Saladin 5e Extended Outline

Chapter 11 Muscular Tissue

I. Types and Characteristics of Muscular Tissue (pp. 404 405! A. In animals, muscle cells have evolved that are specialized for movement. (p. 404) B. Skeletal muscle is the type of muscle that holds the ody erect a!ainst the pull of !ravity and produces its out"ardly visi le movement. (p. 404) #. $uscle cells have five universal characteristics. (p. 404) %. &esponsiveness (e'cita ility). &esponsiveness is a property of all livin! cells, ut muscle and nerve cells have developed this property to the hi!hest de!ree. (. #onductivity. Stimulation of a muscle fi er produces a "ave of e'citation that travels alon! the fi er and initiates muscle contraction. ). #ontractility. $uscle fi ers are uni*ue in their a ility to shorten "hen stimulated, allo"in! them to pull on ones and other tissues. 4. +'tensi ility. In order to contract, a muscle cell must e a le to stretch a!ain et"een contractions, skeletal muscle fi ers can stretch to as much as three times their contracted len!th. -. +lasticity. .hen a muscle cell is stretched and then the tension released, it recoils to its ori!inal restin! len!th. #. Skeletal muscle may e defined as voluntary striated muscle that is usually attached to one or more ones. (pp. 404/40-) %. Skeletal muscle has li!ht and dark transverse ands called striations. (0i!. %%.%) (. Skeletal muscle is called voluntary ecause it is usually su 1ect to conscious control, other types of muscle are involuntary, and they are never attached to ones. ). A typical skeletal muscle cell is a out %00 2m in diameter and ) cm lon!, some are a thick as -00 2m and )0 cm lon!. 4. Because of their len!th, skeletal muscle cells are usually called muscle fi ers of myofi ers. -. A skeletal muscle is composed of not only muscle cells ut also fi rous connective tissue. a. 3he endomysium surrounds each muscle fi er. . 3he perimysium undles muscle fi ers to!ether into fascicles. c. 3he epimysium encloses the entire muscle.

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d. 3hese connective tissues are continuous "ith colla!en fi ers of the matri', "hich in turn are continuous "ith the colla!en of the one matri'.

6. 3he colla!en of muscles is neither e'cita le nor contractile, ut it is some"hat e'tensi le and elastic. a. 3he colla!en resists e'cessive stretchin! "hen a muscle len!thens and protects the muscle from in1ury. . Some elieve that the recoil of tendons contri ute to po"er output and efficiency of a muscle, others feel that the elasticity in humans is ne!li!i le. II. Microscopic "nato#y of S$eletal Muscle (pp. 405 410! A. 3he muscle fi er has a comple', ti!htly or!anized internal structure that is closely tied to its contractile function (pp. 40-/406) %. 3he plasma mem rane is called the sarcolemma, and its cytoplasm is the sarcoplasm. a. 3he sarcoplasm is occupied mainly y lon! protein undles called myofi rils a out % 2m in diameter. (0i!. %%.() . It also contains !lyco!en, a starchlike car ohydrate, "hich stores ener!y, and the red pi!ment myo!lo in, "hich stores o'y!en. (. $uscle fi ers have multiple flattened or sausa!e7shaped nuclei pressed a!ainst the inside of the sarcolemma. a. 8urin! em ryonic development, several stem cells called myo lasts fuse to produce each muscle fi er, each contri utin! a nucleus. . Some myo lasts remain as unspecialized satellite cells et"een the muscle fi er and endomysium, these can multiply and produce ne" muscle fi ers to some de!ree. ). $ost other or!anelles are packed into the spaces et"een the myofi rils. a. 3he smooth endoplasmic reticulum, called the sarcoplasmic reticulum (S&), forms a net"ork around each myofi ril, it periodically has dilated end7sacs called terminal cisternae. . 4steocytes function to resor or deposit one matri', contri utin! to the homeostatisis of one density and lood concentrations of calcium and phosphate ions c. 3he sarcolemma has tu ular infoldin! called transverse (3) tu ules that penetrate throu!h the cell and open onto the other side. i. +ach tu ule is closely associated "ith t"o terminal cisternae. ii. A 3 tu ule and its associated t"o terminal cisternae constitute a triad.

