fig.2 . 8.

Skeletal Muscle Structure • Thick filaments: extend the entire length of an A band • Thin filaments: extend across the I band and partway into the A band • Z-disc: a coin-shaped sheet of proteins (connectins) that anchors the thin filaments and connects myofibrils to one another • Thin filaments do not overlap thick filaments in the lighter H zone • M lines appear darker due to the presence of the protein desmin • The arrangement of myofibrils within a fiber is so organized a perfectly aligned repeating series of dark A bands and light I bands is evident .

3bc . 8.Fig.

the muscle shortens • The I band and H zones become narrower during contraction. myosin heads bind to actin and sliding begins – Each myosin head binds and detaches several times during contraction (acting like a ratchet to generate tension and propel the thin filaments to the center of the sarcomere) • In the relaxed state. thin and thick filaments overlap only slightly • As this event occurs throughout the sarcomeres. and the A band remains constant in length .Sliding Filament Model • Actin and myosin myofilaments do not change in length during contraction • Thin filaments slide past the thick ones so that the actin and myosin filaments overlap to a greater degree – Upon stimulation.

Fig.4 . 8.

4 .Sliding Filament Model • Actin and myosin myofilaments in a relaxed muscle (below) and a contracted muscle are the same length. 8. Myofilaments do not change length during muscle contraction Fig.

and the sarcomere shortens Fig. 8. As a result.4 . actin myofilaments at each end of the sarcomere slide past the myosin myofilaments toward each other. the Z disks are brought closer together.Sliding Filament Model • During contraction.

which are equal to the length of the myosin myofilaments. The A bands. 8.Sliding Filament Model • As the actin myofilaments slide over the myosin myofilaments.4 . do not narrow because the length of the myosin myofilaments does not change Fig. the H zones (yellow) and the I bands (blue) narrow.

4 . the ends of the actin myofilaments overlap at the center of the sarcomere and the H zone disappears Fig.Sliding Filament Model • In a fully contracted muscle. 8.

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Physiology of Skeletal Muscle Fibers • Membrane Potentials – The nervous system stimulates muscles to contract through electric signals called action potentials – Plasma membranes are polarized. which means there is a charge difference (resting membrane potential) across the plasma membrane – The inside of the plasma membrane is negative as compared to the outside in a resting cell – An action potential is a reversal of the resting membrane potential so that the inside of the plasma membrane becomes positive .

Physiology of Skeletal Muscle Fibers • Ion Channels – Assist with the production of action potentials • Ligand-gated channels • Voltage-gated channels .

Fig.5 . 8.

Fig.6 . 8.

an action potential is produced – The depolarization phase of the action potential results from the opening of many Na+ channels Fig.6 .Physiology of Skeletal Muscle Fibers • Action Potentials – Depolarization results from an increase in the permeability of the plasma membrane to Na+ – If depolarization reaches threshold. 8.

8.Physiology of Skeletal Muscle Fibers • Action Potentials – The repolarization phase of the action potential occurs when the Na+ channels close and K+ channels open briefly Fig.6 .

Fig.6 . 8.

Physiology of Skeletal Muscle Fibers • Action Potentials – Occur in an all-or-none fashion • A stimulus below threshold produces no action potential • A stimulus at threshold or stronger will produce an action potential – Propagate (travel) across plasma membranes .

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Physiology of Skeletal Muscle Fibers • Nerve Stimulus of Skeletal Muscle – Skeletal muscles are stimulated by motor neurons of the somatic nervous system – Axons of these neurons travel in nerves to muscle cells – Axons of motor neurons branch profusely as they enter muscles – Each axonal branch forms a neuromuscular junction with a single muscle fiber .

Physiology of Skeletal Muscle Fibers • The neuromuscular junction is formed from: – Axonal endings • Have small membranous sacs (synaptic vesicles) • Contain the neurotransmitter acetylcholine (ACh) – Motor end plate of a muscle • Specific part of the sarcolemma • Contains ACh receptors • Though exceedingly close. axonal ends and muscle fibers are always separated by a space called the synaptic cleft .

