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HISTOLOGY BY DR. ZANA LECTURE -3MUSCULAR SYSTEM INTRODUCTION • Although all cells are capable of some sort of movement, the dominant function of several cell types is to generate motile forces through contraction. • In these specialized contractile cells, motile forces are generated by the interaction of the proteins actin and myosin (contractile proteins). PROPERTIES OF MUSCLE • Contractility • Ability of a muscle to shorten with force • Excitability • Capacity of muscle to respond to a stimulus • Extensibility • Muscle can be stretched to its normal resting length and beyond to a limited degree • Elasticity • Ability of muscle to recoil to original resting length after stretched 3 • Certain forms of contractile cell function as single-cell contractile units. • Myoepithelial cells are an important component of certain secretory glands where they function to expel secretions from glandular acini. • Pericytes are smooth muscle-like cells that surround blood vessels • Myofibroblasts are cells that have a contractile role in addition to being able to secrete collagen. This type of cell is generally inconspicuous in normal tissues but comes to be a dominant cell type when tissues undergo repair after damage in the formation of a scar • Layers CONNECTIVE • External lamina. Delicate, reticular fibers. Surrounds sarcolemma TISSUE • Endomysium. Loose C.T. with reticular fibers. • Perimysium. Denser C.T. surrounding a group of muscle fibers. Each group called a fasciculus • Epimysium. C.T. that surrounds a whole muscle (many fascicles) 9-5 • Fascia: connective tissue sheet • Forms layer under the skin • Holds muscles together and separates them into functional groups. • Allows free movements of muscles. • Carries nerves (motor neurons, sensory neurons), blood vessels, and lymphatics. • Continuous with connective tissue of tendons and periosteum. • Other forms of contractile cell function by forming multicellular contractile units termed muscles. Such muscle cells can be divided into three types: 1. Skeletal muscle. 2. Smooth muscle. 3. Cardiac muscle. SKELETAL MUSCLE • responsible for the movement of the skeleton and organs such as the globe of the eye and the tongue. Skeletal muscle is often referred to as voluntary muscle since it is capable of voluntary (conscious) control. • The arrangement of the contractile proteins gives rise to the appearance of prominent cross-striations in some histological preparations and so the name striated muscle is often applied to skeletal muscle. • The highly developed functions of the cytoplasmic organelles of muscle cells has led to the use of a special terminology for some muscle cell components: plasma membrane or plasmalemma = sarcolemma; cytoplasm = sarcoplasm; endoplasmic reticulum = sarcoplasmic reticulum. • Skeletal muscles have a wide variety of morphological forms and modes of action; nevertheless all have the same basic structure. • Skeletal muscle is composed of extremely elongated, multinucleate contractile cells, often described as muscle fibers, bound together by collagenous supporting tissue. • Individual muscle fibers vary considerably in diameter from 10 to 100 μm and may extend throughout the whole length of a muscle reaching up to 35 cm in length. Nuclei. Due to their syncytial development, skeletal myocytes are polyploid cells, which contain multiple ovoid nuclei. In mammals, the nuclei are typically located in the peripheral sarcoplasm just beneath the sarcolemma. This characteristic distinguishes skeletal muscle cells from cardiac muscle cells , which usually have one centrally- located nucleus. STRIATIONS. • Composed of alternating dark and light bands, the repetitive striations seen in myocytes result from the vertical summation of smaller bands that occur in myofibrils. • Myofibrils are the smallest structural and functional subunits of myocytes seen at the LM level. They are arranged in register within the sarcoplasm, parallel to the long axis of the muscle fiber. Dark bands are called A bands because they are anisotropic in polarized light (i.e., they rotate plane polarized light). The parallel array of thick filaments are mainly responsible for the LM appearance of the A band. In the center of this band a paler region, called the H band [formerly Hensen disk], can be seen in relaxed muscle. It represents an area of thick filaments that are not overlapped by thin filaments. At the middle of the A band (and the H band) is a middle line called the M line. Light bands are called I bands because they are isotropic in polarized light (i.e., they do not rotate plane polarized light). A dark transverse line, called the Z line (Z disk) bisects each I band. • Thin filaments are composed of several proteins, but primarily of actin and two important regulatory proteins called tropomyosin and troponin. • Thick filaments are composed primarily of composites of the protein myosin