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Skeletal Muscle Tissue
- Attached to bones and skin
- Striated
- Voluntary (i.e., conscious control)
- Powerful
- Primary topic of this chapter
- Each muscle is served by one artery, one nerve, and one or more veins
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Cardiac Muscle
- Only in the heart
- Striated
- Involuntary
- More details in Chapter 18
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Smooth Muscle Tissue
- In the walls of hollow organs, e.g., stomach, urinary bladder, and airways
- Not striated
- Involuntary
- More details later in this chapter
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Special Characteristics of Muscle Tissue
- Excitability (responsiveness or irritability): ability to receive and respond to stimuli
- Contractility: ability to shorten when stimulated
- Extensibility: ability to be stretched
- Elasticity: ability to recoil to resting length
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Muscle Functions
- Movement of bones or fluids (e.g., blood)
- Maintaining posture and body position
- Stabilizing joints
- Heat generation (especially skeletal muscle)
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Connective tissue sheaths of skeletal muscle:
- Epimysium: dense regular connective tissue surrounding entire muscle
- Perimysium: fibrous connective tissue surrounding fascicles (groups of muscle fibers)
- Endomysium: fine areolar connective tissue surrounding each muscle fiber
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Muscle Attachments
Directly—epimysium of muscle is fused to the periosteum of bone or perichondrium of cartilage
Indirectly—connective tissue wrappings extend beyond the muscle as a ropelike tendon or sheetlike aponeurosis
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Charectoristics of a Muscle Cell
*Cylindrical cell 10 to 100 micrometers in diameter, up to 30 cm long
*Multiple peripheral nuclei
*Many mitochondria
*Glycosomes for glycogen storage, myoglobin for O2 storage
*Also contain myofibrils, sarcoplasmic reticulum, and T tubules
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Myofibrils
- *Densely packed, rodlike elements
- *~80% of cell volume
- *Exhibit striations: perfectly aligned repeating series of dark A bands and light I bands
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Sarcomere
- *Smallest contractile unit (functional unit) of a muscle CELL
- *The region of a myofibril between two successive Z discs
- *Composed of thick and thin myofilaments made of contractile proteins
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Features of a Sarcomere
- *Thin filaments: run the length of the I band and partway into the A band
- *Z disc: coin-shaped sheet of proteins that anchors the thin filaments and connects myofibrils to one another
- *Thick filaments: run the entire length of an A band
- *H zone: lighter midregion where filaments do not overlap
- *M line: line of protein myomesin that holds adjacent thick filaments together
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Ultrastructure of Thick Filament
- Composed of the protein myosin
- *Myosin tails contain: 2 interwoven, heavy polypeptide chains
- *Myosin heads contain:
- - 2 smaller, light polypeptide chains that act as cross bridges during contraction
- -Binding sites for actin of thin filaments
- -Binding sites for ATP
- -ATPase enzymes
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Ultrastructure of Thin Filament
- *Twisted double strand of fibrous protein F actin
- *F actin consists of G (globular) actin subunits
- *G actin bears active sites for myosin head attachment during contraction
- *Tropomyosin and troponin: regulatory proteins bound to actin
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Sarcoplasmic Reticulum (SR)
- *Network of smooth endoplasmic reticulum surrounding each myofibril
- *Pairs of terminal cisternae form perpendicular cross channels
- *Functions in the regulation of intracellular Ca2+ levels
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T Tubules
- *Continuous with the sarcolemma
- *Penetrate the cell’s interior at each A band–I band junction
- *Associate with the paired terminal cisternae to form triads that encircle each sarcomere
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Triad Relationships
- *T tubules conduct impulses deep into muscle fiber
- *Integral proteins protrude into the intermembrane space from T tubule and SR cisternae membranes
- *T tubule proteins: voltage sensors
- *SR foot proteins: gated channels that regulate Ca2+ release from the SR cisternae
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Contraction
- *The generation of force
- *Does not necessarily cause shortening of the fiber
- *Shortening occurs when tension generated by cross bridges on the thin filaments exceeds forces opposing shortening
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Sliding Filament Model of Contraction
*In the relaxed state, thin and thick filaments overlap only slightly
*During contraction, myosin heads bind to actin, detach, and bind again, to propel the thin filaments toward the M line
*As H zones shorten and disappear, sarcomeres shorten, muscle cells shorten, and the whole muscle shortens
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Requirements for Skeletal Muscle Contraction
1. Activation: neural stimulation at aneuromuscular junction
- 2. Excitation-contraction coupling:
- -Generation and propagation of an action potential along the sarcolemma
-Final trigger: a brief rise in intracellular Ca2+ levels
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Events at the Neuromuscular Junction
- *Skeletal muscles are stimulated by somatic motor neurons
- *Axons of motor neurons travel from the central nervous system via nerves to skeletal muscles
- *Each axon forms several branches as it enters a muscle
- *Each axon ending forms a neuromuscular junction with a single muscle fiber
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Events at NeuroMuscular Junction
 - 1. Actionpotential arrives at the axon terminal of a motor neauron
- 2.Voltage Gated Calcium (Ca2) chammels open and the Ca2 enters th axon terminal
- 3.Ca2+ entry causes some synaptic vesicles to release their contents (acetylcholine)by exocytosis.