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d. 3he sarcoplasmic reticulum is a reservoir of calcium ions, it has !ated

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channels in its mem ranes that allo" a flood of calcium ion into the cytosol "here it activates muscle contraction. e. 3he 3 tu ule si!nals that S& "hen to release these calcium ursts. B. +ach myofi ril is a undle of parallel protein microfilaments called myofilaments, there are three kinds of myofilaments. (pp. 406/409) %. 3he thick filaments are a out %- nm in diameter is each is made of several hundred myosin molecules. (0i!. %%.)a, , d) a. A myosin molecule is shaped like a !olf clu , "ith t"o chains intert"ined to form a shaftlike tail and a dou le !lo ular head pro1ectin! from the tail at an an!le. . A thick filament may e likened to a undle of (00 to -00 such :!olf clu s,; "ith their heads directed out"ard around the undle. c. <alf of the heads an!le to the left and half to the ri!ht, "ith a are zone in the middle. (. 3he thin filaments, 9 nm in diameter, are composed of t"o intert"ined strands of fi rous (0) actin. (0i!. %%.)c, d) a. +ach 0 actin strand is strin! of su units called !lo ular (=) actin. . +ach = actin has an active site that can ind to the head of a myosin molecule. c. A thin filament also has 40 to 60 molecules of the protein tropomyosin, "hich locks active sites of some = actins "hen a muscle fi er is rela'ed. d. A calcium7 indin! protein, troponin, is ound to every tropomyosin molecule. ). +lastic filaments, % nm in diameter, are made of a lar!e protein called titin (connectin). (0i!. %%.- ) a. +lastic filaments flank each thick filament and anchor it to a structure called the > disc, helpin! to sta ilize the thick filament. 4. $yosin and actin are called contractile proteins ecause they accomplish the shortenin! of the muscle fi er. -. 3ropomyosin and troponin are called re!ulatory proteins ecause they act like a s"itch to determine "hen the fi er can contract or not contract. a. 3he action of these re!ulatory proteins depends on the availa ility of calcium ions, "hich ind to troponin. 6. At least seven other accessory proteins occur in the thick and thin filaments or are associated "ith them, the most clinically important is dystrophin.

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a. 8ystrophin is an enormous protein located et"een the sarcolemma and the outermost myofilaments, it links actin filaments to a peripheral protein on the inner face of the sarcolemma. (0i!. %%.4) . 3he peripheral protein in turn is linked to transmem rane proteins that connect "ith proteins e'ternal to the fi er that ultimately link "ith the asal lamina and to the endomysium. c. 8ystrophin is therefore a key element in transferrin! the forces of myofilament movement to the connective tissue of the muscle as a "hole. d. =enetic defects in dystrophin are responsi le for muscular dystrophy.

#. Striations are a precise array of the a undant myosin and actin found in skeletal and cardiac muscle. (pp. 409/4%0) (0i!. %%.-) %. Striated muscle has dark A (anisotropic) ands and li!hter I (isotropic) ands, referrin! to their effect on polarized li!ht. a. +ach A and consists of thick filaments lyin! side y side. i. 5art of the A and is especially dark, h"ere each thick filament is surrounded y thin filaments. ii. In the middle of the A and is a li!ht re!ion called the < and, into "hich thin filaments do not reach. iii. 3he thick filaments ori!inate at a dark $ line in the middle of the < and. . +ach li!ht I and is isected y a dark narro" > disc (> line), "hich provides anchora!e for the thin filaments and elastic filaments. c. +ach se!ment of a myofi ril from one > disc to the ne't is called a sarcomere, it is the functional contractile unit of the muscle fi er. i. A muscle shortens ecause its individual sarcomeres shorten and pull the > discs closer to!ether. ii. 8ystrophin and the linkin! proteins pull on the e'tracellular proteins of the muscle. iii. 3he > discs pull on the sarcolemma to achieve overall shortenin! of the cell. d. 3he terminolo!y of muscle fi er structure is sho"n in 3a le %%.%. III. The %er&e Muscle 'elationship (pp. 410 41(! A. 3he relationship et"een nerve and muscle cells is important to understandin! muscle contraction ecause skeletal muscle never contracts unless it is stimulated y a nerve (or artificially "ith electrodes). (p. 4%0)

