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Skeletal Muscle Physiology Muscular System Functions • Body movement (Locomotion) • Maintenance of posture • Respiration – Diaphragm and intercostal contractions • Communication (Verbal and Facial) • Constriction of organs and vessels – Peristalsis of intestinal tract – Vasoconstriction of b.v. and other structures (pupils) • Heart beat • Production of body heat (Thermogenesis) Properties of Muscle • Excitability: capacity of muscle to respond to a stimulus • Contractility: ability of a muscle to shorten and generate pulling force • Extensibility: muscle can be stretched to its original length (and beyond to a certain degree) • Elasticity: ability of muscle to recoil to original resting length after stretched Types of Muscle • Skeletal – Attached to bones – Makes up 40% of body weight – Responsible for locomotion, facial expressions, posture, respiratory movements, other types of body movement – Voluntary in action; controlled by somatic motor neurons • Smooth – In the walls of hollow organs, blood vessels, eye, glands, uterus, skin – Some functions: propel urine, mix food in digestive tract, dilating/constricting pupils, regulating blood flow, – In some locations, autorhythmic – Controlled involuntarily by endocrine and autonomic nervous systems • Cardiac – Heart: major source of movement of blood – Autorhythmic – Controlled involuntarily by endocrine and autonomic nervous systems Muscle Tissue Types Connective Tissue Sheaths • Connective Tissue of a Muscle – Epimysium. Dense regular c.t. surrounding entire muscle • Separates muscle from surrounding tissues and organs • Connected to the deep fascia – Perimysium. Collagen and elastic fibers surrounding a group of muscle fibers called a fascicle • Contains b.v and nerves – Endomysium. Loose connective tissue that surrounds individual muscle fibers • Also contains b.v., nerves, and satellite cells (embryonic stem cells function in repair of muscle tissue • Collagen fibers of all 3 layers come together at each end of muscle to form a tendon or aponeurosis. Parts of a Muscle Z disc Z disc Microanatomy of Skeletal Muscle Muscle Fiber Anatomy • • Sarcolemma - cell membrane – Surrounds the sarcoplasm (cytoplasm of fiber) • Contains many of the same organelles seen in other cells • An abundance of the oxygen-binding protein myoglobin – Punctuated by openings called the transverse tubules (T-tubules) • Narrow tubes that extend into the sarcoplasm at right angles to the surface • Filled with extracellular fluid Myofibrils -cylindrical structures within muscle fiber – Are bundles of protein filaments (=myofilaments) • Two types of myofilaments 1. Actin filaments (thin filaments) 2. Myosin filaments (thick filaments) – At each end of the fiber, myofibrils are anchored to the inner surface of the sarcolemma – When myofibril shortens, muscle shortens (contracts) • Sarcomeres: Z Disk to Z Disk Sarcomere - repeating functional units of a myofibril – About 10,000 sarcomeres per myofibril, end to end – Each is about 2 µm long • Banded or striated appearance: – A bands: a dark band; full length of thick (myosin) filament – M line - protein to which myosins attach – H zone - thick but NO thin filaments – I bands: a light band; from Z disks to ends of thick filaments • Thin but NO thick filaments • Extends from A band of one sarcomere to A band of the next sarcomere – Z disc: filamentous network of protein. Serves as attachment for actin myofilaments – Titin filaments: elastic chains of amino acids; keep thick and thin filaments in proper alignment Sarcoplasmic Reticulum (SR) Sarcoplasmic Reticulum (SR) • SR is an elaborate, smooth endoplasmic reticulum – runs longitudinally and surrounds each myofibril – Form chambers called terminal cisternae on either side of the T-tubules • A single T-tubule and the 2 terminal cisternae form a triad • SR stores Ca++ when muscle not contracting – When stimulated, calcium released into sarcoplasm – SR membrane has Ca++ pumps that function to pump Ca++ out of the sarcoplasm back into the SR after contraction Structure of Actin and Myosin • Myosin (Thick) Myofilament • • • Many elongated myosin molecules shaped like golf clubs. Single filament contains roughly 300 myosin molecules Molecule consists of two heavy myosin molecules wound together to form a rod portion lying parallel to the myosin myofilament and two heads that extend laterally. Myosin heads 1. Can bind to active sites on the actin molecules to form cross-bridges. (Actin binding site) 2. Attached to the rod portion by a hinge region that can bend and straighten during contraction. 