II. Sarcomeres are defined as the basic units of contraction in striated muscles. A single sarcomere extends from one Z line to the next and contains one A band separating two semi-I bands. Thus, myofibrils are composed of a series of tandomly-arranged sarcomeres consisting of interdigitating polarized thin filaments (plus end toward Z line; minus end toward A band) and bipolar thick filaments (myosin heads toward each semi-I band). Arrangement of filaments in sarcomeres. Thick filaments occupy central portions of the sarcomere; thin filaments attach at each end to the Z lines and run parallel to, and between, the thick filaments. i. I bands are composed of thin filaments. Each sarcomere has ½ of an I band at its ends. Thus, a whole I band is shared between adjacent sarcomeres. ii. A bands are composed mostly of thick filaments and the thin filaments between them. iii. H bands are composed of only thick filaments, and in relaxed muscle represent the area between the ends of thin filaments that are attached to each Z line at the other ends. In contracted muscle, when the thin filaments merge or overlap, the H band disappears. iv. Actin and myosin together represent approximately 55% of the total proteins in striated muscle. Electron microscopic appearance. In electron micrographs, the repeating pattern of bands and sarcomeres is due to the arrangement of the myofilaments. Myofilaments represent the thick myosin filaments and the thin actin filaments. These structures are not resolved at the LM level. • Skeletal muscle contraction is controlled by large motor nerves, individual nerve fibers branching within the muscle to supply a group of muscle fibers, collectively described as a motor unit. • Excitation of any one motor nerve results in simultaneous contraction of all the muscle fibers of the corresponding motor unit. • The structure of neuromuscular junctions is described in The vitality of skeletal muscle fibers is dependent on the maintenance of their nerve supply which, if damaged, results in atrophy of the fibers. • Skeletal muscle contains highly specialized stretch receptors known as neuromuscular spindles • The individual muscle cells (muscle fibers) are grouped together into elongated bundles called fasciculi with delicate supporting tissue called endomysium occupying the spaces between individual muscle fibers. • Each fascicle is surrounded by loose collagenous tissue called perimysium. Most muscles are made up of many fasciculi and the whole muscle mass is invested in a dense collagenous sheath called the epimysium. Large blood vessels and nerves enter the epimysium and divide to ramify throughout the muscle in the perimysium and endomysium. • The size of the fasciculi reflects the function of the particular muscle concerned. Muscles responsible for fine, highly controlled movements, e.g. the external muscles of the eye, have small fasciculi and a relatively greater proportion of perimysial supporting tissue. In contrast, muscles responsible for gross movements only, e.g. the muscle of the buttocks, have large fasciculi and relatively little perimysial tissue. • Muscle fibers are anchored to the support tissue so that contractile forces can be transmitted. The connective tissue framework contains both collagen and elastic fibers. This connective tissue becomes continuous with that of the tendons and muscle attachments which distribute and direct the motive forces of the muscle to bone, skin etc. SKELETAL MUSCLE EMBRYOGENESIS • During embryological development, mesenchymal cells in each myotome differentiate into long, mononuclear skeletal muscle precursors called myoblasts which then proliferate by mitosis. Subsequently, the myoblasts fuse end to end forming elongated multinucleate cells called myotubes. • Mature muscle cells can regenerate if damaged, by proliferation of stem cells which remain in adult muscles. These muscle stem cells resemble myoblasts and are called satellite cells. They enter mitosis after muscle damage and several fuse to form differentiated muscle fibers. Muscle fibers which are the result of regeneration after damage have nuclei in the center of the fiber rather than at the periphery. SMOOTH MUSCLE • Smooth muscle is so named because, unlike other forms of muscle, the arrangement of contractile proteins does not give the histological appearance of cross-striations. This type of muscle forms the muscular component of visceral structures such as blood vessels, the gastrointestinal tract, the uterus and the urinary bladder, giving rise to the alternative name of visceral muscle. Since smooth muscle is under inherent autonomic and hormonal control, it is also described as involuntary muscle. • In contrast to skeletal muscle, which is specialized for relatively forceful contractions of short duration and under fine voluntary control, smooth muscle is specialized for continuous contractions of relatively low force, producing diffuse movements