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B. ?erve cells called somatic motor neurons, "ith cell odies in the rainstem and spinal cord, stimulate muscle fi ers via their a'ons, called somatic motor fi ers, a sin!le motor fi er and all the muscle fi ers it innervates are collectively called a motor unit. (pp. 4%0/4%%) (0i!. %%.6) %. 3he muscle fi ers of a sin!le motor unit are not clustered to!ether ut are dispersed throu!hout a muscle, so that their stimulation causes a "eak contraction over a "ide area. (. 4n avera!e, a out (00 muscle fi ers are innervated y each motor neuron, ut "ide variation e'ists for different purposes. a. Small motor units are present "here fine control is needed, such as in the muscles of eye movement () to 6 muscle fi ers per neuron). . @ar!e motor units are present "here stren!th is more important than fine control, such as in the !astrocnemius (%,000 muscle fi ers per neuron). ). $ultiple motor units also have the advanta!e of allo"in! :shifts; in muscle contraction, so that "hen some units ecome fati!ued, others can take over. #. 3he point "here a nerve fi er meets its tar!et cell is a synapse, and "hen the tar!et cell is a muscle fi er, the synapse is called a neuromuscular 1unction (?$A) or motor end plate. (pp. 4%%/ 4%)) (0i!. %%.9) (3a le %%.() %. 4ne nerve fi er stimulates the muscle fi er at several points "ithin the ?$A, each terminal ranch of the nerve fi er has its o"n synapse point to the muscle fi er. (. At each synapse, the nerve fi er ends in a ul ous s"ellin! called a synaptic kno , "hich is separated from the muscle fi er y a narro" space 60/%00 nm "ide called the synaptic cleft. a. A third cell, a Sch"ann cell, envelopes the entire 1unction. . A synaptic kno contains spheroidal or!anelles called synaptic vesicles, "hich are filled "ith acetylcholine (A#h). c. A#h is released into the synaptic cleft "hen the nerve impulse reaches the nerve endin!s. d. 3he muscle fi er has a out -0 million A#h receptors incorporated into its sarcolemma, "hich contains numerous infoldin!s at the 1unction called 1unctional folds that increase surface area for receptors. e. A deficiency of A#h receptors leads to the muscle paralysis of myasthenia !ravis. ). 3he muscle fi er and the Sch"ann cell of the ?$A are surrounded y a asal lamina that separates them from the surroundin! connective tissue. a. 3he asal lamina ases throu!h the synaptic cleft and fills it. . Both the sarcolemma and the asal lamina "ithin the cleft contain the enzyme acetylcholinesterase (A#h+), "hich reaks do"n A#h. Insight 11.1 ?euromuscular 3o'ins and 5aralysis

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8. $uscle fi ers and neurons are considered electrically e'cita le cells ecause their plasma mem ranes e'hi it volta!e chan!es in response to stimulation. (p. 4%(/4%)) %. 3he study of the electrical activity of cells is called electrophysiolo!y. (. In an unstimulated (restin!) cell, more anions (ne!ative ions) are found on the inside of the plasma mem rane than on the outside, the mem rane is therefore polarized, or char!ed. a. In a restin! muscle cell, ?aB is in e'cess in the e'tracellular fluid (+#0) and CB is in e'cess in the intracellular fluid (I#0). . Anions such a proteins, nucleic acids, and phosphates are also inside the I#0 and cannot cross the mem rane. ). A difference in electrical char!e et"een t"o points is called an electrical potential, or volta!e. a. 3he volta!e across the sarcolemma of a muscle cell is only a out /D0 mE and is called the restin! mem rane potential (&$5). . 3he ne!ative si!n indicates that the ne!ative char!e is !reater on the inside of the mem rane. c. 3he restin! mem rane potential is maintained y the sodium/potassium pump. 4. .hen a nerve or muscle cell is stimulated, ion !ates open in the plasma mem rane and ?aB diffuses do"n its concentration !radient into the cell. a. 3hese cations override the ne!ative char!es in the I#0, and the inside of the mem rane riefly ecomes positive, a chan!e termed depolarization. -. 3he ?aB !ates then close and the CB !ates open, CB rushes out of the cell turnin! the inside of the mem rane ne!ative a!ain. a. 3his chan!e is termed repolarization. 6. 3he volta!e shift of depolarization follo"ed y repolarization is called an action potential. 9. Action potentials perpetuate themselves alon! a mem rane, a "ave of action potentials movin! alon! a nerve fi er is called a nerve impulse or nerve si!nal. I). *eha&ior of S$eletal Muscle +i,ers (pp. 414 4-0! A. 3he process of muscle contraction and rela'ation has four ma1or phasesF e'citation, e'citation/ contraction couplin!, contraction, and rela'ation. (p. 4%4) (0i!s. %%.G, %%.D, %%.%0, %%.%%) B. +'citation is the process in "hich action potentials in the nerve fi er lead to action potentials in the muscle fi er, it can e divided into five steps. (p. 4%4) (0i!. %%.G) %. A nerve si!nal arrives at a synaptic kno and stimulates volta!e7re!ulated #a(B !ates to open, calcium ions enter the synaptic kno .