3. Have ATPase activity: activity that breaks down adenosine triphosphate (ATP), releasing energy. Part of the energy is used to bend the hinge region of the myosin molecule during contraction • • • • Thin Filament: composed of 3 major proteins 1. F (fibrous) actin 2. Tropomyosin 3. Troponin Two strands of fibrous (F) actin form a double helix extending the length of the myofilament; attached at either end at sarcomere. – Composed of G actin monomers each of which has a myosin-binding site (see yellow dot) – Actin site can bind myosin during muscle contraction. Tropomyosin: an elongated protein winds along the groove of the F actin double helix. Troponin is composed of three subunits: – Tn-A : binds to actin – Tn-T :binds to tropomyosin, – Tn-C :binds to calcium ions. Actin (Thin) Myofilaments Now, putting it all together to perform the function of muscle: Contraction Neuromuscular Junction Neuromuscular Junction • Region where the motor neuron stimulates the muscle fiber • The neuromuscular junction is formed by : 1. End of motor neuron axon (axon terminal) • Terminals have small membranous sacs (synaptic vesicles) that contain the neurotransmitter acetylcholine (ACh) 2. The motor end plate of a muscle • A specific part of the sarcolemma that contains ACh receptors • Though exceedingly close, axonal ends and muscle fibers are always separated by a space called the synaptic cleft Neuromuscular Junction Motor Unit All the muscle cells controlled by one nerve cell Muscle Contraction Summary 1. Nerve impulse reaches myoneural junction 2. Acetylcholine (Ach) is released from motor neuron (axon terminals) and crosses synaptic cleft. 3. Ach binds with receptors in the muscle membrane to allow sodium to enter synaptic cleft 4. Sodium influx will generate an action potential in the sarcolemma Muscle Contraction (Cont’d) 5. Action potential travels down T tubule and to sarcoplamic reticulum (SR) 6. SR releases calcium ions 7. Calcium binds with troponin to move the troponin/tropomyosin complex on the actin 8. Binding sites on the actin filament are exposed Muscle Contraction (cont’d) 9. ATP helps myosin head attach to binding sites (forming a crossbridge) and create a power stroke which moves actin, causing contraction 10. ATP detaches myosin heads and energizes them for another contraction 11. When action potentials cease the muscle stop contracting Sarcomere Relaxed Sarcomere Partially Contracted Sarcomere Completely Contracted Binding Site Troponin Ca2+ Tropomyosin Myosin Excitation-Contraction Coupling Muscle contraction •Alpha motor neurons release Ach •ACh produces large EPSP in muscle fibers (via nicotinic Ach receptors •EPSP evokes action potential •Action potential (excitation) triggers Ca2+ release, leads to fiber contraction •Relaxation, Ca2+ levels lowered by organelle reuptake Excitation-Contraction Coupling Excitation-Contraction Coupling Sliding Filament Model of Contraction • Thin filaments slide past the thick ones so that the actin and myosin filaments overlap to a greater degree • In the relaxed state, thin and thick filaments overlap only slightly • Upon stimulation, myosin heads bind to actin and sliding begins How striated muscle works: The Sliding Filament Model The lever movement drives displacement of the actin filament relative to the myosin head (~5 nm), and by deforming internal elastic structures, produces force (~5 pN). Thick and thin filaments interdigitate and “slide” relative to each other. Three Potential Actions During Muscle Contraction: • isotonic Biceps muscle shortens during contraction *shortening against fixed load = muscle building *No movement (but as tension increases muscles grow larger/stronger) • isometric • lengthening Biceps muscle lengthens during contraction *Most likely to cause muscle injury Developmental Aspects: Male and Female • These differences are due primarily to the male sex hormone testosterone • With more muscle mass, men are generally stronger than women – Women’s skeletal muscle makes up 36% of their body mass – Men’s skeletal muscle makes up 42% of their body mass • Body strength per unit muscle mass, however, is the same in both sexes Developmental Aspects: Age Related • With age, connective tissue increases and muscle fibers decrease • Muscles become stringier and more lean • By age 80, 50% of muscle mass is lost (sarcopenia) • Decreased density of capillaries in muscle • Reduced stamina • Increased recovery time • Regular exercise reverses sarcopenia