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Functions of Skeletal Muscle |
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Body movement Move bones, make facial expressions, speak, breathe, swallow Maintenance of posture Stabilize joints, maintain body position Protection and support Package internal organs and hold them in place Regulating elimination of materials Circular sphincters control passage of material at orifices Heat production Help maintain body temperature |
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Characteristics of Skeletal Muscle Tissue |
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Excitability: ability to respond to a stimulus by changing electrical membrane potential Conductivity: involves sending an electrical change down the length of the cell membrane Contractility: exhibited when filaments slide past each other Enables muscle to cause movement Elasticity: ability to return to original length following a lengthening or shortening Extensibility: ability to be stretched |
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Hierarchy of muscle structures |
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What is the hierarchy of structures in a muscle? A whole muscle contains many fascicles A fascicle consists of many muscle fibers A muscle fiber is a muscle cell
In addition to the muscle cells, a skeletal muscle contains nerves, blood vessels, and connective tissue [image] |
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Gross Anatomy of Skeletal Muscle |
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Connective tissue components Three concentric layers of wrapping Epimysium Dense irregular connective tissue wrapping whole muscle Perimysium Dense irregular connective tissue wrapping fascicle Houses many blood vessels and nerves Endomysium Areolar connective tissue wrapping individual fiber Delicate layer for electrical insulation, capillary support, binding of neighboring cells |
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Attachments of muscle to bone (or to skin or to another muscle) can be tendons or aponeuroses Tendon: cordlike structure of dense regular connective tissue Aponeurosis: thin, flattened sheet of dense irregular tissue |
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Sheet of dense irregular connective tissue Located external to epimysium Separates different muscles while binding them together Contains nerves, blood vessels, and lymph vessels |
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Areolar and adipose tissue Located superficial to deep fascia Separates muscles from skin |
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Skeletal muscle has extensive blood vessels Deliver oxygen and nutrients, removing waste products Skeletal muscle is innervated my somatic neurons Axons of neurons branch, terminate at neuromuscular junctions Can allow for voluntary control of contraction |
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Sarcoplasm (cytoplasm) Has typical organelles plus contractile proteins and other specializations Multiple nuclei (individual cells are multinucleated) Cell is formed in embryo when multiple myoblasts fuse Some nearby myoblasts become undifferentiated satellite cells for support and repair of muscle fibers Sarcolemma (plasma membrane) Has T-tubules (transverse tubules) that extend deep into the cell Sarcolemma and its T-tubules have voltage-gated ion channels that allow for conduction of electrical signals The interior of T-tubules is part of the extracellular environment |
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Development of Skeletal Muscle |
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Structure of Muscle Fiber |
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Myofibrils (hundreds to thousands per cell) Bundles of myofilaments (contractile proteins) enclosed by sarcoplasmic reticulum Make up most of the cell’s volume Sarcoplasmic reticulum Internal membrane complex similar to smooth endoplasmic reticulum Terminal cisternae: blind sacs of sarcoplasmic reticulum Serve as reservoirs for calcium ions Combine in twos with central T-tubule to form triads |
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Has a pump – sarcoplasmic endoplasmic reticulum calcium ATPase (SERCA) that import Ca2+ into sarcoplasmic reticulum where it binds to calmodulin and calquestrin Has channels (Ryanodine receptors) that allow Ca2+ to be released into surrounding sarcoplasm to trigger contraction |