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(. #a(B stimulates e'ocytosis of synaptic vesicles, "hich release A#h into the synaptic cleft. ). A#h diffuses across the synaptic cleft and inds to receptor proteins on the sarcolemma. 4. 3he receptors are li!and7re!ulated ion !ates that ind t"o A#h molecules to open. a. .hen the !ates are opened, ?aB diffuses into the cell and CB diffuses out, the sarcolemma reverses polarity from /D0 mE to B9- mE, then falls ack a!ain as CB diffuses out. . 3his rapid fluctuation in mem rane volta!e at the motor end plate is called end7plate potential (+55). -. Areas ad1acent to the ?$A have ion7specific volta!e7re!ulated !ates that open in response to the +55, allo"in! flo" of ?aB in and CB out, !eneratin! an action potential, the muscle fi er is no" e'cited. #. +'citation/contraction couplin! refers to the events that link the action potentials on the sarcolemma to activation of the myofilaments, this process has four steps that follo" from e'citation. (p. 4%4) (0i!. %%.D) %. (6) A "ave of action potentials spreads from the end plate in all directions, and enters the 3 tu ules, continuin! do"n them into the sarcoplasm. (. (9) Action potentials open volta!e7re!ulated ion !ates in the 3 tu ules. a. 3hese !ates are linked to calcium channels in the terminal cisternae of the sarcoplasmic reticulum (S&). a. .hen the channels in the S& open, #a(B diffuses out of the S& and into the cytosol do"n its concentration !radient. ). (G) #alcium inds to the troponin of the thin filaments. 4. (D) 3he troponin/tropomyosin comple' chan!es shape, e'posin! active sites on the actin filaments that can ind to myosin heads. #. #ontraction is the step in "hich the muscle fi er develops tension and may shorten, the mechanism of contraction "as proposed in %D-4 y <anson and <u'ley as the slidin! filament theory. 3he process can e divided into four steps that follo" e'citation/contraction couplin!. (pp. 4%4/4%G) (0i!. %%.%0) %. (%0) $yosin A35ase hydrolyzes A35 that is ound to the myosin head, the ener!y released activates the head y chan!in! its shape into a :cocked; position. (. (%%) .ith A85 and phosphate still ound, the activated myosin head inds to an e'posed active site on the thin filament, formin! a cross rid!e. ). (%() $yosin releases the A85 and phosphate and fle'es into a ent, lo" ener!y shape, tu!!in! the thin filament alon! "ith it, this is called the po"er stroke.

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4. (%)) Hpon indin! to another A35, myosin releases the actin, it is no" prepared to repeat the process y hydrolyzin! the A35 and recockin! (the recovery stroke). It "ill then attach to a ne" active site farther do"n. a. .hen one myosin releases an actin, many other heads on the same thick filament are still ound to actin on the thin filament so it does not slide ack. . +ven thou!h the muscle fi er contracts, the myofilaments do not ecome shorter, instead, the thin filaments slide over the thick ones. c. 3he cycle of po"er and recovery is repeated many times y each myosin head, at a rate of a out five strokes per second. 8. .hen stimulation ceases, a muscle fi er rela'es and returns to it restin! len!th, the process can e divided into five steps that follo" the contraction phase. (p. 4%D) (0i!. %%.%%) %. (%4) ?erve si!nals stop arrivin! at the ?$A, so the synaptic kno stops releasin! A#h. (. (%-) As A#h dissociates from the receptor, A#h+ reaks it do"n, the synaptic kno rea sor s the fra!ments as usual, ut no" no ne" A#h replaces that "hich is roken do"n. ). (%6) Active transport pumps in the S& pump #a(B from the cytosol ack into the cisternae. a. 3he #a(B in the cisternae inds to a protein called calse*uestrin and is stored until stimulation occurs a!ain. . Active transport re*uires A35, therefore A35 is needed for oth muscle contraction and muscle rela'ation. 4. (%9) As #a(B dissociates from troponin, it is pumped into the S& and not replaced. -. (%G) 3ropomyosin moves ack into position, lockin! the active sites of the actin filament and preventin! myosin indin!. 6. A muscle returns to its restin! len!th "ith the aid of t"o forces. a. Its intracellular and perhaps e'tracellular elastic components stretch it, like a recoilin! ru er and. . 3he contraction of an anta!onist muscle len!thens it, for e'ample, contraction of the triceps rachii stretches the iceps rachii. Insight 11.2 &i!or $ortis +. 3he amount of tension a muscle !enerates depends on ho" stretched or contracted it "as efore it "as stimulated, this principle is termed the len!th/tension relationship. (p. 4(0) (0i!ure %%.%() %. If a muscle fi er is overly contracted at rest, then upon stimulation the thick filaments can contract no farther than the > discs and the contraction is "eak. (. If a muscle fi er is too stretched, then upon stimulation there is little overlap et"een thick and thin filaments, and the myosin heads cannot !et a !ood !rip on the actin and the contraction is "eak.

