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    Muscular System Reading

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    cones and joints form the framework of the body, but they cannot move Its from the contraction and relax. iy themselves. Motion results ea py ieles, Muscle tissue constitutes about 40 t0 50 percent of srovtoal body weight and is composed of highly specialized cell The scientific study of muscles is known as myology (mFOL-O-jé; myo = muscle; logos = study of). CHARACTERISTICS Muscle tissue has four principal characteristics that are i its functions: important in understanding itabilit i ive and respond to stimuli 1. Excitability is the ability of muscle tissue to receive an 2. Contractility is the ability to shorten and thicken, or contract. 3. Extensibility is the ability of muscle tissue to stretch or extend. 4, Elasticity is the ability of muscle tissue to return to its original shape after contraction or extension. a FUNCTIONS ‘Through contraction, muscle performs three important functions: 1. Motion. 2. Maintenance of posture. 3. Heat production. Motion is obvious in movements involving the whole body, such as walking and running, and in localized movements, such as grasping a pencil or nodding the head. Less noticeable kinds of motion produced by muscles are the beating of the heart, chumning of food in the stomach, and contraction of the urinary bladder to expel urine. ‘Muscle tissue also enables the body to maintain posture. The contraction of skeletal muscles holds the body in stationary positions, such as standing and sitting. The third function of muscle tissue is heat production. Skeletal muscle contrac- tions produce heat and thereby help maintain normal body temperature. As much as 85 percent of all body heat is generated by muscle contractions. TYPES There are three types of muscle tissue. Skeletal muscle tissue, which is named for its location, is attached to bones and moves the skeleton. It is striated; that is, striations, of alternating light and dark bands, are visible under a microscope. It is voluntary because it can be made to contract and relax by conscious control. Cardiac muscle tissue forms the bulk of the wall of the heart. It is striated and involuntary; that is, its contractions are not under conscious control ‘Smooth muscle tissue is involved with internal processes. It is located in the walls of hollow internal structures, such as blood vessels, the stomach, and the intestines. It is nonstriated, because it lacks striations, and involuntary SKELETAL MUSCLE TISSUE To understand how muscles move, you need some knowledge of their connective tissue coverings, nerve and blood supply, and the structure of individual muscle fibers (cells). CONNECTIVE TISSUE COMPONENTS The term fascia (FASH-€-a) is applied to a sheet or broad band of fibrous connective tissue beneath the skin or around muscles and other organs of the body. There are two types of fascia. superficial fascia, immediately under the skin, and more important tothe study of muscles, deep fascia, which holds muscles together separating them into functioning groups. Functionally, deep fascia allows free movement of muscles, carries nerves and blood and lymphatic vessels, fills spaces between muscles, and sometimes provides the origin (one point of attachment to a bone) for muscles Skeletal muscles are further protected, strengthened, and at- tached to other structures by several other connective tissue cover- ings (Figure 8-1). The entire muscle is wrapped in fibrous connective tissue called the epimysium (ep’-i-MIZ-8-um), which is deep fascia Bundles of fibers (cells) called fasciculi (fa-SIK-yoo-Ii) or fascicles (FAS-i-kuls) are covered by a fibrous connective tissue called the perimysium (per'-i-MIZ-é-um), Finally, a fibrous connective tissue called the endomysium (en'-d6-MIZ-¢-um) wraps each individual muscle fiber (cell). Epimysium, perimysium, and endomysium extend beyond the muscle as a tendon (1endere = to stretch out)—a cord of connective Skeletal Muscle Tissue * 143 tissue that attaches a muscle to a bone. When a tendon extends as a broad, flat layer, itis called an aponeurosis. An example of ‘an aponeurosis is the galea aponeurotica (see Figure 8-9). Certain tendons, especially those of the wrist and ankle, are enclosed in tubes of fibrous connective tissue called tendon sheaths, which