8.7 .Fig.

8. ACh is released into the synaptic cleft by exocytosis Fig.8 . Calcium ions enter the presynaptic terminal and initiate the release of the neurotransmitter acetylcholine (ACh) from synaptic vesicles 3.Neuromuscular Junction Physiology 1. An action potential (orange arrow) arrives at the presynaptic terminal and causes voltage-gated Ca2+ channels in the presynaptic membrane to open 2.

Calcium ions enter the presynaptic terminal and initiate the release of the neurotransmitter acetylcholine (ACh) from synaptic vesicles 3.8 . An action potential (orange arrow) arrives at the presynaptic terminal and causes voltage-gated Ca2+ channels in the presynaptic membrane to open 2. 8. ACh is released into the synaptic cleft by exocytosis Fig.Neuromuscular Junction Physiology 1.

8.8 .Neuromuscular Junction Physiology 1. Calcium ions enter the presynaptic terminal and initiate the release of the neurotransmitter acetylcholine (ACh) from synaptic vesicles 3. An action potential (orange arrow) arrives at the presynaptic terminal and causes voltage-gated Ca2+ channels in the presynaptic membrane to open 2. ACh is released into the synaptic cleft by exocytosis Fig.

8 . Ligand-gated Na+ channels open and Na+ enters the postsynaptic cell. ACh is removed from the ligand-gated Na+ channels. an action potential is generated along the postsynaptic membrane 6.Neuromuscular Junction Physiology 4. ACh diffuses across the synaptic cleft and binds to ligand-gated Na+ channels on the postsynaptic membrane 5. 8. If depolarization passes threshold. causing the postsynaptic membrane to depolarize. which then close Fig.

If depolarization passes threshold.Neuromuscular Junction Physiology 4.8 . an action potential is generated along the postsynaptic membrane 6. Ligand-gated Na+ channels open and Na+ enters the postsynaptic cell. 8. ACh is removed from the ligand-gated Na+ channels. ACh diffuses across the synaptic cleft and binds to ligand-gated Na+ channels on the postsynaptic membrane 5. which then close Fig. causing the postsynaptic membrane to depolarize.

Neuromuscular Junction Physiology 4. causing the postsynaptic membrane to depolarize. an action potential is generated along the postsynaptic membrane 6. ACh diffuses across the synaptic cleft and binds to ligand-gated Na+ channels on the postsynaptic membrane 5. Ligand-gated Na+ channels open and Na+ enters the postsynaptic cell. 8. ACh is removed from the ligand-gated Na+ channels. If depolarization passes threshold.8 . which then close Fig.

Choline is symported with Na+ into the presynaptic terminal. ACh is then taken up by the synaptic vesicles Fig.8 . which is attached to the postsynaptic membrane. 8. Acetic acid diffuses away from the synaptic cleft 9. removes acetylcholine from the synaptic cleft by breaking it down into acetic acid and choline 8. The enzyme acetylcholinesterase. where it can be recycled to make ACh. ACh is reformed within the presynaptic terminal using acetic acid generated from metabolism and choline recycled from the synaptic cleft.Neuromuscular Junction Physiology 7.

8. which is attached to the postsynaptic membrane. ACh is then taken up by the synaptic vesicles Fig.Neuromuscular Junction Physiology 7. Acetic acid diffuses away from the synaptic cleft 9. Choline is symported with Na+ into the presynaptic terminal. ACh is reformed within the presynaptic terminal using acetic acid generated from metabolism and choline recycled from the synaptic cleft.8 . where it can be recycled to make ACh. The enzyme acetylcholinesterase. removes acetylcholine from the synaptic cleft by breaking it down into acetic acid and choline 8.