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Thick filaments Consist of bundles of many myosin protein molecules Myosin has ATPase activity and when it hydrolyzes ATP, it is in a high energy state with a high affinity for actin Each myosin molecule has two heads and two intertwined tails Heads have binding site for actin of thin filaments and ATPase site Heads point toward ends of the filament Thin filaments Consist mostly of two twisted strands of fibrous actin (F-actin) Each strand is a necklace of hundreds of actin globules (G-actin) Each G-actin has a myosin binding site to which myosin heads attach during contraction Troponin and tropomyosin are regulatory proteins of thin filament Tropomyosin: twisted string-like protein covering the myosin-binding site on actin in a noncontracting muscle Troponin: globular protein attached to tropomyosin Troponin T binds tropomyosin, Troponin I binds actin and inhibits contraction, Troponin C binds Ca++ When Ca2+ binds to troponin it pulls tropomyosin off actin allowing contraction |
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Myofilaments arranged in repeating units, sarcomeres Composed of overlapping thick and thin filaments Delineated at both ends by Z discs Specialized proteins perpendicular to myofilaments Anchors for thin filaments The positions of thin and thick filaments give rise to alternating I-bands and A-bands [image] |
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Organization of Sarcomere |
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Definition
I bands Light-appearing regions that contain only thin filaments Bisected by Z disc Get smaller when muscle contracts (can disappear with maximal contraction) A band Dark-appearing region that contains thick filaments and overlapping thin filaments Contains H zone and M line Makes up central region of sarcomere H zone: central portion of A band Only thick filaments present; no thin filament overlap Disappears with maximal muscle contraction M line: middle of H zone Protein meshwork structure Attachment site for thick filaments [image] [image] |
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Connectin Extends from Z disc to M line Stabilizes thick filaments and has “springlike” properties (passive tension) Dystrophin Anchors some myofibrils to sarcolemma proteins A large protein that connects thin filaments to β-dystroglycan in the sarcolemma by syntrophins in complex with sarcoglycans It provides a scaffold for the myofibrils Abnormalities of this protein cause muscular dystrophy |
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Titin (largest known protein) anchors myosin and is important in skeletal muscle elasticity Actinin binds actin to the Z lines Desmin attaches the Z lines to the sarcolemma |
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Mitochondria and other structures associated with energy production |
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Muscle fibers have abundant mitochondria for aerobic ATP production Myoglobin within cells allows storage of oxygen used for aerobic ATP production Glycogen is stored for when fuel is needed quickly Creatinine phosphate can quickly give up its phosphate group to help replenish ATP supply |
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A motor neuron and all the fibers it controls. Axons of motor neurons from spinal cord (or brain) innervate numerous muscle fibers The number of fibers a neuron innervates varies Small motor units have less than five muscle fibers Allow for precise control of force output Large motor units have thousands of muscle fibers Allow for production of large amount of force (but not precise control) Fibers of a motor unit are dispersed throughout the muscle (not just in one clustered compartment) |
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Location where motor neuron innervates muscle Usually mid-region of muscle fiber Has synaptic knob, synaptic cleft, motor end plate[image] [image] |
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Expanded tip of the motor neuron axon Houses synaptic vesicles Small sacs filled with neurotransmitter acetylcholine (ACh) Has Ca2+ pumps in plasma membrane Establish calcium gradient, with more outside the neuron Has voltage-gated Ca2+ channels in membrane Ca2+ flows into cell (down concentration gradient) if channels open |
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Specialized region of sarcolemma with numerous folds Has many ACh receptors[image] Plasma membrane protein channels Opened by binding of ACh Allow Na+ entry and K+ exit |
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Narrow fluid-filled space Separates synaptic knob from motor end plate Acetylcholinesterase resides here Enzyme that breaks down ACh molecules |
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Resting Membrane Potential |
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Definition
Fluid inside cell is negative compared to fluid outside cell RMP of muscle cell is about –90 mV RMP set by leak channels and Na+/K+ pumps (voltage-gated channels are closed) |
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Skeletal Muscle Contraction |
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Definition
Calcium enters synaptic knob Nerve signal travels down axon, opens voltage-gated Ca2+ channels Ca2+ diffuses into synaptic knob Ca2+ binds to proteins on surface of [image]synaptic vesicles