Permit tendons to slide easily and prevent them from slipping out of place. NERVE AND BLOOD SUPPLY ‘Skeletal muscles are well supplied with nerves and blood vessels, both of which are directly related to contraction, the chief character- istic of muscle. For a skeletal muscle fiber (cell) to contract, it ‘must first be stimulated by an electric current called an action potential. An action potential that is associated with a nerve cell is called a nerve action potential or nerve impulse. One associated with a muscle cell is called a muscle action potential. Muscle contraction also requires good deal of energy and therefore large amounts of nutrients and oxygen. Moreover, the waste products ‘of these energy-producing reactions must be eliminated. Thus, FIGURE 8-1 Relationships of connective fssve to skeletal muscle depicted by a three-dimen- sional drawing of a muscle indicating the relative positions of the epimysium, perimysium, ‘ond endomysivm. EpimysiuM (cep tas) PeRMYSIUM ewoomysium: Myotis (ruse tber real) Sarcolemma loos capillary Fascielus PERIMSIUM ee a See ee na 144 » Chopter 8 The Muscular System prolonged muscle action depends on a rich blood supply to deliver nutrients and oxygen and remove wastes. Generally, an artery and one or two veins accompany each nerve that penetrates a skeletal muscle, Microscopic blood vessels called capillaries ae distributed within the endomysium. Each mus- cle fiber is thus in close contact with one or more capillaries. Each skeletal muscle fiber also makes contact with a portion of nerve called a synaptic end bulb. HISTOLOGY Ifyou tease apart skeletal muscle and view it under a microscope, ‘you will ee it consists of thousands of elongated, cylindrical cells called muscle fibers or myofibers (see Exhibit 4-3). Each fiber is covered by a plasma membrane called the sarcolemma (sarco flesh; lemma = sheath). The cytoplasm, called sarcoplasm, of a ‘muscle fiber contains many nuclei (multinucleate). Inthe sarcoplasm fare numerous mitochondria. The large numbers are related to the large amounts of energy (ATP) that muscle tissue must generate in order to contract. The sarcoplasm also contains myofibrils (de- scribed shortly), special high-energy molecules (also tobe described short), enzymes, and sarcoplasmic reticulum (sar’-k0-PLAZ-mik re-TIK-yoo-lum), a network of membrane-enclosed tubules compa- rable to smooth endoplasmic reticulum (Figure 8-2a). Perpendic- tular to the sarcoplasmic reticulum are transverse tubules (T tu- ules). The tubules are tunnellke extensions of the sarcolemma that passthrough the muscle fiber a ight angles tothe sarcoplasmic reticulum and also open to the exterior of the muscle fiber. Skeletal muscle fibers also contain myoglobin, a reddish pigment similar to hemoglobin in blood. Myoglobin stores oxygen until needed by mitochondria to generate ATP. Each skeletal muscle fiber is composed of cylindrical structures called myofibrils which run longitudinally through the muscle fiber. They, in tum, consist of two kinds of even smaller structures called thin myofilaments and thick myofilaments. ‘The myofilaments do not extend the entire length of a muscle fiber: they are arranged in compartments called sarcomeres (Figure 8-28). Sarcomeres are separated from one another by narrow zones ‘of dense material called Z lines. Within a sarcomere is a dark area, called the A band, composed of the thick myofilaments. The ends of the A band are darker because of overlapping thick and thin myofilaments. The length of darkening depends on the extent of overlapping. A light-colored arca called the 1 band is ‘composed of thin myofilaments. This combination of alternating dark A bands and light I bands gives the muscle fiber its striated (striped) appearance. Thin myofilaments are anchored to the Z lines and project in ‘both directions. They are composed mostly of the protein actin arranged in two single strands that entwine like a rope (Figure 8-32), Each actin molecule contains a myosin-binding site for a ‘myosin molecule (described shortly). Besides actin, the thin myofi- laments contain two other protein molecules, tropomyosin and troponin, that help regulate muscle contraction. “Thick myofilaments are composed mostly ofthe protein myosin, which is shaped like a golf club. The tails (handles of the golf club) are arranged parallel to each other, forming the shaft