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). $uscle has an optimum restin! len!th at "hich it can respond "ith !reatest force, the central nervous system continually ad1usts the len!th of restin! muscles in a state of partial contraction called muscle tone. ). *eha&ior of .hole Muscles (pp. 4-0 4-5! A. A nerve/muscle preparation, such as the !astrocnemius and sciatic nerve from a fro! attached to stimulatin! electrodes, can e used to record a chart of stimulation and muscle contraction called a myo!ram. (p. 4(%) %. A "eak (su threshold) electrical stimulus causes no contraction, as volta!e is increased the threshold is reachedIthe minimum volta!e necessary to !enerate an action potential and cause contraction. a. 3he action potential tri!!ers the release of #a(B into the cytosol and activates the slidin! filament mechanism. . At threshold or hi!her, a stimulus causes a *uick cycle of contraction and rela'ation called a t"itch. (0i!. %%.%)) (. A delay, or latent period, of a out ( milliseconds occurs et"een the onset of the stimulus and the onset of the t"itch. a. 8urin! this time e'citation, e'citation/contraction couplin!, and tensin! of elastic components occurs. . 3he force !enerated is called internal tension, and it does not sho" up on the myo!ram ecause the muscle does not yet shorten. ). 4nce elastic components are taut, the muscle e!ins to produce e'ternal tension, this is called the contraction phase of the t"itch. a. 3he resistin! load in a preparation is the sensor of the recordin! apparatus, so the movement is recorded on the myo!ram. . In the ody, the resistin! load is usually a one. 4. 3he contraction phase is short7lived ecause the S& rea sor s #a(B efore the muscle develops ma'imum force, as the #a(B level falls, muscle tension declines durin! the rela'ation phase. a. 3he muscle is *uicker to contract than it is to rela'. . 3he entire t"itch lasts from 9 to %00 msec. Insight 11.3 =alvani, Eolta, and Animal +lectricity B. Althou!h electrical e'citation of a muscle fi er o eys an all7or7none la", muscle fi ers do not e'hi it all7or7none t"itches in response to e'citation. (pp. 4(%/4()) %. 3"itches vary in stren!th for a num er of reasonsF a. 3"itch stren!th varies "ith stimulation fre*uencyF stimuli arrivin! close to!ether produce stron!er t"itches than those arrivin! far apart.

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. 3"itches vary "ith the concentration of #a(B in the sarcoplasm, "hich can vary "ith stimulation fre*uency. c. 3"itch stren!th depends on ho" stretched the muscle "as 1ust efore stimulation (len!th/tension relationship). d. 3"itches vary "ith the temperature of the muscle, "armer muscle contracts more stron!ly. e. 3"itches are "eaker "hen the p< of the sarcoplasm falls elo" normal, producin! fati!ue. f. 3"itches vary "ith the state of hydration of the muscle, "hich affect overlap et"een filaments and a ility of myosin to form cross7 rid!es "ith actin.