8 . ACh is then taken up by the synaptic vesicles Fig. Choline is symported with Na+ into the presynaptic terminal. which is attached to the postsynaptic membrane. The enzyme acetylcholinesterase. where it can be recycled to make ACh.Neuromuscular Junction Physiology 7. Acetic acid diffuses away from the synaptic cleft 9. 8. ACh is reformed within the presynaptic terminal using acetic acid generated from metabolism and choline recycled from the synaptic cleft. removes acetylcholine from the synaptic cleft by breaking it down into acetic acid and choline 8.

Fig.8 . 8.

along its sarcolemma – Have a rise in intracellular Ca2+ levels.Excitation-Contraction Coupling • In order to contract. a skeletal muscle must: – Be stimulated by a nerve ending – Propagate an electrical current. the final trigger for contraction • Linking the electrical signal to the contraction is excitation-contraction coupling . or action potential.

smooth endoplasmic reticulum that mostly runs longitudinal and surrounds each myofibril – Paired terminal cisternae form perpendicular cross channels – Functions in the regulation of intracellular calcium levels • A triad is a T tubule and two terminal cisternae Fig. 8.Excitation-Contraction Coupling • Invaginations of the sarcolemma form T tubules.9 . which wrap around the sarcomeres and penetrate into the cell’s interior at each A band –I band junction • Sarcoplasmic reticulum (SR) is an elaborate.

resulting in an increase in the permeability of the SR to Ca2+. Once active sites on G actin molecules are exposed. the heads of the myosin myofilaments bind to them to form cross-bridges . Calcium ions then diffuse from the SR into the sarcoplasm 3. causing tropomyosin to move.Excitation-Contraction Coupling 1. Calcium ions released from the SR bind to troponin molecules. especially in the terminal cisternae. The depolarization also spreads along the membrane of the T tubules 2. exposing the active sites on G actin 4. The depolarization of the T tubule causes gated Ca2+ channels in the SR to open. The troponin molecules bound to G actin molecules are released. An action potential produced at the presynaptic terminal in the neuromuscular junction is propagated along the sarcolemma of the skeletal muscle.

causing tropomyosin to move. The depolarization also spreads along the membrane of the T tubules 2. resulting in an increase in the permeability of the SR to Ca2+. Calcium ions then diffuse from the SR into the sarcoplasm 3. An action potential produced at the presynaptic terminal in the neuromuscular junction is propagated along the sarcolemma of the skeletal muscle. the heads of the myosin myofilaments bind to them to form cross-bridges . The depolarization of the T tubule causes gated Ca2+ channels in the SR to open. The troponin molecules bound to G actin molecules are released. exposing the active sites on G actin 4. Calcium ions released from the SR bind to troponin molecules.Excitation-Contraction Coupling 1. especially in the terminal cisternae. Once active sites on G actin molecules are exposed.

causing tropomyosin to move. The depolarization also spreads along the membrane of the T tubules 2. Once active sites on G actin molecules are exposed. The depolarization of the T tubule causes gated Ca2+ channels in the SR to open.Excitation-Contraction Coupling 1. exposing the active sites on G actin 4. The troponin molecules bound to G actin molecules are released. especially in the terminal cisternae. Calcium ions released from the SR bind to troponin molecules. the heads of the myosin myofilaments bind to them to form cross-bridges . resulting in an increase in the permeability of the SR to Ca2+. An action potential produced at the presynaptic terminal in the neuromuscular junction is propagated along the sarcolemma of the skeletal muscle. Calcium ions then diffuse from the SR into the sarcoplasm 3.

The depolarization also spreads along the membrane of the T tubules 2. exposing the active sites on G actin 4. Once active sites on G actin molecules are exposed. An action potential produced at the presynaptic terminal in the neuromuscular junction is propagated along the sarcolemma of the skeletal muscle.Excitation-Contraction Coupling 1. Calcium ions released from the SR bind to troponin molecules. causing tropomyosin to move. The depolarization of the T tubule causes gated Ca2+ channels in the SR to open. the heads of the myosin myofilaments bind to them to form cross-bridges . Calcium ions then diffuse from the SR into the sarcoplasm 3. resulting in an increase in the permeability of the SR to Ca2+. especially in the terminal cisternae. The troponin molecules bound to G actin molecules are released.