of the thick myofilament. The heads of the golf clubs project outward on the surface of the shaft. These projecting heads are referred to a eros bridges and contain an actin-binding ste and an ATP. binding site (Figure 8-3). In ee el attention hasbeen focused on the use of muscle, suing anabolic steroids by amateur athletes. These steroids, a arsvatve of the hormone testosterone, are used 0 build muscle frotens and therefore increase strength and endurance, However, Prot steroids have a number of side effects, including liver ariter, kidney damage, increased risk of heart disease, imitability sad aggressive behavior, psychotic symptoms (including hallucina- fiong)_and mood swings. In females, additional side effects include Silty, the development of facial hai, deepening of the voice, frophy of the breasts and uterus, enlargement of the clitoris, and imegularities of menstruation. In males, additional side effects in- ‘clude testicular atrophy, baldness, excessive development of breast glands, and diminished hormone secretion and sperm production by the testes. Steroids may also be addictive _ CONTRACTION SLIDING-FILAMENT MECHANISM During muscle contraction, thin myofilaments slide inward toward the center of a sarcomere (see Figure 8-5d). The sarcomere shortens, but the lengths of the thin and thick myofilaments themselves do rot change. The myosin cross bridges of the thick myofilaments connect with portions of actin of the thin myofilaments. The myosin cross bridges move like the oars of a boat on the surface of the thin myofilaments, and the thin and thick myofilaments slide past cach other. The myosin cross bridges may pull the thin myofilaments of each sarcomere so far inward toward the center of a sarcomere that their ends overlap. AS the thin myofilaments slide inward, the Z lines are drawn toward each other and the sarcomere is shortened. The sliding of myofilaments and shortening of sarco- meres cause the shortening of the muscle fibers. This isthe slidin Silament mechanism of muscle contraction. This process occurs ‘only when there are sufficient calcium ions (Ca®*) and an adequate supply of energy. NEUROMUSCULAR JUNCTION For a skeletal muscle fiber to contract, it must be stimulated by a nerve cell, or neuron. The particular type of neuron that stimulates muscle tissue is called a motor neuron. ‘When the axon of a motor neuron enters a skeletal muscle, it branches into axon terminals that approach—but do not touch— the sarcolemma of a muscle fiber. This region of the sarcolemma near the axon terminal is known as the motor end plate, The term neuromuscular junction refers to the axon terminal of @ ‘motor neuron together with the motor end plate (Figure 8-4). The ends of the axon terminals are expanded into bulblike structures called synaptic end bulbs (see Figure 9-7). These bulbs contain sacs called synaptic vesicles filled with chemicals called neuro ‘transmitters. The space between the axon terminal and sarcolemma is known as a synaptic cleft. ‘When a nerve impulse (nerve action potential) reaches an axon teeminal, Ca" ions from interstitial fd emer the synaptic end Contraction * 145 URE 22 Hsloay of skeletal muscle tae. (a) Enlarged a «muscle fiber (cel) based on an electron ne, spect of several myofbris of mo hie ond ee ee rograph. (b) Enlarged aspect of a sarcomere Sarcoplasmic TH ad reticulum, Mitochondrion Nucleus Sarcolemma Thin myofilament Myofibrit ‘Thick myotilament band Aband band Sarcomere @ ‘and Abana band “Tick myeflament Cross tipo zie » "TT OF what doos the A band consist? OF what does the bond consis Where are clcium (Co?*) ions stored? 146 * Chapter 8 The Muscular System FIGURE 8.3 Detailed structure of portions of Thi imyoflament. (6 Thick myoflament eigen ‘Act Troponin Tropemyosin Myosin-binding site, EI hin myoflaments ore composed of wo strands of molecules of which substance? Thick myofilaments ore composed of molecules of which substance? bulb, causing synaptic vesicles to release a neurotransmitter (Figure 8-4c). The neurotransmitter released at neuromuscular junctions is acetylcholine (as'-é+til-KO-Ién), or ACh. The ACh diffuses across the synaptic cleft to combine with receptors on the sarco- lemma. This combination alters the permeability ofthe sarcolemma to sodium and potassium ions and ultimately produces a muscle ‘action potential that travels along the sarcolemma