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Muscle System th th 12 & 13 lectures February 11 & 18, 2016 Muscle tissue, one of the four major tissue types, plays the vital role of providing movement and heat generation to the organs of the body. There are 3 types of muscle tissue 1. Skeletal muscle = voluntary striated muscle 2. Cardiac muscle = involuntary striated muscle 3. Smooth muscle = involuntary non-striated muscle Characteristics of all muscle tissues: 1. Specialized cells: elongated, high density of myofilaments = cytoplasmic filaments of actin and myosin 2. Excitability/irritability: receive and respond to stimulus 3. Contractility: shorten and produce force upon stimulation 4. Extensibility: can be stretched 5. Elasticity: recoil after stretch Skeletal Muscle Tissue -forms skeletal muscles (44% of body mass) A skeletal muscle = an organ: composed of skeletal muscle cells (fibers), CT, nerves and blood vessels. Functions of skeletal muscles: 1. Produce skeletal movement 2. Maintain posture and upright position 3. Stabilize joints 4. Support soft tissues 5. Guard entrances and exits 6. Generate heat (maintain body temp) Skeletal Muscle Fibers -Huge cells: up to 100µm diameter, 30cm long -Multinucleate -Formed by fusion of 100 myoblasts-nuclei of each myoblast retained to provide enough mRNA for protein synthesis in large fiber -Unused myoblasts in adult = satellite cells -Satellite cells capable of division and fusion to fiber for repair but cannot generate new fibers -Cell membrane = sarcolemma Development of skeletal muscle. • Skeletal muscle begins to differentiate when mesenchymal cells called myoblasts align and fuse together to make longer, multinucleated tubes called myotubes. • Myotubes synthesize the proteins to make up myofilaments and gradually begin to show cross striations by light microscopy. • Myotubes continue differentiating to form functional myofilaments and the nuclei are displaced against the sarcolemma. • Part of the myoblast population does not fuse and differentiate, but remains as a group of mesenchymal cells called muscle satellite cells located on the external surface of muscle fibers inside the developing external lamina. • Satellite cells proliferate and produce new muscle fibers following muscle injury. Development of skeletal muscle Structure or Parts of Skeletal Muscle Endomysium –a delicate connective tissue sheath that encloses each skeletal muscle fiber. Perimysium – a coarse fibrous membrane that wraps the sheathed muscle fibers. Fascicle –bundle of fibers formed from group of sheathed muscle fibers wrapped by perimysium. Epimysium –a tough overcoat of connective tissue that bounds together fascicles. This is the connective tissue that covers the entire muscle. Tendons –these are cordlike structures that are formed from epimysia. These are composed of mostly collagenic fibers that can cross rough bony projections. Aside from anchoring muscles, tendons are very important in providing durability. Aponeuroses –these are sheet like structures that attaches muscles indirectly to bones, cartilages or connective tissue coverings of each other. Organization of skeletal muscle (a): Micrograph shows a cross section of striated muscle demonstrating connective tissue and cell nuclei. The endomysium around individual muscle fibers is indicated by arrowheads. At left is a portion of the epimysium. All three of these tissues contain collagen types I and III (reticulin). X200. H&E. Mechanism of Contraction • Resting sarcomeres consist of partially overlapping thick and thin filaments. • During contraction, neither the thick nor thin filaments changes their length. • Contraction is the result of an increase in the amount of overlap between the filaments caused by the sliding of thin and thick filaments past one another. • Contraction is induced by an action potential produced at a synapse, the neuromuscular junction, between the muscle fiber and a terminus of a motor axon. • Key molecular events in muscle contraction are summarized in the next slide Sliding Filament Theory -Contraction of skeletal muscle is due to thick filaments and thin filament sliding past each other, not compression of the filaments -evidence: 1. H-zones and I-bands decrease width during contraction 2. Zones of overlap increase 3. Z-lines move closer together 4. A-band remains constant -Sliding causes shortening of every sarcomere in every myofibril in every fiber -Overall result = shortening of whole skeletal muscle Skeletal muscle fibers of humans are classified into three types based on their physiological, biochemical, and histochemical characteristics All three fiber types are normally found throughout most muscles. §Type I or slow, red oxidative fibers contain many mitochondria and abundant myoglobin, a protein with iron groups that bind O2 and produce a dark red color. they are adapted for slow, continuous contractions over prolonged periods, as required for example in the postural muscles of the back. §Type IIa or fast, intermediate oxidative-glycolytic fibers have many mitochondria and much myoglobin, but also have considerable glycogen. They are adapted for rapid contractions and short bursts of activity, such as those required for athletics. §Type IIb or fast, white glycolytic fibers have fewer mitochondria and less myoglobin, but abundant glycogen, making them very pale in color. They are typically small muscles with a relatively large number of neuromuscular junctions, such as the muscles that move the eyes and