(. $uscles must e a le to contract "ith varia le stren!th for different tasks, so it is not surprisin! that t"itches vary in stren!th. ). Stimulus intensity and stimulus fre*uency have contrastin! effects. (0i!. %%.%4) a. At threshold, a "eak t"itch occurs, and if volta!e is increased, t"itches are stron!er. i. <i!her volta!es e'cite more and more nerve fi ers in the motor nerve and thus stimulate more motor units. ii. 3his effect is called recruitment or multiple motor unit ($$H) summation. . +ven at constant volta!e, a hi!her fre*uency of stimulation produces stron!er t"itches than does a lo"er fre*uency. c. Hp to %0 stimuli per second, a muscle produces an identical t"itch for each stimulus and recovers fully et"een t"itches. (0i!. %%.%-a) d. Bet"een %0 and (0 stimuli per second, the muscle recovers fully et"een t"itches, ut each t"itch develops more tension than the one efore it, this pattern is called treppe or the staircase phenomenon. (0i!. %%.%- ) i. 4ne cause of treppe is that the S& does not have time to completely rea sor all the #aB( released. ii. Another factor is that the heat of each t"itch causes muscle enzymes to "ork more efficiently e. At hi!her stimulus fre*uency ((0/40 stimuli per second) each ne" stimulus arrives efore the previous t"itch is over, each ne" t"itch :pi!!y acks; on the previous one and !enerates hi!her tension. (0i!. %%.%-c) i. 3his phenomenon is called temporal summation or "ave summation. ii. It produces a state of sustained flutterin! contraction called incomplete tetanus.

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f. At still hi!her fre*uency (40/-0 stimuli per second) the muscle has no time to rela' at all and the t"itches fuse into a smooth, prolon!ed contraction called complete tetanus. (0i!. %%.%-d) i. 3his state should not e confused "ith the disease tetanus caused y the tetanus to'in. ii. #omplete tetanus rarely if ever occurs in the ody.

#. #ontraction does not al"ays mean the shortenin! of a muscle, it may mean only that the muscle is producin! internal tension. (pp. 4()/4(-) %. 5hysiolo!ists speak of isotonic versus isometric contraction, and concentric versus eccentric contraction. (. Isometric contraction is contraction "ithout a chan!e in len!th. (0i!. %%.%6a) a. Isometric contraction of anta!onistic muscles at a 1oint maintains 1oint sta ility. . Isometric contraction of postural muscles keeps the ody erect. ). Isotonic contraction is contraction "ith a chan!e in len!th ut no chan!e in tension. (0i!. %%.%6 ) a. Isotonic contraction moves a load as the muscle shortens. 4. Isometric and Isotonic contraction are oth phases of normal muscular action. (0i!. %%.%9) -. Isotonic contraction has t"o formsF concentric and eccentric. a. In concentric contraction, a muscle shortens as it maintains tension, such as "hen the iceps rachii contracts and fle'es the el o" to lift a "ei!ht. . In eccentric contraction, a muscle len!thens as it maintains tension, such as "hen the iceps rachii len!thens as a "ei!ht is lo"ered. )I. Muscle Meta,olis# (pp. 4-5 4(0! A. All muscle contraction depends on A35, and the supply of A35 depends on the availa ility of o'y!en and or!anic ener!y sources such as !lucose and fatty acids. (pp. 4(-/4(6) %. 3he t"o main sources of A35 synthesis are anaero ic fermentation and aero ic respiration, durin! the course of e'ercise, different mechanisms produce A35 dependin! on duration. (0i!. %%.%G) a. Anaero ic fermentation allo"s the cell to produce A35 in the a sence of o'y!en, ut yield is limited and lactic acid, a to'ic end product, is a ma1or factor in muscle fati!ue. . Aero ic respiration produces more A35 and less to'ic end products, ut re*uires a continual supply of o'y!en. c. In a restin! muscle, most A35 is !enerated y the aero ic respiration of fatty acids.