and results in contraction. The details of nerve impulse generation and muscle action potential are discussed in Chapter 9. ‘As long as ACh is present in the neuromuscular junction, it will stimulate the muscle fiber. Continuous stimulation by ACh is prevented by an enzyme called acetylcholinesterase (AChE), or simply cholinesterase (k0'-lin-ES-ter-As). ACHE is found on the sarcolemma surface. AChE inactivates ACh within Ys second by breaking it down into its components, acetate and choline ‘This permits the muscle fiber to prepare itself so another muscle ion potential may be generated. When the next nerve impulse ‘comes through, the synaptic vesicles release more ACh, another muscle action potential is generated, and ACHE again inactivates ‘ACh. This cycle is repeated over and over again. Eventually, choline reenters the synaptic end bulb, where it combines with acetate produced in the end bulb to form more ACh that is distributed into synaptic vesicles. PHYSIOLOGY OF CONTRACTION Let us look more closely at the neuromuscular junction and how the molecular events there cause a muscle fiber to contract. Ca?* FIGURE 8-4 Neuromuscular junction. (a} Diagram based on « photon. Grooreph (b) Enlerged aspect based on on electron micrograph Nuclei of ‘muscle fiber yobs of muscle fiber ‘co weil Synaptic vesicle Sarcalemm Synaptic ena oa Sarceptsm nae ‘vale et © TI Wars the motor end plaet ions and energy, in the form of ATP, are required for a muscle ‘o contract. When a muscle fiber is relaxed (not contracting). the" is @ low concentration of Ca* ions in the sarcoplasm. Also: i? # relaxed muscle, the concentration of ATP is high and attached (© ‘the ATP-binding sites on myosin cross bridges (Figure 8-54). T myosin cross bridges cannot combine with actin ofthe thin myo"! Contraction * 14 FAGURE 8-4 (Continued) (¢) Release, ction, by {1)A nerve action potential couses (2) celery soon srthosis of acehchotne (ACH ond (4) induce the release of ACh from @ tronic sy hte @ smaptc end bulb © synaptic vesicle. (5) ACh diffuses across the pic deft and two molecules of ACK 0s cross the cbiece of ACh the henna reins eee YA AcePI he srcenma, In the Sarcolemma: ‘Synaptic ‘end bulb ‘Synaptic Neurotransmitter ae (ACh) receptor UI Witaris ACh and how does i foncin ina contraction? What is AChE andi function? ments as long as tropomyosin and troponin are attached to actin and the ATP is attached to the myosin cross bridges. ‘What causes the concentration of Ca** ions to increase enough for contraction to occur? To understand, we need to back up 0 when ACh diffuses across the synaptic clef. ‘When ACh diffuses across the synaptic cleft and combines with Teeeptors of the sarcolemma, it causes a muscle action potential '0 develop and spread over the surface of the sarcolemma, into the transverse tubules, and to the sarcoplasmic reticulum. The reticulum then releases Ca** ions into the sarcoplasm. ‘The Ca?* ions cause the troponin on the thin myofilaments to ‘change structurally in such a way thatthe troponin and the atached tropomyosin move into a groove between actin strands, thus expos- ing the myosin-binding sites on actin (Figure 8-5). ‘Thevenergy for muscle contraction is supplied by ATP. ATP is found on myosin cross bridges. When a muscle action potential stimulates a muscle fiber, the myosin cross bridges split ATP into ADP + P, releasing energy. The myosin cross bridges, activated (energized) by the energy from the splitting of ATP, combine ‘with myosin-binding sites on actin (Figure 8-5c). As a result, the ADP + Ps released, and the myosin cross bridge moves toward the center of the sarcomere. This movement of the myosin cross bridges, called the power stroke, is the force that causes the thin actin myofilaments to slide past the thick myosin myofilaments. 148 + Chapter 8 The Muscular System ich FIGURE 8.5 Mechanism of muscle contraction. (o) Relaxed state of a muscle fiber in whi ‘tropomyosin and troponin cover the myosin-binding site on actin and the A Sarde eet of the myosin cross bridge is ocuped.