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組織學 Historlogy 台北醫學大學/解剖學科 教授:邱瑞珍 分機號碼:3261 電子郵件信箱:rueijen@tmu.edu.tw 1 Muscle Tissue 台北醫學大學/解剖學科 教授:邱瑞珍 分機號碼:3261 電子郵件信箱:rueijen@tmu.edu.tw 2 學習目的 • The morphological and functional characters of skeletal, cardiac and smooth muscles. 3 參考資料 • Junqueira's Basic Histology, twelfth edition, text and atlas, Anthony L. Mescher, McGraw-Hill Companies 4 Summary I. Skeletal muscle 1. nuclei: multi-nucleation, peripherally located 2. Myofibrils:myosin (thick) filaments, actin (thin) filaments 3. striation: dark band – A band (anisotropic); H band; M-line light band – I band (isotropic); Z-line sarcomere 4. Sarcoplasm :mitochondria, Golgi complex, glycogen, lipid droplets & myoglobin sarcoplasmic reticulum 5. Sarcolemma transverse (T) tubules – located at A-I junctiontriad→triad II. Cardiac muscle 1. nucleus: one or two, centrally located 2. myofibrils: similar to skeletal muscle 3. sarcoplasm: abundant, numerous mitochondria and lipid droplets, sarcoplasmic reticulum 4. sarcolemma: T- tubule located at Zline, intercalated disc – desmosome + gap junction Heart wall: epicardium, myocardium & endocardium Purkinje fiber: specialized cardiac muscle cells conduct impulses III. Smooth muscle •nucleus: one, centrally located •myofibrils: less orderly arranged, no striation, with dense body •sarcoplasm: various organelles, poor sarcoplasmic reticulum 4.sarcolemma: no T-tubule, caveolae or 5 vesicles (Ca2+), gap junction Muscle tissue: differentiated cell property: contractility microfilaments & associated proteins Æ generate forces drive movement within organs & body Nearly all muscle cells: from mesoderm differentiate by cell lengthening with synthesis of myofibrillar proteins 6 3 types of muscle tissue: morphologic & functional characteristics structure adapted to physiologic role Skeletal muscle: bundles of very long, cylindrical multinucleated cells show cross-striations contraction: quick, forceful, under voluntary control interaction of thin actin filament & thick myosin filament sliding 7 Cardiac muscle: also cross-striations elongated, branched cells lie parallel to each other end-to-end contact intercalated disks only in cardiac muscle contraction: involuntary, vigorous, rhythmic 8 Smooth muscle: collections of fusiform cells not show striations contraction: slow, not subject to voluntary control 9 Cytoplasm of muscle cells: Sarcoplasm Smooth ER: Sarcoplasmic reticulum Cell membrane: Sarcolemma 10 SKELETAL MUSCLE Consist of muscle fibers: long, cylindrical multinucleated cells diameter : 10-100µm Multinucleation: fusion of embryonic mesenchymal cells myoblasts Long oval nuclei: found at peripheral of cell under cell membrane 11 Organization Epimysium : an external sheath of dense connective tissue surrounding entire muscle Perimysium : surrounding fascicles (bundles) within muscle Endomysium : surrounded each muscle cell basal lamina 12 Connective tissue: transmit mechanical forces Blood vessels: penetrate the muscle within connective tissue form a rich capillary network in endomysium lymphatic vessels & larger blood vessels: other layers Muscle-tendon: myotendinous junctions 13 Muscle fibers Longitudinally sectioned: cross-striations: light & dark bands Dark bands : A bands Light bands : I bands Within I band: Z line Sarcomere: Z line to Z line 2.5 µm long in resting muscle 14 Sarcoplasm: little rER or free ribosome filled with long cylindrical filamentous bundles called myofibrils parallel to long axis of fiber Myofibrils : 1-2 µm consist of end to end sarcomeres pattern of transverse striations A & I banding pattern: regular arrangement of myofilaments thin & thick 15 Thick myofilament : 1.6μm long, 15nm wide occupy A band: central portion of sarcomere Thin myofilament: run between & parallel to thick filaments one end attached to Z line 1.0μm long, 8nm wide I band: portions of thin filaments not overlap the thick filaments A band: composed mainly thick overlapping portions of thin 16 A band: a lighter zone in center H zone only rod-like portions of myosin molecule no thin filaments Bisecting H zone: M line lateral connection of thick major protein: myomesin myosin-binding protein hold thick creatine kinase supply energy 17 Thin & thick: overlap within A band A cross section: each thick filament surrounded by 6 thin a hexagon 18 Thin filaments: composed of F-actin associated with tropomyosin long fine polymer also troponin globular complex of 3 subunits Thick filaments: consist primarily myosin Myosin + actin: 55% of total protein of striated muscle 19 F-actin: long filamentous polymers containing 2 strands of globular (G-actin) monomers 5.6 nm in diameter twisted around each other: double helical formation each G-actin: a binding site for myosin anchored on Z line: α-actinin 20 Each tropomyosin subunit: a long, thin molecule: 40nm long containing 2 polypeptide chains assemble to form a long polymer located in groove between actin strands Troponin: complex of 3 subunits TnT: attach to tropomyosin TnC: bind calcium ions TnI: inhibit actin-myosin interaction 21 Myosin : much larger complex: molecule weight ~500kDa 2 heavy chains, 2 pairs of light chains Heavy chain: thin, rod-like molecules 150 nm long, 2-3 nm thick made of 2 heavy chains twisted as myosin tails small globular projections: heads ATP binding site enzymatic capacity to hydrolysis ATP binding actin 22 4 light chains: associated with head Several hundred myosin molecules: arranged within each thick filament rodlike