resulting in contraction of the whole muscle mass rather than contraction of individual motor units. • Contractility is an inherent property of smooth muscle, occurring independently of neurological innervation often in a rhythmic or wave-like fashion. Superimposed on this inherent contractility are the influences of the autonomic nervous system, hormones and local metabolites which modulate contractility to accommodate changing functional demands. • For example, the smooth muscle of the intestinal wall undergoes continuous rhythmic contractions which result in waves of constriction passing along the bowel, propelling the luminal contents distally. • This activity is enhanced by parasympathetic stimulation and influenced by a variety of hormones released in response to changes in the nature and volume of the gut contents. • The cells of smooth muscle are relatively small with only a single nucleus. The fibers are bound together in irregular branching fasciculi, the arrangement varying considerably from one organ to another according to functional requirements. • smooth muscle fibers are elongated, spindle-shaped cells with tapered ends which may occasionally be bifurcated. Smooth muscle fibers are generally much shorter than skeletal muscle fibers and contain only one nucleus which is elongated and centrally located in the cytoplasm at the widest part of the cell; however, depending on the contractile state of the fibers at fixation, the nuclei may sometimes appear to be spiral-shaped. • Smooth muscle fibers are bound together in irregular, branching fasciculi and these fasciculi, rather than individual fibers, are the functional contractile units. • Within the fasciculi, individual muscle fibers are arranged roughly parallel to one another with the thickest part of one cell lying against the thin parts of adjacent cells. • The contractile proteins of smooth muscle are not arranged in myofibrils as in skeletal and cardiac muscle, and thus visceral muscle cells are not striated. CARDIAC MUSCLE • Cardiac muscle has many structural and functional characteristics intermediate between those of skeletal and smooth muscle and provides for the continuous, rhythmic contractility of the heart. Although striated in appearance, cardiac muscle is readily distinguishable from skeletal muscle and should not be referred to by the term 'striated muscle' • Cardiac muscle or myocardium exhibits many structural and functional characteristics intermediate between those of skeletal and visceral muscle. • Like the former, its contractions are strong and utilize a great deal of energy, and like the latter the contractions are continuous and initiated by inherent mechanisms, although they are modulated by external autonomic and hormonal stimuli. • Cardiac muscle fibers are essentially long cylindrical cells with one or at most two nuclei, centrally located within the cell. The ends of the fibers are split longitudinally into a small number of branches, the ends of which abut onto similar branches of adjacent cells giving the impression of a continuous three-dimensional cytoplasmic network; this was formerly described as a syncytium before the discrete intercellular boundaries were recognized • Between the muscle fibers, delicate collagenous tissue analogous to the endomysium of skeletal muscle supports the extremely rich capillary network necessary to meet the high metabolic demands of strong continuous activity. • Cardiac muscle fibers have an arrangement of contractile proteins similar to that of skeletal muscle and are consequently striated in a similar manner. However, this is often difficult to see with light microscopy due to the irregular branching shape of the cells and their myofibrils • Cardiac muscle fibers also have a system of T tubules and sarcoplasmic reticulum analogous to that of skeletal muscles. In the case of cardiac muscle, however, there is a slow leak of calcium ions into the cytoplasm from the sarcoplasmic reticulum after recovery from the preceding contraction; this causes a succession of automatic contractions independent of external stimuli. The rate of this inherent rhythm is then modulated by external autonomic and hormonal stimuli. • Between the ends of adjacent cardiac muscle cells are specialized intercellular junctions, called intercalated discs, which not only provide points of anchorage for the myofibrils but also permit extremely rapid spread of contractile stimuli from one cell to another. • Thus, adjacent fibers are triggered to contract almost simultaneously, thereby acting as a functional syncytium. In addition, a system of highly modified cardiac muscle cells constitutes the pacemaker regions of the heart and ramifies throughout the organ as the Purkinje system, thus coordinating contraction of the myocardium as a whole in each cardiac cycle. • Cardiac muscle cells in certain locations in the heart are responsible for secreting hormones into the bloodstream.