1. Neuromuscular Junction: Excitation of Skeletal Muscle Fiber: Release of ACh from synaptic vesicles and subsequent binding of ACh to ACh receptors on motor end plate.Vesicles merge with cell membrane at synaptic knob: exocytosis Thousands of ACh molecules released from about 300 vesicles
2. Sarcolemma, T-tubules, and sarcoplasmic reticulum: excitation-contraction coupling: ACh binding triggers propagation of an action potential along the sarcolemma and T-tubules to the sarcoplasmic reticulum, which is stimulated to release Ca2+ 3. Sarcomere: Crossbridge Cycling: Ca2+ binding to troponin triggers sliding of thin filaments past thick filaments of sarcomeres. As sarcomeres shorten, the muscle contracts |
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Autoimmune disease, primarily in women Antibodies bind to ACh receptors in neuromuscular junctions Receptors removed from muscle fiber by endocytosis Results in decreased muscle stimulation Rapid fatigue and muscle weakness Eye and facial muscles often involved first May be followed by swallowing problems, limb weakness |
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End-Plate Potential (EPP) |
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Definition
ACh receptors are chemically gated channels that open when ACh binds to them Na+ diffuses into the cell through the channels (while a little K+ diffuses out) Cell membrane briefly becomes less negative at the end plate region EPP is local but it does lead to the opening of voltage-gated ion channels in the adjacent region of the sarcolemma |
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becomes less negative, eventually becomes +30 mV This results in the opening of adjacent voltage-gated Na+ channels and more Na+ entry A chain reaction occurs as depolarization is propagated down the membrane and T-tubules |
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K+ diffuses out of the cell |
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Definition
Cell repolarizes: returns to –90mV Repolarization is then propagated down the membrane and T-tubules While the cell is depolarizing and repolarizing it is in a refractory period—unable to respond to another stimulation |
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[image] Release of Ca2+ from the sarcoplasmic reticulum The spreading action potential causes a conformational change in the voltage-sensitive dihydropyridine (DHP)receptors on the sarcolemma to cause an unblocking of the RyR and Ca++ efflux from the sarcoplasmic reticulum (Action potential opens voltage-gated Ca2+ channels of sarcoplasmic reticulum) Ca2+ diffuses out of cisternae into sarcoplasm Ca2+ interacts with myofilaments triggering contraction |
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When Ca2+ binds to troponin C, it triggers crossbridge cycling Troponin and tropomyosin move so actin is exposed. Crossbridge formation Myosin head attaches to exposed binding site on actin Power stroke Myosin head pulls thin filament toward center of sarcomere ADP and Pi released 3) Release of myosin head ATP binds to myosin head causing its release from actin 4) Reset myosin head ATP split into ADP and Pi by myosin ATPase Provides energy to “cock” the myosin head
[image] Cycling continues as long as Ca2+ and ATP are present Results in sarcomere shortening as Z discs move closer together Narrowing (or disappearance) of H zone and I band Thick and thin filaments remain the same length but slide past each other |
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Spastic paralysis caused by toxin from Clostridium tetani Blocks release of inhibitory neurotransmitter in spinal cord, resulting in overstimulation of muscles Vaccination prevents this life-threatening condition |
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Muscular paralysis caused by toxin from Clostridium botulinum Prevents release of ACh at synaptic knobs Although toxin ingestion can be life-threatening, careful injections of it can treat spasticity (e.g., due to cerebral palsy) or can be used for cosmetic purposes (diminishing wrinkles) |
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Definition
Termination of nerve signal and ACh release from motor neuron Hydrolysis of ACh by acetylcholinesterase Closure of ACh receptor causes cessation of end plate potential No further action potential generation Closure of calcium channels in sarcoplasmic reticulum Return of Ca2+ to sarcoplasmic reticulum by pumps Return of troponin to original shape Return of tropomyosin blockade of actin’s myosin binding sites Return of muscle to original position due to its elasticity |
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Muscle cells have only a little ATP in storage Stored ATP is spent after about 5 seconds of intense exertion Three ways to generate ATP in skeletal muscle fiber Immediate supply via phosphate transfer Short-term supply via glycolysis Long-term supply via aerobic cellular respiration |