(b) Calcium ions combine with troponin, ond troponin tnd the attached tropomyosin move, exposing the myosi-binding st.) A myosin ross bridge splits ATP into ADP + P. The energy from the epliting of ATP acvaos the myosin «10s bridge and it combines with the myotn-bndng ste on oct, As a res the myosin cross bridge moves toward the center of the sarcomere (power stroke), which slides ‘actin myofilament past the myosin myofilament, CO What i the immediate soure y for contraction? What substance is used to generate ATP if contraction continues for more than a few seconds? ‘Once the power stroke is complete, the myosin cross bridge detaches from actin. A given myosin ctoss bridge then combines with another ‘myosin-binding site further along the actin strand. Again, ATP is split and the cycle repeats itself. The myosin cross bridges keep ‘moving back and forth like the cogs of a ratchet with each power stroke, moving the thin actin myofilaments toward the center of sarcomere. The Zines ae drawn toward each other, the sarcomere shortens, the muscle fibers contract, and thus, the muscle itself contracts (Figure 8-54) What happens when a muscle fiber goes from a contracted state back to a relaxed state? The process simply reverses itself from the point at which AChE destroys the ACh that initiated the events leading to contraction. Once ACh is inactivated, no more muscle action potentials are generated. Ca** ions are pumped back into the sarcoplasmic reticulum for storage, myosin cross bridges detach from actin, ADP is resynthesized to ATP, and the thin myofilaments slide back to their relaxed positions. Thus, the muscle resumes its original length Following death, muscles become rigid, unable to contract or stretch, a condition called rigor mortis. Without ATP, myosin ‘ross bridges remain attached to the actin myofilaments, thus pre venting relaxation. The time elapsing between death and the onset of rigor mortis varies greatly among individuals. Those who have hhad long, wasting illnesses undergo rigor mortis more quickly. © ENERGY FOR CONTRACTION ‘Skeletal muscle fibers, unlike other body cells, alternate between Virtual inactivity and continuous activity, Although ATP is the immediate source of energy for muscular contraction, muscle fibers usually contain only enough ATP to sustain activity for about 5 to 6 seconds, Skeletal muscle fibers contain the enzyme ATPase and a high-energy molecule called phosphocreatine (fos'-f0-KRE- ‘tin, both of which can be used to quickly produce more ATP during prolonged exercise. In the presence of ATPase, ATP is broken down into ADP + P and energy is released as follows: ATP— ADP + P + Energy Phosphocreatine breaks down into creatine and phosphate. and in Zine Contraction * 149 cond maximally ofthe thin myofilaments and the relative se of te CONTRACTING === Spree MAXIMALLY CONTRACTED Shin myotiament ‘Thick myoflament er ‘Two sarcomeres @ ET in the sicing-flamont mochonism of contraction, which floments side? the process, large amounts of energy are released as follows: Phosphocreatine —> Creatine + Phosphate + Energy The released energy is used to convert ADP to ATP and the ATP is then used as a source of energy for contraction. This mechanism Provides enough energy for muscles t0 contract maximally for about 15 seconds, as in a 100-meter dash. In other words, itis ‘good for maximal short bursts of activity ‘When muscle activity continues so that even the supply of phos- Phocreatine is depleted, glucose must be broken down. Skeletal muscles store glucose in the form of glycogen. During exercise, lycogen is converted back to glucose. Once this occurs, glucose is split into two molecules of pyruvic acid in the sarcoplasm, a process called glycolysis. In the process, energy is released and used to form ATP. Glycolysis does not require oxygen, $0 it is ‘an anaerobic process. For this reason, glycolysis is referred to 4&8 anaerobic respiration. It may be summarized as follows: 1 Glucose —> 2 Pyruvic Acid + Energy (ATP) Normally, muscle fibers contain oxygen (03). This allows the Pyruvic acid formed by glycolysis to enter mitochondria, where completely catabolized to carbon dioxide and water. Since this breakdown requires O,, it is referred to as aerobic respiration cor cellular respiration. The process is assisted by the presence ‘of myoglobin, which helps store O; for use during vigorous exercise. ‘The complete catabolism of pyruvic acid also yields energy that is used to generate most of a muscle fiber’s ATP: Pyruvic acid + O,—> CO, + H;0 + Energy (ATP) If there is insufficient oxygen for the complete catabolism of pyruvic acid, most of the pyruvic acid is converted to lactic acid, some of which diffuses out of the