digits. Cross-section of skeletal muscle stained histochemically to detect the density of myofibrillar myosin-ATPase can be used to demonstrate the distribution of slow (S) type I fibers, intermediate (I) type IIa fibers, and fast (F) type IIb fibers. Events of muscle contraction Cardiac Muscle Tissue -forms the majority of heart tissue cells =cardiocytes -one or two nuclei -no cell division -long branched cells -myofibrils organized into sarcomeres (striated) -aerobic respiration only -mitochondria and myoglobin rich -glycogen and lipid energy reserves -intercalated discs at cell junctions (gap junctions and desmosomes) allow transmission of action potentials & link myofibrils from one cell to next Features of cardiac muscle: 1. Can contract without neural stimulation; automaticity due to pacemaker cells that generate action potentials spontaneously 2. Pace and amount of tension can be adjusted by nervous system 3. Contractions 10X longer than skeletal muscle Diagram of cardiac muscle cells indicates characteristic features of this muscle type. The fibers consist of separate cells with interdigitating processes where they are held together. These regions of contact are called the intercalated discs, which cross an entire fiber between two cells. Other Heart Tissue Structures 1- Epicardium: CT layer that surrounds the heart 2- Purkinje Fibers: the purkinje fibers are found in the sub-endocardium. They are larger than cardiac muscle cells, but have fewer myofibrils, lots of glycogen and mitochondria. These cells are connected together by desmosomes and gap junctions, but not by intercalated discs. They are specialized conducting fibers, which is responsible for conduction of signal used for heart contraction; located at bottom (base) of heart. • IV. Smooth Muscle- A tissue formed by uninucleated spindle shaped cells found in six areas of the body: blood vessel walls, respiratory tract, digestive tubes, urinary organs, reproductive organs, and the eye. A. It exist as two layers with fibers running perpendicular to each other. One layer, the longitudinal layer is parallel to the axis, the circular layer is perpendicular. As they alternate contractions they shorten and constrict the organ. Main Charectristic of Smooth Muscle Tissue -lines hollow organs: regulates blood flow and movement of materials in organs -forms arrectorpili muscles -usually organized into two layers: Circular & longitudinal -spindle shaped cells -central nucleus -cells capable of division -no myofibrils, sarcomeres thick filaments scattered -thin filaments attached to dense bodies on desmin cytoskeleton (web) -adjacent cells attach at dense bodies with gap junctions (firm linkage and communication) -no tendons -contraction compresses whole cell B. There are no striations and no sarcomeres. Calcium ions signal contraction and the forces are not high. It can contract for a long time before fatiguing. Typically cells are not individually innervated and contraction may be signaled by stretching or hormones. Most molecules that allow contraction are similar in the three types of muscle, but the filaments of smooth muscle are arranged differently and appear less organized. The diagram shows that thin filaments attach to dense bodies located at the cell membrane and deep in the cytoplasm. Dense bodies contain a-actinin for thin filament attachment. Dense bodies at the membrane are also attachment sites for intermediate filaments and for adhesive junctions between cells. This arrangement of both the cytoskeleton and contractile apparatus allows the multicellular tissue to contract as a unit, providing better efficiency and force. (b) Micrograph showing a contracted (C) region of smooth muscle, with contraction decreasing the cell length and deforming the nuclei. The long nuclei of individual fibers assume a cork-screw shape when the fibers contract, reflecting the reduced cell length at contraction. Connective tissue (CT) of the perimysium outside the muscle fascicle is stained blue. X240. Mallory trichrome. Regeneration of Muscular Tissue • Hyperplasia - An increase in the number of fibers • Skeletal muscle has limited regenerative abilities - Growth of skeletal muscle after birth is due mainly to hypertrophy - Satellite cells divide slowly and fuse with existing fibers • Assist in muscle growth • Repair of damaged fibers • Cardiac muscle can undergo hypertrophy in response to increased workload - Many athletes have enlarged hearts • Smooth muscle in the uterus retain their capacity for division Copyright 2009, John Wiley & Sons, Inc. Aging and Muscular Tissue • Aging - Brings a progressive loss of skeletal muscle mass A decrease in maximal strength A slowing of muscle reflexes A loss of flexibility • With aging, the relative number of slow oxidative fibers appears to increase • Aerobic activities and strength training can slow the decline in muscular performance Copyright 2009, John Wiley & Sons, Inc. Muscle tissue throughout life 1. Mesoderm cells called myoblast fuse to form skeletal muscle tissues or join at cellular ends to form cardiac and smooth muscle. 2. Cardiac muscle contracts by the week 3 and skeletal muscle by week 7 of development! 3. Mitosis= skeletal muscle stops dividing once formed but has limited regenerated capacity in case of injury, cardiac muscle stops dividing by age 9, and smooth muscle divides as needed and has great regenerative abilitiy. 4. Muscle tissue is replaced with connective tissue as one ages. This is called sarcopenia and is reversible with exercise