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Muscle Overview Function: to transform chemical energy (ATP) into mechanical energy The three types of muscle tissue are: Skeletal Cardiac smooth These types differ in structure, location, function, and means of activation PLAY InterActive Physiology ®: Anatomy Review: Skeletal Muscle Tissue, page 3 Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Muscle Similarities Skeletal and smooth muscle cells are elongated and are called muscle fibers Muscle contraction depends on two kinds of myofilaments – actin and myosin Muscle terminology is similar Sarcolemma – muscle plasma membrane Sarcoplasm – cytoplasm of a muscle cell Prefixes – myo, mys, and sarco all refer to muscle Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Skeletal Muscle Tissue Packaged in skeletal muscles that attach to and cover the bony skeleton Longest muscle cells Has obvious stripes called striations Is controlled voluntarily (i.e., by conscious control) Contracts rapidly but tires easily Is responsible for overall body motility Is extremely adaptable and can exert forces ranging from a fraction of an ounce to over several hundred pounds Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Cardiac Muscle Tissue Occurs only in the heart Is striated like skeletal muscle but is not voluntary Contracts at a fairly steady rate set by the heart’s pacemaker Neural controls allow the heart to respond to changes in bodily needs Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Smooth Muscle Tissue Found in the walls of hollow visceral organs, such as the stomach, urinary bladder, and respiratory passages Forces food and other substances through internal body channels It is not striated and is involuntary Contractions are slow and sustained Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Functional Characteristics of Muscle Tissue Muscles respond to neurotransmitter stimulus via nerve cells generating an electrical impulse that passes along the sarcolemma (plasma membrane) of the muscle cell causing it to contract Excitability, or irritability – the ability to receive and respond to stimuli Contractility – the ability to shorten forcibly Extensibility – the ability to be stretched or extended Elasticity – the ability to recoil and resume the original resting length Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Muscle Function i) Produce movement ii) Maintain posture iii) Stabilize joints iv) Generate heat v) Protection of viscera vi) act as valves Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Structure and Organization of Skeletal Muscle Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Table 9.1a Structure and Organization of Skeletal Muscle Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Table 9.1b Skeletal Muscle Each muscle is a discrete organ composed of muscle tissue, blood vessels, nerve fibers, and connective tissue Each muscle is served by: 1 nerve 1 artery 1 or more veins (much waste to be removed) With all 3 entering at the central region of the muscle, then branching outward Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Skeletal Muscle The three connective tissue sheaths are (internal to external): Endomysium – fine sheath of connective tissue composed of reticular fibers surrounding each muscle fiber Perimysium – fibrous connective tissue that surrounds groups of muscle fibers called fascicles Epimysium – an overcoat of dense regular connective tissue that surrounds the entire muscle. This may blend with the deep fascia that lies between neighboring muscles Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Skeletal Muscle The sheaths are continuous with one another and with the tendons Thus, when muscle fibers contract, they pull on the sheaths which transmit force to the bone They also contribute to the elasticity of muscles Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Skeletal Muscle PLAY InterActive Physiology ®: Anatomy Review: Skeletal Muscle Tissue, pages 4-6 Figure 9.2a Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Skeletal Muscle: Attachments Skeletal muscles span joints Attached to bones in at least 2 places Upon contraction, the muscle’s insertion (movable bone) moves towards the muscle’s origin (less movable bone) Direct attachment- epimysium (see slide 11 again) is fused to the perichondrium or periosteum Indirect attachment (more common)- tendon or aponeurosis anchors muscle to the connective tissue covering the skeletal element Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Microscopic Anatomy of a Skeletal Muscle Fiber Unit = muscle fiber = a skeletal muscle cell Multinucleated Plasma membrane = sarcolemma 10x bigger than the average body (epithelial) cell Up to 30 cm long (e.g. that is 12 inches!) Cytoplasm = sarcoplasm: contains high stores of glycosomes (glycogen granules) and myoglobin (an oxygen storing pigment) Contains modified organelles: i) Myofibrils ii) Sarcoplasmic reticulum & T-tubles Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Myofibrils Densly packed, 80% of cell volume Contain the contractile elements Striations are repeating series of A bands (dark) and I bands (light) The A band possesses an H zone which is bisected vertically by an M line I bands are interrupted by the Z disc (line) A, I, H, M, Z (what’s that now?) Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sarcomeres The smallest contractile unit of a muscle The region of a myofibril between two successive Z discs Composed of myofilaments made up of contractile proteins Contain: A band & ½ I band Myofilaments are of two types – thick and thin Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sarcomeres PLAY InterActive Physiology ®: Anatomy Review: Skeletal Muscle Tissue, page 9 Figure 9.3c Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Myofilaments: Banding Pattern Thick filaments (myosin) – extend the entire length of an A band Thin filaments (actin) – extend across the I band and partway into the A band. Anchored by Z disc Z-disc – coin-shaped sheet of proteins (connectins) that anchors the thin filaments and connects myofibrils to one another H band is less dense because actin (thin) filaments do not extend there M line in the center of H band is dense due to protein (desmin) strands that bind adjacent myosin filaments together Myofilaments are attached to the sarcolemma at the Z discs and the M lines Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Myofilaments: Banding Pattern Figure 9.3c,d Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Ultrastructure of Myofilaments: Thick Filaments Thick filament contains 300 myosin molecules The tails form the central part of the thick filament and consist of 2 interwoven, heavy polypeptide chains The heads face outward and in opposite directions and consist of 2 smaller, light polypeptide chains called cross bridges The central portion is smooth, and the ends are studded with myosin heads The heads possess actin binding sites & ATPase activity sites (the latter splits ATP to generate energy) Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Ultrastructure of Myofilaments: Thick Filaments Figure 9.4a,b Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Ultrastructure of Myofilaments: Thin Filaments Thin filaments are chiefly composed of the protein actin Each actin molecule is a helical polymer of globular subunits called G actin G actin monomers are polymerized into long actin filaments called fibrous (F) actin Each thin filament is formed from 2 intertwined actin filaments The subunits contain the active sites to which myosin heads attach during contraction Tropomyosin and troponin are regulatory subunits bound to actin Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Ultrastructure of Myofilaments: Thin Filaments Figure 9.4c Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Ultrastructure of Myofilaments: Thin Filaments Tropomyosin: 2 strands spiral about actin Help stiffen actin Block myosin binding to actin in the relaxed state Troponin: 3 polypeptide complex Tn1: inhibitory subunit bound to actin TnT: binds to tropomyosin and helps position it on actin TnC: binds calcium ions. Tn1 TnC TnT Figure 9.4c Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Arrangement of the Filaments in a Sarcomere Longitudinal section within one sarcomere Figure 9.4d Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Elastic filament Elastic filaments are composed of titin Extends from the Z disc to the thick filament and runs through the thick filament and is attached to the M line 2 functions: Holds the thick filament in place thus maintaining the A band Recoils the muscle cell after contraction terminates. Helps to resist excessive stretching Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Elastic filament Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sarcoplasmic Reticulum (SR) SR is an elaborate, smooth endoplasmic reticulum that mostly runs longitudinally and surrounds each myofibril Paired terminal cisternae form perpendicular cross channels at the A band – I band junction Functions in the regulation of intracellular calcium levels Stores Ca++ and releases it on demand during contraction (Ca++ is the final “Go” signal for contraction…are you thinking about the troponin subunit TnC…you should be!) Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sarcoplasmic Reticulum (SR) Elongated tubes called T (transverse) tubules penetrate into the cell’s interior at each A band–I band junction (where the sarcolemma penetrates into the cell interior Lumen of T-tubule is continuous with the extracellular space They conduct nerve impulses from the sarcolemma deep into the cell interior and to every sarcomere Impulses signal for the release of Ca++ from adjacent terminal cisternae Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sarcoplasmic Reticulum (SR) Figure 9.5 Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sliding Filament Model of Contraction During contraction the 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 at the A bands Upon stimulation (contraction), the myosin heads bind and “ratchet” actin toward the center of the sarcomere Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sliding Filament Model of Contraction Z discs are pulled toward the thick filaments I bands shorten Distance between Z bands is reduced H zone disappears A bands move closer together, but do not change length Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Sliding Filament Model of Contraction 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 As this event occurs throughout the sarcomeres, the muscle shortens Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Skeletal Muscle Contraction In order to contract, a skeletal muscle must: Be stimulated by a nerve ending Propagate an electrical current, or action potential, along its sarcolemma Have a rise in intracellular Ca2+ levels, the final trigger for contraction Linking the electrical signal to the contraction is excitation-contraction coupling Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings 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 Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Neuromuscular Junction The neuromuscular junction is formed from: Axonal endings, which have small membranous sacs (synaptic vesicles) that contain the neurotransmitter acetylcholine (ACh) The motor end plate of a muscle, which is a specific part of the sarcolemma that contains ACh receptors and helps form the neuromuscular junction Though exceedingly close, axonal ends and muscle fibers are always separated by a space called the synaptic cleft Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Neuromuscular Junction Figure 9.7 (a-c) Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings Neuromuscular Junction Events i) Nerve impulse reaches axon terminus ii) voltage-gated Ca++ channels in membrane open iii) Ca++ flows into axon from extracellular fluid iv) ACh is released into synaptic cleft after synaptic vesicles fuse w/ axonal membrane (exocytosis) v) ACh diffuses across cleft and attaches to receptors on sarcolemma vi) sarcolemma “fires” and creates an action potential vii) ACh splits into acetic acid and choline via acetylcholine esterase (located in the synaptic cleft). This destruction prevents continued muscle fiber contraction in the absence of additional stimuli Copyright © 2006 Pearson Education, Inc., publishing as Benjamin Cummings