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Immediate supply of ATP: phosphate (Pi) transfer Myokinase transfers Pi from one ADP to another Creatine kinase transfers Pi from creatine phosphate to ADP |
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Does not require oxygen Glucose (from muscle’s glycogen or through blood) is converted to two pyruvate molecules 2 ATP released per glucose molecule Occurs in cytosol |
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Aerobic Cellular Respiration |
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Requires oxygen Occurs within mitochondria Pyruvate oxidized to carbon dioxide Transfer of chemical bond energy to NADH and FADH2 Energy used to generate ATP by oxidative phosphorylation Produces a net of 30 ATP Triglycerides can also be used as fuel to produce ATP More ATP from triglycerides with longer fatty acid chains |
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For a 50-meter sprint (less than 10 seconds) ATP supplied primarily by phosphate transfer system For a 400-meter sprint (less than a minute) ATP supplied primarily by glycolysis after first few seconds For a 1500-meter run (more than a minute) ATP supplied primarily by aerobic processes after first minute |
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Amount of additional oxygen needed after exercise to restore pre-exercise conditions Additional oxygen required to Replace oxygen on hemoglobin and myoglobin Replenish glycogen Replenish ATP and creatine phosphate Convert lactic acid back to glucose |
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Differences in power, speed, and duration Power related to diameter of muscle fiber (larger are more powerful) Speed and duration related to type of myosin ATPase (some catalyze faster), quickness of action potential propagation, and quickness of Ca2+ release and reuptake by sarcoplasmic reticulum Fast-twitch fibers are more powerful and have quicker and briefer contractions than slow twitch fibers. Speed of contraction determined by Vmax of myosin ATPase. High Vmax (fast, white) All fibers in a particular motor unit will be of the same type, i.e., fast or slow. |
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Oxidative vs Glycolytic Fibers |
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Oxidative fibers (fatigue-resistant) use aerobic cellular respiration Extensive capillaries Many mitochondria Large supply of myoglobin (red fibers) Smaller diameter Glycolytic fibers (fatigable) use anaerobic cellular respiration Fewer capillaries Fewer mitochondria Smaller supply of myoglobin (white fibers) Large glycogen reserves |
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3 types of Skeletal Muscle |
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1. Slow oxidative fibers (type I) Contractions are slower and less powerful High endurance since ATP supplied aerobically About half the diameter of other fibers, red in color due to myoglobin 2. Fast oxidative fibers (type IIa, intermediate) Contractions are fast and powerful Primarily aerobic respiration, but delivery of oxygen lower Intermediate size, light red in color 3. Fast glycolytic fibers (type IIb, fast anaerobic) Contractions are fast and powerful Contractions are brief, as ATP production is primarily anaerobic Largest size, white in color due to lack of myoglobin
Hand muscles have a high percentage of fast glycolytic fibers for quickness Back muscles have a high percentage of slow oxidative fibers to continually maintain postural support |
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Long-distance runners Higher proportion of slow-oxidative fibers in legs Sprinters Higher percentage of fast glycolytic fibers Determined primarily by genes Determined partially by training |
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Tension is the force generated when a muscle is stimulated to contract Lab experiments measure tension and graph it (myogram) |
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A twitch is a brief contraction to a single stimulus The minimum voltage that triggers a twitch is threshold stimulus Periods of the twitch Latent period Time after stimulus but before contraction begins No change in tension Contraction period Time when tension is increasing Begins as power strokes pull thin filaments Relaxation period Time when tension is decreasing to baseline Begins with release of crossbridges Generally lasts a little longer than contraction period |
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Recruitment is also called multiple motor unit summation It explains how muscles exhibit varying degrees of force Recruit just a few motor units to lift a pencil Recruit many motor units to lift a suitcase Above a certain voltage, all units are recruited, and so maximum contraction occurs (regardless of how much higher voltage is) |
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