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(. Immediate ener!y such as that needed for a %00 m dash relies on o'y!en stored in myo!lo in. a. 3he muscle orro"s phosphate !roups from other molecules and transfers them to A85 to form A35, t"o enzymes systems control these transfers. (0i!. %%.%D) i. $yokinase transfers phosphate from one A85 to another to form A35. ii. #reatine kinase o tains phosphate from creatine phosphate (#5) and donates it to A85 to make A35. . A35 and #5, collectively called the phospha!en system, provide nearly all the ener!y used for short ursts of intense activity, such as sprintin! for 6 seconds. ). As the phospha!en system is e'hausted, the muscles shift into anaero ic fermentation for short7term ener!y until cardiopulmonary function can catch up "ith the o'y!en demand a. 8urin! this period, muscles o tain !lucose from the lood and from their o"n stored !lyco!en. . 3he path"ay from !lyco!en to lactic acid, called the !lyco!en/lactic acid system, produces enou!h A35 for )0 to 40 seconds of ma'imum activity. 4. After 40 seconds or so, the respiratory and cardiovascular system deliver o'y!en to the muscles fast enou!h for aero ic respiration to meet most of the A35 demand. a. Aero ic respiration produces much more A35 and is a very efficient means of meetin! the A35 demands of prolon!ed e'ercise. . 4'y!en consumption rises for ) to 4 minutes and then levels off at a steady state in "hich A35 production keeps pace "ith demand. c. @ittle lactic acid accumulates under steady state, ut the depletion of !lyco!en and lood !lucose, to!ether "ith loss of fluid and electrolytes, set limits to endurance and performance even "hen lactic acid does not. B. $uscle fati!ue is the pro!ressive "eakness and loss of contractility that results from prolon!ed use of the muscles. (pp. 4(6/4(9) %. 0ati!ue has multiple causes. a. A35 synthesis declines as !lyco!en is consumed. . An A35 shorta!e slo"s do"n the sodium/potassium pumps, "hich affects the restin! mem rane potential and muscle e'cita ility. c. @actic acid lo"ers the p< of the sarcoplasm, inhi itin! enzymes. d. 3he accumulation of CB in the +#0 lo"ers the mem rane potential. e. $otor nerve fi ers use up their A#h (1unctional fati!ue).

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f. 3he #?S fati!ues y processes not yet understood, so that less si!nal output to the muscles occurs.

(. 3he ma'imum o'y!en uptake (Eo(ma') is the point at "hich the rate of o'y!en consumption reaches a plateau and does not increase further, it determines a personJs a ility to maintain hi!h7intensity e'ercise for more than 4 to - minutes. a. Eo(ma' is proportional to ody size, peaks at around a!e (0, and is usually !reater in males than in females. . It can e t"ice as !reat in a trained endurance athlete as in an untrained person. i. A typical sedentary adult "ei!hin! %60 pounds has a Eo(ma' of a out )- m@KminKk!. ii. An elite endurance athlete of the same "ei!ht can have a Eo(ma' of a out 90 m@KminKk!. iii. @ance Armstron!Js Eo(ma' "as measured at G).G m@KminKk!. Insight 11.4 Beatin! 0ati!ueISome Athletic Strate!ies and 3heir <azards #. 4'y!en de t is the difference et"een the restin! rate of o'y!en consumption and the elevated rater after e'ercise, it is also kno"n as e'cess poste'ercise o'y!en consumption (+54#). (pp. 4(9/4(G) %. 3ypically a out %% @ of e'tra o'y!en is consumed after strenuous e'ercise and is used for the follo"in! purposesF a. &eplacin! the odyJs depleted o'y!en reserves, such as o'y!en in myo!lo in and hemo!lo in, dissolved in the lood plasma, and in the air in lun!s. . &eplenishin! the phospha!en system y synthesizin! A35 and transferrin! phosphates to creatine to restore A35 and #5 levels. c. 4'idizin! lactic acid to pyruvic acid in the kidneys, cardiac muscle, and especially the liver, "here pyruvic acid is then converted into !lucose. d. Servin! the elevated meta olic rate resultin! from ody heatin!. 8. $uscle fi ers can e classed accordin! to their physiolo!ical characteristics. (pp. 4(G/4(D) (3a le %%.)) %. Slo" o'idative (S4), slo"7t"itch, red, or type I fi ers have relatively a undant mitochondria, myo!lo in, and lood capillaries and a deep red color. a. S4 fi ers do not fati!ue easily and e'hi it a relatively lon! t"itch (L%00 msec) in response to a sin!le stimulus . +'amples are the soleus muscle of the calf and the postural muscles. (. 0ast !lycolytic (0=), fast7t"itch, "hite, or type II fi ers are adapted for *uick responses ut not for fati!ue resistance.

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5a!e %4
a. 3hey are rich in enzymes of the phospha!en and !lyco!en/lactic acid systems. . 3heir S& releases and rea sor s #a(B *uickly. c. 0= fi ers are poorer in mitochondria, myo!lo in, and lood capillaries than S4 fi ers, so they are relatively pale. e. 3hey produce t"itches as short as 9.- msec. f. +'amples are the !astrocnemius of the calf, rachii of the arm, and muscles of eye movement.

