Muscle Tissue. General concepts. Classification of muscle. I. Functional classification is based on the type of neural control.

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1 Muscle Tissue LEARNING OBJECTIVES 1. Identify the three types of muscle tissue at the light microscopic level. 2. List and compare the structural and functional features of each of the three muscle fiber types. 3. Name and describe the connective tissues present in muscle tissue 4. Describe the banding pattern of striated muscle and indicate the arrangement of myofilaments in a sarcomere. 5. Describe and compare the T-tubule system and its function in striated muscle General concepts Muscle tissue is specialized for the ability to shorten or contract. While all cells possess the cellular machinery necessary for shape change and contraction, these structures are significantly more prominent in muscle cells. For some muscle types, the cells are non-proliferative due to this high degree of specialization and differentiation. Muscle contraction is accomplished by the reciprocating sliding of intracellular filaments composed of actin and myosin. Classification of muscle I. Functional classification is based on the type of neural control. A. Voluntary B. Involuntary II. Structural classification is based on the presence or absence of cross-striations. A. Striated B. Non-striated (smooth) III. Combined functional and structural classification A. Skeletal muscle 1. Striated and voluntary 2. Found mostly attached to the skeleton. B. Cardiac muscle 1. Striated and involuntary

2 2. Composes the majority of the heart wall (myocardium) C. Smooth (visceral) muscle 1. Non-striated and involuntary 2. Found mostly in the walls of hollow organs and blood vessels Skeletal muscle I. Structure of skeletal muscle fibers A. Largest fiber type, fibers can be 1-30 mm in length and microns in diameter. B. Each muscle fiber is cylindrical, unbranched and multinucleated (syncytium) C. Each fiber has multiple nuclei located at the periphery of the muscle fiber immediately beneath the sarcolemma. D. Cytoplasm of the fibers is filled with myofibrils 1. Visible with the light microscope, 1-2 microns wide 2. Hundreds of myofibrils per cell and are oriented parallel to each other and to the long axis of the muscle fiber 3. Composed of bundles of overlapping myofilaments a. Visible only with the electron microscope b. Thin filament composed primarily of actin; Thick filaments composed primarily of myosin D. Extensive smooth endoplasmic reticulum is called the sarcoplasmic reticulum. E. Each fiber is surrounded by an external lamina (basal lamina) which functions to transmit the force of contraction. II. Banding pattern of skeletal (striated) muscle A. Fibers show prominent, alternating light and dark bands (cross striations) due to the alignment and overlap of the myofilaments within myofibrils. Myofilaments within a

3 myofibril are arranged in register and adjacent myofibrils are similarly aligned, causing the banding pattern seen at both the light and electron microscopic levels. B. Bands: 1. A band appears dark and contains both actin and myosin myofilaments. 2. I band appears light and contains actin myofilaments only. 3. Z line (Z disc) is located in the center of the I Band contains alpha-actinin and CapZ proteins. 4. H band is located in the center of the A band and contains myosin myofilaments only. C. Sarcomere 1. Contractile unit of striated muscle fibers, seen in both skeletal and cardiac muscle fibers 2. Extends from Z-line to Z-line 3. Sarcomeres are repeated in series along the length of each myofibril. 4. Alterations in sarcomeres during contraction a. Sarcomeres shorten b. Z-line to Z-line interval narrows c. Width of H and I bands decrease as actin myofilaments are pulled past the myosin myofilaments and area of myofilament overlap increases. d. A band width remains unchanged.

4 III. Regulation of skeletal muscle fiber contraction A. A complex system of intracellular, membranous structures called the triad insures coordinated contraction throughout the muscle fiber by (1) allowing the nervous impulse to penetrate and simultaneously reach all parts of the muscle fiber; and (2) releasing calcium in response to the nervous impulse. These functions are accomplished by the triad. B. Triads - composed of one T-tubule plus two adjacent terminal cisterns of the sarcoplasmic reticulum 1. T-tubules are invaginations of the sarcolemma 2. Terminal cisterns are expanded portions of the sarcoplasmic reticulum which lie adjacent to the T tubule and release calcium to initiate contraction. C. Role of triad in muscle contraction. 1. Nerve impulse arriving at the muscle fiber depolarizes the sarcolemma at the neuromuscular junction. 2. The membrane depolarization propagates along the sarcolemma and extends down the T-tubules. 3. T-tubule depolarization is transmitted to the terminal cisterns and the remainder of the sarcoplasmic reticulum, causing release of stored calcium. 4. Calcium initiates the interaction between actin and myosin myofilaments, leading to muscle contraction.

