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Skeletal Muscle Muscle Tissue Dr. Patrick C. Nahirney Oct. 27, 2014 Island Medical Program, UVic Department of Cellular & Physiological Studies, UBC Objectives 1. Compare and contrast the 3 general types of muscle 2. Describe muscle fascicles, muscle fibers, myofibrils, myofilaments & sarcomeres in skeletal muscle 3. Describe epimysium, perimysium & endomysium 4. Relate arrangement of myofilaments, sarcoplasmic reticulum, T-tubules & triads to function in contraction 5. Outline myogenesis (muscle fiber development) 6. Describe neuromuscular junction and muscle spindle Images from Sections 4.2 & 4.3, Pages 73 & 74, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Muscle Tissue Classified into 3 categories based on structure, function & location • Skeletal Muscle: (Striated, Voluntary) - Attached to skeleton - 40% body wt. • Cardiac Muscle: (Striated, Involuntary) - In myocardium of heart • Smooth Muscle: (No striations, Involuntary) - In hollow tubes & viscera Images from Sections 4.3, 8.6 & 13.11, Pages 74, 179 & 296, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Skeletal Muscle 1° Function: Generate Force for Movement Skeletal muscle fibers: • Long cylindrical cells with tapered ends - 50-200 µm in diam and up to several cm long • Multinucleated with nuclei in peripheral position • Cytoplasm packed with myofibrils (cylindrical bundles of filaments) along length of fiber (highly eosinophilic) • 2 sets of contractile filaments containing actin & myosin that interdigitate and form cross striations Image from Section 4.2, Page 73, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Skeletal Muscle Design Rectus femoris muscle Muscle fiber (Myofiber) Nuclei Muscle fascicle Myofibril The 4 F’s: Fascicles Fibers Fibrils Myofilaments Filaments Images from Section 4.3, Page 74, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Non-pennate Skeletal Muscle Design Pennate Figure 2-5, Basic Human Anatomy, Copyright © 2009,p 74, O’Rahilly Ch et4, al. Netter's Essential Histology Skeletal Muscle - H&E Epimysium Muscle fascicle Perimysium Transverse section, Slide #27, UBC CPS Histology Virtual Slides Skeletal Muscle - H&E Muscle Muscle fascicle fibers Endomysium Nerve Blood fascicle vessel Perimysium Transverse section, Slide #27, UBC CPS Histology Virtual Slides Endomysium NaOH digested bovine skeletal muscle from Purslow, 2010 Skeletal Muscle - H&E Capillary Myonuclei Alternating light (Isotropic) and dark (Anisotropic) bands Muscle fiber Longitudinal section, Slide #28, UBC CPS Histology Virtual Slides Skeletal Muscle - EM Myofibrils, Z! Sarcomeres & Myofilaments Z! A band = dark I! I band = light A! Z line = thin I! A sarcomere is delineated by 2 consecutive Z lines TEM of skeletal muscle sectioned in the longitudinal plane (P.C. Nahirney, Personal Collection) Sarcomere I band A band I band Z H zone Z M • Thin (actin) (7nm x 1µm) interdigitate with thick (myosin) (15nm x 1.5µm) filaments - (sliding filament model) • During contraction the inward sliding of thin filaments along fixed thick filaments decreases distance between Z lines & shortens sarcomere TEM of sarcomere in skeletal muscle (UBC CPS Histology EM Magnifier) Muscle Contraction 1. Binding of calcium to troponin C exposes actin binding site on thin filament 2. In the presence of ATP, myosin head (which is energized from ATP) binds to actin 3. Binding of myosin head to actin initiates a powerstroke (bending of the myosin from Jeff Sales, SDSU http://www.sci.sdsu.edu/ head) and subsequent movies/actin_myosin.html release of ADP + Pi Rigor mortis: 4. Another molecule of ATP Lack of ATP prevents binds to myosin and myosin head from releases its attachment to releasing actin actin - ready for next stroke Animation courtesy of Jeff Sale, UCSD http://www.oercommons.org/courses/actin-myosin-crossbridge-animation/view Sarcoplasmic Reticulum (SR) • Analagous to smooth ER in other cells • Internal membrane system that regulates Ca levels - (Ca stimulates contraction within the muscle fiber) • Consists of flattened cisternae & anastamosing tubules • 2 pairs of terminal cisternae encircle each sarcomere at A-I junctions forming ‘muscle triads’ with T-tubules Transverse (T) Tubules • Tubular invaginations of sarcolemma that extend transversely into interior of muscle fiber • 1° role: conducts waves of depolarization