The depolarization of the T tubule causes gated Ca2+ channels in the SR to open. resulting in an increase in the permeability of the SR to Ca2+. An action potential produced at the presynaptic terminal in the neuromuscular junction is propagated along the sarcolemma of the skeletal muscle. Once active sites on G actin molecules are exposed. Calcium ions then diffuse from the SR into the sarcoplasm 3. Calcium ions released from the SR bind to troponin molecules. especially in the terminal cisternae. causing tropomyosin to move. the heads of the myosin myofilaments bind to them to form cross-bridges . exposing the active sites on G actin 4. The depolarization also spreads along the membrane of the T tubules 2. The troponin molecules bound to G actin molecules are released.Excitation-Contraction Coupling 1.

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8.Fig.11 .

11 .Fig. 8.

Fig.11 . 8.

8.11 .Fig.

8.11 .Fig.

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11 . 8.Fig.

11 .Fig. 8.

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preventing further cross-bridge formation .Muscle Relaxation • Calcium ions are transported back into the sarcoplasmic reticulum • Calcium ions diffuse away from troponin and tropomyosin moves.

and relaxation phases Table 8. contraction.2 .Muscle Twitch • The contraction of a muscle as a result of one or more muscle fibers contracting • Has lag.

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Strength of Muscle Contraction • For a given condition. a muscle fiber or motor unit contracts with a consistent force in response to each action potential • For a whole muscle. stimuli of increasing strength result in graded contractions of increased force as more motor units are recruited (multiple motor unit summation) • Stimulus of increasing frequency increase the force of contraction (multiple-wave summation) .

Motor Unit Fig.13 . 8.

and complete tetanus is no relaxation between contractions • The force of contraction of a whole muscle increases with increased frequency of stimulation because of an increasing concentration of Ca2+ around the myofibrils • Treppe is an increase in the force of contraction during the first few contractions of a rested muscle .Strength of Muscle Contraction • Incomplete tetanus is partial relaxation between contractions.

Multiple Motor Unit Summation in a Muscle Fig. 8.14 .

8.15 .Multiple-Wave Summation Fig.

8.15 .Treppe Fig.

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steady muscle contractions .Types of Muscle Contraction • Isometric contractions cause a change in muscle tension but no change in muscle length • Isotonic contractions cause a change in muscle length but no change in muscle tension • Concentric contractions are isotonic contractions that cause muscles to shorten • Eccentric contractions are isotonic contractions that enable muscles to shorten • Muscle tone is the maintenance of a steady tension for long periods • Asynchronous contractions of motor units produce smooth.

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17 . 8.Muscle Length and Tension • Muscle contracts with less than maximum force if its initial length is shorter or longer than optimal Fig.

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Fatigue • The decreased ability to do work • Can be caused by – The central nervous system (psychologic fatigue) – Depletion of ATP in muscles (muscular fatigue) • Physiologic contracture (the inability of muscles to contract or relax) and rigor mortis (stiff muscles after death) result from inadequate amounts of ATP .

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Energy Sources • Creatine phosphate – ATP is synthesized when ADP reacts with creatine phosphate to form creatine and ATP – ATP from this source provides energy for a short time Fig. 8.18 .

