Chapter 10 Muscular Tissue Motion results from alternating contraction (shortening) and relaxation of muscles; the skeletal system provides leverage and a supportive framework for this movement. The scientific study of muscles is known as myology. Types of <a href="/keyword/muscle-tissue/" >muscle tissue</a> Skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> is primarily attached to bones. It is striated and voluntary. Cardiac <a href="/keyword/muscle-tissue/" >muscle tissue</a> forms the wall of the heart. It is striated and involuntary. Smooth (visceral) <a href="/keyword/muscle-tissue/" >muscle tissue</a> is located in viscera. It is nonstraited (smooth) and involuntary. Table 4.4 compares the different types of muscle. Functions of <a href="/keyword/muscle-tissue/" >muscle tissue</a> Through sustained contraction of alternating contraction and relaxation, muscle performs four key functions. These functions are production of body movements, stabilizing body positions, moving substances within the body, and generating heat. Properties of <a href="/keyword/muscle-tissue/" >muscle tissue</a> 1. Electrical excitability is the ability to respond to certain stimuli by producing electrical signals such as action potential (impulse). 2. Contractility is the ability to shorten and thicken (contract), generating force to do work. a. In an isometric contraction, the muscle develops tension but does not shorten. b. In an isotonic contraction, the tension remains constant while the muscle shortens. Properties of <a href="/keyword/muscle-tissue/" >muscle tissue</a> 3. Extensibility is the ability to be extended (stretched) without damaging the tissue. 4. Elasticity is the ability to return to original shape after contraction or extension. Skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> * Each skeletal muscle is a separate organ composed of cells called fibers. * Connective Tissue Components - Fascia is a sheet or band of fibrous connective tissue that is deep to the skin and surrounds muscles and other organs of the body. Skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> * Superficial fascia (or subcutaneous layer) separates muscle from skin and functions to provide a pathway for nerves and blood vessels, stores fat, insulates, and protects muscles from trauma. * Deep fascia, which lines the body wall and limbs and holds muscles with similar functions together, allows free movement of muscles, carries nerves, blood vessels, and lymph vessels, and fills spaces between muscles. Skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> 2. Other connective tissue components are epimysium, covering the entire muscle; perimysium, covering fasciculi; and endomysium, covering individual <a href="/keyword/muscle-fibers/" >muscle fibers</a> ; all are extensions of deep fascia Tendons &amp; Aponeuroses *Tendons and aponeuroses are extensions of connective tissue beyond muscle cells that attach muscle to bone or other muscle. * A tendon is a cord of dense connective tissue that attaches a muscle to the periosteum of a bone. Skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> An aponeurosis is a tendon that extends as a broad, flat layer. Nerve &amp; Blood Supply * Nerves (containing motor neurons) convey impulses for muscular contraction. * Blood provides nutrients and oxygen for contraction. Microscopic Anatomy of a Skeletal Muscle Fiber * During embryonic development, skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> arise from myoblasts. A few myoblasts persist in mature skeletal muscle as satellite cells. Microscopic Anatomy of a Skeletal Muscle Fiber * Sarcolemma, T Tubules, and Sarcoplasm - Skeletal muscle consists of fibers (cells) covered by a sarcolemma. - The fibers contain T tubules and sarcoplasm Microscopic Anatomy of a Skeletal Muscle Fiber The fibers contain T tubules and sarcoplasm 1) T tubules are tiny invaginations of the sarcolemma that quickly spread the muscle action potential to all parts of the muscle fiber. 2) Sarcoplasm is the muscle cell cytoplasm and contains a large amount of glycogen for energy production and myoglobin for oxygen storage. Microscopic Anatomy of a Skeletal Muscle Fiber Myofibrils and Sarcoplasmic Reticulum Each fiber contains myofibrils that consist of thin and thick filaments (myofilaments). The sarcoplasmic reticulum encircles each myofibril. It is similar to smooth endoplasmic reticulum in nonmuscle cells and in the relaxed muscle stores calcium ions. 3. Microscopic Anatomy of a Skeletal Muscle Fiber 4. Muscular atrophy is a wasting away of muscles, whereas muscular hypertrophy is an increase in the diameter of <a href="/keyword/muscle-fibers/" >muscle fibers</a> (Clinical Application). Sarcomeres Myofibrils are composed of thick and thin filaments