rapidly into all parts of muscle fiber SR SR-T T-tubule Triad System The SR stores and releases I calcium in response to A sarcolemmal depolarization Sarcolemma Image from Section 4.4, Page 75, Ovalle & Myofibrils Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. HRSEM of SR-T System and Mitochondrion in Skeletal Muscle HRSEM of part of a Type IIA (intermediate) skeletal muscle fiber in the longitudinal plane (UBC CPS Histology EM Magnifier & in Section 4.13, Page 84, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved.) EM of Z Triads at A-I ! Band Mito! Junctions SR! DHPR! RyR1! T-T! I Band! Ca A band! Z! I band! ATP SR Lumen! Z! Modified from web.squ.edu TEM of part of a skeletal muscle fiber in longitudinal section (UBC CPS Histology EM Magnifier & in Section 4.7, Page 78, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved.) Myogenesis Myoblasts differentiate and align! Myoblasts fuse into myotubes! Myofibril synthesis & alignment Myofiber hypertrophy! Embryo at ~4 wks development • Somites contain myoblasts (muscle stem cells) • Undergo differentiation, fusion & hypertrophy • Some cells fail to fuse and become myosatellite cells Images from Section 4.2, Page 73, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Myogenesis of Skeletal Muscle in vitro Timelapse movie of myoblast fusion ~15 hr growth in culture Imaged at 5 min intervals Movie by P.C. Nahirney, Personal Collection Myogenesis of Skeletal Muscle in vitro Timelapse movie of myoblast fusion ~15 hr growth in culture Imaged at 5 min intervals Movie by P.C. Nahirney, Personal Collection Myosatellite Cells - EM • Mononucleated undifferentiated (stem) cells on surface of skeletal muscle fibers Myosatellite cell (only seen by EM or antibody labeling methods) • Normally quiescent - ‘resting myoblasts’ • Activated in injury Muscle fiber - undergo mitosis and provide limited muscle EM of the periphery of a skeletal muscle fiber in regeneration transverse section Image courtesy of Dr. W.K. Ovalle Neuromuscular Junction (NMJ) • Synapse between motor neurons of Motor spinal cord and axon muscle fibers Myelin Motor • A single motor axon sheath neuron cell innervates up to 100 body in spinal cord muscle fibers = Motor Schwann Unit cell • At NMJ, motor axon loses myelin sheath & ends in shallow depression on muscle sarcolemma • Axon terminal – covered externally by processes of Schwann cell Sarcolemma Image from Section 4.16, Page 87, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Neuromuscular Junction – con’t • Synaptic vesicles (40-60nm in diameter) Synaptic fill the axon terminal vesicles – contain Motor acetylcholine neuron cell • Synaptic cleft – Synaptic body in spinal cord narrow space cleft between axon terminal & sarcolemma of muscle fiber • Post-junctional infoldings of the sarcolemma increase surface area Post-junctional folds Image from Section 4.16, Page 87, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Neuromuscular Junction – EM • Nerve impulse causes vesicles Schwann to release Ach cell into synaptic cleft • Ach activates receptors on sarcolemma & depolarizes membrane NT NT • T-tubules carry signal to triads & activate ryanodine receptors (Ca Muscle fiber channels) of SR Image from Section 4.17, Page 88, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Muscle Spindles • Encapsulated sensory Intrafusal receptors (have outer & fibers inner capsules) Outer capsule • Contain specialized small diameter ‘intrafusal’ Inner muscle fibers enwrapped capsule by annulospiral sensory nerve endings (The bulk of muscle consists of ‘extrafusal’ fibers) • Integral component of the ‘monosynaptic Extrafusal fibers stretch reflex’ Skeletal muscle transverse section Slide #29, UBC CPS Histology Virtual Slides Sensory Innervation of Intrafusal Fibers • Nerve endings deform when muscle stretches ST IF! and initiate a train of ! action potentials • Signal travels up sensory neurons to synapse on dendrites of motor neurons in spinal cord (monosynaptic reflex) ST – sensory terminal IF – intrafusal fiber IC! IC – inner capsule of spindle! Images from Sections 20.22 & 20.23, Pages 477 & 488, Ovalle & Nahirney, Netter’s Essential Histology, 2nd Edition. Used with permission. Copyright © 2013 Elsevier Inc. All rights reserved. Muscle Pathology Normal Dystrophic Images by P.C. Nahirney, Personal Collection .
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