Energy Sources
• Anaerobic respiration
– ATP synthesized provides
energy for a short time at
the beginning of exercise
and during intense
exercise
– Produces ATP less
efficiently but more rapidly
than aerobic respiration
– Lactic acid levels increase
because of anaerobic
respiration
Fig. 8.18

Energy Sources
• Aerobic respiration
– Requires oxygen
– Produces energy
for muscle
contractions under
resting conditions
or during
endurance exercise

Fig. 8.18

Fig. 8.18

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Speed of Contraction • The three main types of skeletal muscle fibers are – Slow-twitch oxidative (SO) fibers – Fast-twitch glycolytic (FG) fibers – Fast-twitch oxidative glycolytic (FOG) fibers • SO fibers contract more slowly than FG and FOG fibers because they have slower myosin ATPases than FG and FOG fibers .

and myoglobin • FG fibers are fatigable – Rely on anaerobic respiration and have a high concentration of glycogen • FOG fibers have fatigue resistance intermediate between SO and FG fibers – Rely on aerobic and anaerobic respiration . a rich blood supply.Fatigue Resistance • SO fibers are fatigue-resistant and rely on aerobic respiration – Many mitochondria.

Functions • SO fibers maintain posture and are involved with prolonged exercise – Long-distance runners have a higher percentage of SO fibers • FG fibers produce powerful contractions of short duration – Sprinters have a higher percentage of FG fibers • FOG fibers support moderate-intensity endurance exercises – Aerobic exercise can result in the conversion of FG fibers to FOG fibers .

3 .Tab. 8.

Muscular Hypertrophy and Atrophy • Hypertrophy is an increase in the size of muscles – Due to an increase in the size of muscle fibers resulting from an increase in the number of myofibrils in the muscle fibers • Aerobic exercise – Increases the vascularity of muscle – Greater hypertrophy of slow-twitch fibers than fast-twitch fibers • Intense anaerobic exercise – Greater hypertrophy of fast-twitch fibers than slow-twitch • Atrophy is a decrease in the size of muscle – Due to a decrease in the size of muscle fibers or a loss of muscle fibers .

Effects of Aging on Skeletal Muscle • By 80 years of age 50% of the muscle mass is gone – Due to a loss in muscle fibers – Fast-twitch muscle fibers decrease in number more rapidly than slow-twitch fibers • Can be dramatically slowed if people remain physically active .

such as the stomach. and respiratory passages – Forces food and other substances through internal body channels – It is not striated and is involuntary . urinary bladder.Types of Smooth Muscle • Visceral smooth muscle fibers have many gap junctions and contract as a single unit • Multiunit smooth muscle fibers have few gap junctions and function independently – Found in the walls of hollow visceral organs.

Regulation of Smooth Muscle • Contraction is involuntary – Multiunit smooth muscle contracts when externally stimulated by nerves. hormones. or other substances – Visceral smooth muscle contracts autorhythmically or when stimulated externally • Hormones are important in regulating smooth muscle .

Structure of Smooth Muscle Cells • Spindle-shaped with a single nucleus • Have actin and myosin myofilaments – Actin myofilaments are connected to dense bodies and dense areas • Not striated • No T tubule system and most have less SR than skeletal muscle • No troponin .

cross-bridges form • Relaxation results when myosin phosphatase removes a phosphate group from the myosin molecule .Contraction and Relaxation of Smooth Muscle • Calcium ions enter the cell to initiate contraction – Bind to calmodulin – Activate myosin kinase. which transfers a phosphate group from ATP to myosin – When phosphate groups are attached to myosin.

19 . 8.Fig.

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Functional Properties of Smooth Muscle • Pacemaker cells are autorhythmic smooth muscle cells that control the contraction of other smooth muscle cells • Smooth muscle cells contract more slowly than skeletal muscle cells • Smooth muscle tone is the ability of smooth muscle to maintain a steady tension for long periods with little expenditure of energy • Smooth muscle in the walls of hollow organs maintain a relatively constant pressure on the contents of the organ despite changes in content volume • The force of smooth muscle contraction remains nearly constant despite changes in muscle length .

Cardiac Muscle Cells • Occurs only in the heart • Is striated like skeletal muscle but is not voluntary • Have a single nucleus • Connected by intercalated disks that allowing them to function as a single unit • Capable of autorhythmicity • Contracts at a fairly steady rate set by the heart’s pacemaker • Neural controls allow the heart to respond to changes in bodily needs .

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8.Tab.1 .

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