arranged in units called sarcomeres. Sarcomeres Sarcomeres are the basic functional units of a myofibril and show distinct dark (A band) and light (I band) areas. -The darker middle portion is the A band consisting primarily of the thick filaments with some thin filaments overlapping the thick ones . Bands, Discs &amp; Zones -The lighter sides are the I bands that consist of thin filaments only. - A Z disc passes through the center of the I band. - A narrow H zone in the center of each A band contains thick but no thin filaments. Muscle Protein Contractile Proteins generate force during contraction. Myosin, the main component of thick filaments, functions as a motor protein. Motor proteins push or pull their cargo to achieve movement by converting energy from ATP into mechanical energy of motion or force. Muscle Protein Actin, the main component of thin filaments, connects to the myosin for the sliding together of the filaments. Muscle Protein Regulatory proteins help switch the contractions on and off. The regulatory proteins tropomyosin and troponin are a part of the thin filament. In relaxed muscle, tropomysium, which in held in place by troponin, blocks the myosin-binding sites on actin preventing myosin from binding to actin. Structural proteins keep the thick and thin filaments in the proper alignment, give the myofibril elasticity and extensibility, and link the myofibrils to the sarcolemma and extracellular matrix. Titin helps a sarcomere return to its resting length after a muscle has contracted or been stretched. Myomesin forms the M line. Nebulin helps maintain alignment of the thin filaments in the sarcomere. Dystrophin reinforces the sarcolemma and helps transmit the tension generated by the sarcomeres to the tendons. Muscle Contraction During muscle contraction, myosin cross bridges pull on thin filaments, causing them to slide inward toward the H zone; Z discs come toward each other and the sarcomere shortens, but the thick and thin filaments do not change in length. The sliding of filaments and shortening of sarcomeres causes the shortening of the whole muscle fiber and ultimately the entire muscle. This is called the sliding filament mechanism. Contraction Cycle The contraction cycle is a repeating sequence of events that causes the filaments to slide. It consists of ATP hydrolysis, attachment of myosin to actin to form cross bridges, the power stroke, and detachment of myosin from actin. Muscle Contraction An increase in calcium ion concentration in the cytosol starts muscle contraction; a decrease, stops it. The muscle action potential releases calcium ions from the sarcoplasmic reticulum that combine with troponin, causing it to pull on tropomyosin to change its orientation, thus exposing myosin-binding sites on actin and allowing the actin and myosin to bind together. Excitation-Contraction The use of calcium ions to remove the contraction inhibitor and the joining of actin and myosin constitute the excitation-contraction coupling, the steps that connect excitation (a muscle action potential propagation through the T tubules) to contraction of the muscle fiber. Calcium ion active transport pumps return calcium ions to the sarcoplasmic reticulum. Rigor mortis, a state of muscular rigidity following death, results from a lack of ATP to split myosin-actin cross bridges (Clinical Application). Muscle Contraction The forcefulness of muscle contraction depends on the length of the sarcomeres within a muscle before contraction begins. - plot of the lengthtension relationships for skeletal muscle. Neuromuscular Junction Muscle action potentials arise at the neuromuscular junction (NMJ), the synapse between a somatic motor neuron and a skeletal muscle fiber. Neuromuscular Junction A synapse is a region of communication between two neurons or a neuron and a target cell. - Synapses separate cells from direct physical contact. - Neurotransmitters bridge that gap. Neuromuscular Junction The neurotransmitter at a NMJ is acetylcholine (ACh). A nerve action potential elicits a muscle action potential through the release of acetylcholine, activation of ACh receptors, production of a muscle action potential, and termination of ACh activity Muscle Metabolism * Active muscle cells require large quantities of ATP. There are three sources for ATP production in muscle cells. * Creatine phosphate and ATP can power maximal muscle contraction for about 15 seconds and is used for maximal short bursts of energy (e.g., 100-meter dash) *Creatine phosphate is unique to <a href="/keyword/muscle-fibers/" >muscle fibers</a> . Muscle Metabolism * The partial catabolism of glucose to generate ATP occurs in anaerobic cellular respiration. This system can provide enough energy for about 30-40 seconds of maximal muscle