- 4.Acetylcholine, aneurotransmitter, diffuses across the synaptic cleft and binds to receptors in the sarcolemma.
- 5.ACh binding opens ionchannels that allow simultaneous passage of Na+ into the musclefiber and K+ out of the muscle fiber.
- 6.ACh effects are terminated by its enzymatic breakdown in the synaptic cleft by acetylcholinesterase.
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Destruction of Acetylcholine
*ACh effects are quickly terminated by the enzyme acetylcholinesterase
*Prevents continued muscle fiber contraction in the absence of additional stimulation
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Events in Generation of an Action Potential
- 1.Local depolarization (end plate potential):
- -ACh binding opens chemically (ligand) gated ion channels
- -Simultaneous diffusion of Na+ (inward) and K+ (outward)
- -More Na+ diffuses, so the interior of the sarcolemma becomes less negative
- -Local depolarization – end plate potential
- 2.Generation and propagation of an action potential:
- End plate potential spreads to adjacent membrane areas
- Voltage-gated Na+ channels open
- Na+ influx decreases the membrane voltage toward a critical threshold
- If threshold is reached, an action potential is generated
- Local depolarization wave continues to spread, changing the permeability of the sarcolemma
- Voltage-regulated Na+ channels open in the adjacent patch, causing it to depolarize to threshold
- 3.Repolarization:
- Na+ channels close and voltage-gated K+ channels open
- K+ efflux rapidly restores the resting polarity
- Fiber cannot be stimulated and is in a refractory period until repolarization is complete
- Ionic conditions of the resting state are restored by the Na+-K+ pump
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Role of Calcium (Ca2+) in Contraction
- At higher intracellular Ca2+ concentrations:
- Ca2+ binds to troponin Troponin
- changes shape and moves tropomyosin away from active sites
- Events of the cross bridge cycle occur
- When nervous stimulation ceases, Ca2+ is pumped back into the SR and contraction ends
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Cross Bridge Cycle
- Continues as long as the Ca2+ signal and adequate ATP are present
- Cross bridge formation—high-energy myosin head attaches to thin filament
- Working (power) stroke—myosin head pivots and pulls thin filament toward M line
- Cross bridge detachment—ATP attaches to myosin head and the cross bridge detaches
- “Cocking” of the myosin head—energy from hydrolysis of ATP cocks the myosin head into the high-energy state
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Review Principles of Muscle Mechanics
- 1.Contraction happens in single cells and in whole muscles
- 2.Contraction produces tension (force)
- 3.Contraction does not always shorten a muscle
- 4.Force and duration of contraction vary in response to stimuli of different frequencies and intensities
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ATP is regenerated by
ATP is regenerated by:
- Direct phosphorylation of ADP by creatine phosphate (CP)
- Anaerobic pathway (glycolysis)
- Aerobic respiration
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Muscle Cell Types Visually
- White = Fast glycolytic cells = short fast activity
- Red = Slow oxidative cells= long slow activity
- (Pink= Fast oxidative cells)= long and fast!
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