portions overlapping globular heads: directed toward either end Crossbridges: between thin & thick formed by head of thick & part of rod portion involved in conversion chemical energy Æ mechanical energy 23 Sarcoplasmic Reticulum & Transverse Tubule System In muscle: sER: specialized for Ca2+ ion sequestration depolarization Æ release Ca2+ Surface-initiated depolarization signals Æ diffusion to sER In large muscle cells: diffusion of depolarization Æ peripheral earlier than central Uniform contraction: transverse (T) tubules 24 T tubules : fingerlike invaginations of sarcolemma Form a complex network of tubules Encircle every myofibil Near A-I band boundaries of each sarcomere Adjacent to opposite side of each T tubule: expanded terminal cisternae of sarcoplasmic reticulum 2 cisternae + a T tubule : triad At triad: depolarization Æ sarcolemma Æ T tubules Æ sarcoplasmic reticulum 25 Mechanism of Contraction Resting sarcomeres: thick & thin partially overlapping During contraction: thick & thin not change their length Contraction : increase amount of overlap between filaments sliding of thin & thick induced by an action potential produced at synapse 26 27 If no ATP available: actin-myosin complex become stable rigor motis occur after death During contraction: I band decrease in size H band diminishes in width sarcomere, whole cell (fiber) greatly shorten 28 Muscle Fiber Types The most available forms of energy: ATP & phosphocreatine energy-rich phosphate compounds Chemical energy also stored in glycogen particles 0.5-1% muscle weight Muscle tissue obtains energy: Aerobic metabolism of fatty acids & glucose fatty acid Æ acetate Æ citric acid cycle Æ ATP Short-term exercise: Anaerobic metabolism of glucose (from glycogen) produce lactate oxygen debt 29 Skeletal muscles: 3 types physiological, biochemical, histochemical characteristics •Type I (slow, red oxidative) fibers contain many mitochondria abundant myoglobin a protein with iron groups bind O2, produce a dark red color energy from aerobic oxidative phosphorylation of fatty acids adapted for slow, continuous contraction over prolonged periods 30 •Type IIa (fast, intermediate oxidative-glycolytic) fibers many mitochondria much myoglobin, also glycogen both oxidative metabolism & anaerobic glycolysis intermediate between other types both in color & energy metabolism adapted for rapid contractions & short bursts of activity 31 •Type IIb (fast, white glycolytic) fibers fewer mitochondria less myoglobin abundant glycogen pale in color depend largely on glycolysis for energy fatigue quickly typically small muscles Differentiation of muscle Æ red, white, intermediate controlled by frequency of impulses from motor innervation Simple denervation of muscle: fiber atrophy & paralysis 32 CARDIAC MUSCLE During embryonic development: mesodermal cells align into chainlike arrays Cardiac muscle cells: complex junctions between processes Cells: branch & bind to cells in adjacent fibers Cardiac muscle cells: 15μm in diameter, 85-100 μm in length cross-striated banding pale-staining 1 or 2 nuclei centrally located endomysium: rich capillary network 33 Intercalated discs Interface between adjacent muscle cells many junctional complex Desmosomes, fasciae adherens: bind cardiac cells Gap junction: act as electrical synapses allow cells of cardiac muscle act as a multinucleated syncytium contraction signals passing in a wave from cell to cell 34 Contractile proteins: same as skeletal muscle T- tubules: more numerous & larger Sarcoplasmic reticulum: less well developed Contain numerous mitochondria: 40% or more of cytoplasmic volume need for continuous aerobic metabolism occupy 2% volume of skeletal muscle Fatty acids: major fuel of heart store as triglycerides Glycogen particles: also present Lipofuscin pigment granules: often found near the nuclei 35 SMOOTH MUSCLE Smooth muscle fibers: elongated, tapering, nonstriated cells each enclosed by a thin basal lamina & a fine network of reticular fibers Connective tissue: combine force generated by each fibers Æ a concerted action Smooth muscle cells: range in length: 20μm in small blood vessels ~500 μm in pregnant uterus single nucleus in center Contract: border scalloped nucleus distorted 36 A rudimentary sarcoplasmic reticulum: present No T tubules Thin & thick myofilaments : crisscross obliquely through the cell forming a latticelike network Sliding filament mechanism Intermediate filaments: desmin: major protein in all smooth muscle vimentin: additional component in vascular Both & F-actin insert into dense bodies 37 Dense body: membrane-associated or cytoplasmic contain α-actinin functionally similar to Z discs Attachment of thin & intermediate filaments: help transmit contractile force Æ other cells & surrounding network of reticular fibers Not under voluntary control Regulated by autonomic nerves, hormones, local conditions 1.Multiunit smooth muscle contract independently 2.Unitary smooth muscle gap junctions : 38 (X) contract independently In addition to contractile activity: smooth muscle cells also synthesize collagen, elastin, proteoglycans, ECM components normally synthesized by fibroblasts 39 REGENERATION OF MUSCLE TISSUE Skeletal muscles : (X) mitotic activity limited regeneration sparse mesenchymal satellite cells muscle injury or other stimuli Cardiac muscles: lack satellite cells: no regenerative capacity damage : fibrobalsts proliferation Æ myocardial scar Smooth muscle : more active regenerative response (V) mitotic activity 40