activity (e.g., 300-meter race). Muscle Metabolism Muscular activity lasting more than 30 seconds depends increasingly on aerobic cellular respiration (reactions requiring oxygen). This system of ATP production involves the complete oxidation of glucose via cellular respiration (biological oxidation) Muscle Metabolism <a href="/keyword/muscle-tissue/" >muscle tissue</a> has two sources of oxygen: diffusion from blood and release by myoglobin inside <a href="/keyword/muscle-fibers/" >muscle fibers</a> . The aerobic system will provide enough ATP for prolonged activity so long as sufficient oxygen and nutrients are available. Muscle Metabolism * The inability of a muscle to maintain its strength of contraction or tension is called muscle fatigue; it occurs when a muscle cannot produce enough ATP to meet its needs. * Elevated oxygen use after exercise is called recovery oxygen consumption (rather than the formerly used term oxygen debt). Motor Neurons * When considering the contraction of a whole muscle, the tension it can generate depends on the number of fibers that are contracting in unison. * A motor neuron and the <a href="/keyword/muscle-fibers/" >muscle fibers</a> it stimulates form a motor unit. A single motor unit may innervate as few as 10 or as many as 2,000 <a href="/keyword/muscle-fibers/" >muscle fibers</a> , with an average of 150 fibers being innervated by each motor neuron. Twitch Contraction * A twitch contraction is a brief contraction of all the <a href="/keyword/muscle-fibers/" >muscle fibers</a> in a motor unit in response to a single action potential. * A record of a muscle contraction is called a myogram and includes three periods: latent, contraction, and relaxation. * The refractory period is the time when a muscle has temporarily lost excitability with skeletal muscles having a short refractory period and cardiac muscle having a long refractory period. Waves Wave summation is the increased strength of a contraction resulting from the application of a second stimulus before the muscle has completely relaxed after a previous stimulus * A sustained muscle contraction that permits partial relaxation between stimuli is called incomplete (unfused) tetanus; a sustained contraction that lacks even partial relaxation between stimuli is called complete (fused) tetanus. * The process of increasing the number of active motor units is called recruitment (multiple motor unit summation). - It prevents fatigue and helps provide smooth muscular contraction rather than a series of jerky movements. - Aerobic training builds endurance and anaerobic training builds muscle strength. (Clinical Application) F. A sustained partial contraction of portions of a relaxed skeletal muscle results in a firmness known as muscle tone. A any given moment, a few <a href="/keyword/muscle-fibers/" >muscle fibers</a> within a muscle are contracted while most are relaxed. This small amount of contraction is essential for maintaining posture. * Isotonic contractions occur when a constant load is moved through the range of motions possible at a joint and include concentric contractions and eccentric contractions; in an isometric contraction, the muscle does not shorten but tension increases - All skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> are not identical in structure or function -Color varies according to the content of myoglobin, an oxygen-storing reddish pigment. Red <a href="/keyword/muscle-fibers/" >muscle fibers</a> have a high myoglobin content while the myoglobin content of white <a href="/keyword/muscle-fibers/" >muscle fibers</a> is low. - Fiber diameter varies as do the cell's allocations of mitochondria, blood capillaries, and sarcoplasmic reticulum. - Contraction velocity and resistance to fatigue also differ between fibers. * On the basis of structure and function, skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> are classified as slow oxidative, oxidative-glycolytic, and fast glycolytic fibers. * Most skeletal muscles contain a mixture of all three fiber types, their proportions varying with the usual action of the muscle. All fibers of any one motor unit, however, are the same. * Although the number of different skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> does not change, the characteristics of those present can be altered by various types of exercise. * The use of anabolic steroids by athletes to increase muscle size, strength, and endurance has been shown to have very serious side effects, some of which are life-threatening. Cardiac <a href="/keyword/muscle-tissue/" >muscle tissue</a> Cardiac <a href="/keyword/muscle-tissue/" >muscle tissue</a> is found only in the heart wall. - Its fibers are arranged similarly to skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> . - Cardiac <a href="/keyword/muscle-fibers/" >muscle fibers</a> connect to adjacent fibers by intercalated discs which contain desmosomes and gap junctions Cardiac Muscle * Cardiac muscle contractions last longer than the skeletal muscle twitch due to the prolonged delivery of calcium ions from the sarcoplasmic reticulum and the extracellular fluid. * Cardiac <a href="/keyword/muscle-fibers/" >muscle fibers</a> contract when stimulated by their own autorhythmic fibers. * This continuous, rhythmic activity is a major physiological difference between cardiac and skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> . Smooth Muscle * Smooth <a href="/keyword/muscle-tissue/" >muscle tissue</a> is nonstriated and involuntary and is classified into two types: visceral (single unit) smooth muscle and multiunit smooth muscle. Smooth Muscle * Visceral (single unit) smooth muscle is found in the walls of hollow viscera and small blood vessels; the fibers are arranged in a network. * Multiunit smooth muscle is found in large blood vessels, large airways, arrector pili muscles, and the iris of the eye. The fibers operate singly rather than as a unit. Smooth Muscle * Sarcoplasm of smooth <a href="/keyword/muscle-fibers/" >muscle fibers</a> contains both thick and thin filaments which are not organized into sarcomeres. * Smooth <a href="/keyword/muscle-fibers/" >muscle fibers</a> contain intermediate filaments which are attached to dense bodies. Physiology of Smooth Muscle * The duration of contraction and relaxation of smooth muscle is longer than in skeletal muscle. * In smooth muscle, the regulator protein that binds calcium ions in the cytosol is calmodulin (in place of the role of troponin in striated muscle); calmodulin activates the enzyme myosin light chain kinase, which facilitates myosin-actin binding and allows contraction to occur at a relatively slow rate. Physiology of Smooth Muscle * The prolonged presence of calcium ions in the cytosol of smooth <a href="/keyword/muscle-fibers/" >muscle fibers</a> provides for smooth muscle tone, a state of continued partial contraction. * Smooth <a href="/keyword/muscle-fibers/" >muscle fibers</a> can stretch considerably without developing tension; this phenomenon is termed the stress-relaxation response. Regeneration of <a href="/keyword/muscle-tissue/" >muscle tissue</a> * Skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> cannot divide and have limited powers of regeneration; growth after the first year is due to enlargement of existing cells, rather than an increase in the number of fibers (although new individual cells may be derived from satellite cells). * The number of new skeletal <a href="/keyword/muscle-fibers/" >muscle fibers</a> formed from satellite cells is minimal. * Extensive repair results in fibrosis, the replacement of <a href="/keyword/muscle-fibers/" >muscle fibers</a> by scar tissue. * Cardiac <a href="/keyword/muscle-fibers/" >muscle fibers</a> cannot divide or regenerate. * Smooth <a href="/keyword/muscle-fibers/" >muscle fibers</a> have limited capacity for division and regeneration. Development of Muscle * With few exceptions, muscles develop from mesoderm * Skeletal muscles of the head and extremities develop from general mesoderm; the remainder of the skeletal muscles develop from mesoderm of somites. Aging &amp; <a href="/keyword/muscle-tissue/" >muscle tissue</a> * Beginning at about 30 years of age, there is a progressive loss of skeletal muscle, which is replaced by fat. * There is also a decrease in maximal strength and a slowing of muscle reflexes. Disorders * Neuromuscular disease involves problems involving somatic motor neurons, neuromuscular junctions, or <a href="/keyword/muscle-fibers/" >muscle fibers</a> . Myopathy signifies a disease or disorder of the skeletal <a href="/keyword/muscle-tissue/" >muscle tissue</a> itself. * Myasthenia gravis is an autoimmune disorder characterized by great muscular weakness and caused by antibodies directed against ACh receptors at the neuromuscular junction; more ACh receptors are affected as the disease progresses, making the muscle increasingly weaker. Disorders * Muscular dystrophies are inherited muscledestroying diseases that are characterized by degeneration of individual <a href="/keyword/muscle-fibers/" >muscle fibers</a> , leading to progressive atrophy of the skeletal muscle. The most common form is Duchenne muscular dystrophy, for which there is hope that gene therapy may someday be employed to replace the responsible gene with one that halts muscle loss. * Fibromyalgia is a painful, non articular rheumatic disorder that usually appears between the ages of 25 and 50. * Abnormal contractions of skeletal muscle include spasm, cramp, tremor, fasciculation, and fibrillation.