5 5. Calcium is recaptured by sarcoplasmic reticulum during relaxation. IV. Connective tissue investments of a skeletal muscle A. Function 1. Separate muscle into compartments 2. Transmit the force of contraction to insertion points B. Components 1. Endomysium - reticular fibers surrounding each muscle fiber plus the external (basal) lamina produced by the muscle fiber 2. Perimysium - dense connective tissue surrounding groups of fibers and dividing the muscle into fascicles 3. Epimysium - dense connective tissue surrounding the entire muscle, blends with the deep fascia and tendons

6 Cardiac muscle I. Cardiac muscle occurs only in the myocardium of the heart and, to a variable extent, in the roots of large vessels where they join the heart. II. Structure of cardiac muscle fibers A. Intermediate in size between skeletal and smooth muscle B. Fibers are cylindrical, branch, and form interwoven bundles. C. Single, centrally located nucleus D. Myofilament organization into myofibrils is identical to skeletal muscle. Cross-striations and bands identical to skeletal muscle are present, but not as prominent. E. Highly vascular with large numbers of mitochondria and glycogen accumulations, reflecting the high metabolic requirements of cardiac muscle fibers. F. Intercalated discs 1. Junctional complexes which are unique to cardiac muscle fibers. 2. Consist of specialized cell junctions and interdigitations of the sarcolemma at the ends of the fibers. 3. Contain three types of junctions a. Adherent junctions b. Gap junctions - provide ionic coupling between fibers

7 III. Regulation of cardiac muscle contraction A. Sarcomeres, myofibrils and myofilaments are the same as skeletal muscle fibers. B. T-tubules are present, however, the sarcoplasmic reticulum is not as well developed as in skeletal muscle fibers and does not form terminal cisterns. Thus, triads don t exist, however, T-tubules do stimulate release of calcium from the less well developed cisterns of sarcoplasmic reticulum that accompany them. Contraction is initiated by increased intracellular calcium. C. Contraction can spread through the myocardium due to the presence of gap junctions which allow current to flow from one fiber into another. Smooth Muscle I. Smooth muscle occurs mostly as sheets which form the walls of most hollow organs with the exception of the heart. Smooth muscle is also prominent in the walls of blood vessels, many respiratory passageways and some genital ducts.

8 II. Structure of smooth muscle fibers A. Smallest fiber type, length varies from 20 microns in blood vessels to 500 microns in the uterus B. Spindle shaped fibers - elongated with tapering ends and unbranched C. Possess a single, centrally placed, oval nucleus, which can appear spiraled or "inchworm"-shaped when the fiber is contracted. D. Non-striated and no myofibrils are present E. External lamina is present F. Abundant gap junctions G. Capable of both hypertrophy and hyperplasia; skeletal and cardiac muscle fibers can increase in size (hypertrophy) but they cannot proliferate (hyperplasia). III. Organization of the contractile proteins A. Actin and myosin myofilaments are present, but they are not organized into myofibrils. B. Myofilaments overlap as in striated muscle however they are not all oriented parallel to the long axis of the fiber. Rather, myofilaments crisscross throughout the sarcoplasm forming a reticulum. C. Dense bodies 1. Serve as insertion points for myofilaments to transmit the force of filament sliding 2. Contain alpha-actinin and, thus, resemble Z lines of striated muscle 3. Present in the cytoplasm and associated with the sarcolemma

9 IV. Coordination of smooth muscle contraction A. No T-tubules are present, however, fibers do have a rudimentary sarcoplasmic reticulum B. Sliding filament mechanism - regulated by intracellular release of calcium but with some differences from striated muscle fibers V. Types of smooth muscle A. Visceral smooth muscle 1. Occurs in sheets in the wall of hollow organs, e.g., digestive tract 2. Minimally innervated; contraction spreads in peristaltic waves facilitated by large numbers of gap junctions. 3. Specialized for slow, prolonged contraction B. Multiunit smooth muscle 1. Richly innervated 2. Specialized for precise, graded contraction, e.g., iris of the eye VCU Virtual Slide Collection Cardiac muscle: Cardiac muscle: Skeletal muscle: Smooth muscle: Smooth muscle: Cardiac Muscle Heart Skeletal Muscle Ileum Duodenum

10 Muscle tissue laboratory You are responsible for identifying the following structures in the Digital Histology CD package. While additional features may be mentioned in the legend or indicated on the images, you are not required to identify them at this time. Skeletal Muscle Connective tissue components of skeletal muscle Endomysium Epimysium Fascicle Perimysium Skeletal muscle fiber A band Cross striations I band Myofibrils Nucleus (peripheral, multiple) Sarcolemma Sarcomere Z-line Motor end plate Cardiac muscle Cardiac muscle fibers A band Cross striations Glycogen granules H band I band Intercalated disc M band Mitochondria Myofibrils Myofilaments Nucleus Sarcomere Z lines Smooth muscle Inner circular layer of smooth muscle Outer longitudinal layer of smooth muscle Smooth muscle fascicle Smooth muscle fibers Nucleus Sarcolemma Spindle shape