muscle fibers and into blood. ‘The production of lactic acid also releases energy from glucose that can be used to produce ATP, and it occurs anaerobically. ‘Anaerobic respiration (glycolysis) and aerobic respiration (cellular respiration), together, allow for prolonged muscular activity, such as jogging, and will continue as long as nutrients and adequate oxygen last. Anaerobic respiration can provide sufficient energy for 30 to 40 seconds of maximal muscle activity, such as « 400- meter dash. HOMEOSTASIS “Mascle tissue plays a vital role in maintaining the body's homeosta- sis. Three examples are the relationship of muscle tissue to oxygen, to fatigue, and to heat production. 150 + Chapter 8 The Muscular System Oxygen Debt During exercise, blood vessels in muscles dilate and blood flow is increased in order to increase the available oxygen supply. But ‘when muscular exertion is very great, oxygen cannot be supplied to muscle fibers fast enough, and the aerobic breakdown of pyruvic acid cannot produce all the ATP required for further muscle contrac~ tion. Additional ATP is then generated by anaerobic glycolysis. In the process, however, most of the pyruvic acid is converted to lactic acid. About 80 percent ofthis lactic acid diffuses from the skeletal muscles and is transported to the liver for conversion back to glucose or glycogen, but some lactic acid accumulates in muscle tissu. Ultimately, this lactic acid must be broken down cornpletely into carbon dioxide and water. After exercise has stopped, extra oxygen is required to metabolize the lactic acid; replenish ATP, phosphocreatine, and glycogen; and pay back any oxygen that has been borrowed from hemoglobin, myoglobin, air in the lungs, and body fluids. The additional oxygen that must be taken into the body after vigorous exercise to restore all systems to their normal states is called oxygen debt. The debt is paid back by labored breathing that continues after exercise has stopped. Thus, accumulated lactic acid causes hard breathing and sufficient discom- fort to stop muscle activity until homeostasis is restored, ‘The maximum rate of oxygen consumption during the aerobic catabolism of pyruvic acid is called maximal oxygen uptake. Highly trained athletes can have maximal oxygen uptakes that are twice that of average people, probably owing to a combination of genetics and training. As a result, they are capable of greater activity without increasing their lactic acid production, and their oxygen debts are less. For these reasons, they do not become short of breath as readily as untrained individuals. Muscle Fatigue Ifa skeletal muscle or group of skeletal muscles is continuously stimulated for an extended period of time, the contraction becomes Progressively weaker until the muscles no longer respond, The inability of a muscle to maintain its strength of contraction is, called muscle fatigue. It is related to an inability of muscle to Droduce sufficient energy to meet its needs. Although its exact mechanism is not completely understood, it may be related to insufficient oxygen, depletion of glycogen, and/or lactic acid buildup. Increased lactic acid would cause a decrease in the pH ofthe cells" environment. Muscle fatigue may, therefore, be viewed 438 a homeostatic mechanism that prevents pH levels from dropping below the normal acceptable range for the homeostasis of cells, Heat Production Homeostatic mechanisms are used to regulate temperature (as previ- ‘ously described in Chapter 5). Of the total energy released during ‘muscular contraction, only a small amount is used for mechanical work (contraction). As much as 85 percent is released as heat to help maintain normal body temperature, Excessive heat loss by the body results in shivering, an increase in muscle tone, which increases the rate of heat production by several hundred percent as an effort to raise body temperature back to normal. MOTOR UNIT re ‘motor unit is composed of a motor neuron and all the muscle at, engl or neon cmET oma Me fibers. However, muscles that control precise movements, such as the extemal eye muscles, have fewer than 10 muscle fibers in Gach motor unit, Muscles of the body that are responsible for ross movements, such asthe biceps brachii inthe arm and gastro emius in the leg, may have as many as 2000 muscle fibers in

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