). Some authorities reco!nize t"o su types of 0= fi ers called types IIA and IIB. a. 3ype IIB is the common type 1ust descri ed, "hereas IIA, or intermediate fi ers, com ine fast t"itch "ith fati!ue resistance. i. 3ype IIA is rare e'cept in some endurance7trained athletes. . 3he fi er types can e distin!uished histolo!ically "ith stains for mitochondrial enzymes. (0i!. %%.(0) 4. All muscle fi ers of a sin!le motor unit are of the same physiolo!ical type. -. ?early all muscles are composed of oth S4 and 0= fi ers, ut the proportions differ. a. $uscles composed mainly of S4 fi ers are called red muscles and those composed mainly of 0= fi ers are called "hite muscles. . 3he proportions differ even in a sin!le muscle in people "ith different types and levels of physical activity. (3a le %%.4) c. <eredity may play a role in "hether a person is a : orn sprinter; or : orn marathoner.; d. Sometimes "hen t"o or more muscles appear to have the same function, they may have different proportions of S4 to 0= fi ers and so actually allo" a "ider ran!e of function. +. <umans have far more muscular stren!th than is normally used, and muscles can !enerate more tension than the ones and tendons can "ithstand. (pp. 4(D/4)0) %. $uscular stren!th depends on anatomical and physiolo!ical factorsF a. $uscle stren!th is primarily determined y muscle size. . 0ascicle arran!ement contri utes to stren!th, pinnate muscles such as the *uadriceps femoris are stron!er than parallel muscles such as the sartorius, "hich in turn are stron!er than circular muscles such as the or icularis oculi. c. @ar!e motor units produce stron!er contractions than small ones. d. &ecruitment, or multiple motor unit ($$H) summation produces a stron!er muscle contraction. e. 3emporal summation of action potentials causes stron!er contraction.

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5a!e %f. 3he len!th/tension relationship contri utes in that a muscle restin! at optimum len!th can contract more forcefully. !. 0ati!ue contri utes in that fati!ued muscles contract more "eakly.

(. &esistance e'ercise, such as "ei!ht liftin!, can stimulate muscle !ro"th even if only done a fe" minutes at a time a fe" times a "eek. a. =ro"th results primarily form cellular enlar!ement, not cell division. . $yofi rils !ro" thicker and split lon!itudinally "hen they reach a certain size. c. $uscle fi ers are incapa le of mitosis, ut some evidence indicates that they may also split lon!itudinally as they enlar!e. ). +ndurance (aero ic) e'ercise, such as 1o!!in! and s"immin!, improves the fati!ue resistance of the muscles. a. Slo"7t"itch fi ers produce more mitochondria and !lyco!en and ac*uire a !reater density of lood capillaries "ith endurance e'ercise. . 3his form of e'ercise also improves skeletal stren!th, increases red lood cell count and o'y!en transport capacity, and enhances cardiovascular, respiratory, and nervous system function. 4. #ross7trainin! incorporates elements of oth resistance trainin! and endurance trainin! for optimal performance. )II. Cardiac and S#ooth Muscle (pp. 4(0 4(4! A. Any of the three types of muscle cells can e called myocytes, a term prefera le to muscle fi ers for smooth and cardiac muscle cells ecause they are relatively short and have only one nucleus. (p. 4)0) B. #ardiac and smooth muscle are termed involuntary muscles ecause they are usually not su 1ect to conscious control. (p. 4)0) #. #ardiac muscle cells are also called cardiocytes, cardiac muscle is limited to the heart, "here its function is to pump lood. (p. 4)0) %. 3o fulfill its function, cardiac muscle must have five propertiesF (3a le %%.-) a. It must contract "ith a re!ular rhythm. . 3he cells of a !iven heart cham er must contract in unison. c. +ach contraction must last lon! enou!h to e'pel lood from the cham er. d. 3he muscle must function in sleep and "akefulness. e. It must e hi!hly resistant to fati!ue. (. #ardiac muscle is striated like skeletal muscle ut has shorter and thicker cells "ith uneven, notched ends. (0i!. %D.%%) ). +ach myocyte is 1oined to several others at its ends throu!h linka!es called intercalated discs.

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a. 3hese discs appear as thick dark lines in stained tissue sections.

5a!e %6

. An intercalated disc has electrical !ap 1unctions that allo" each myocyte to stimulate nei!h ors and mechanical 1unctions to hold the myocytes to!ether. 4. 3he sarcoplasmic reticulum of cardiocytes is less developed than in skeletal muscle ut has lar!er 3 tu ules. -. 8ama!ed cardiac muscle is repaired y fi rosis, althou!h mitosis has een detected in cardiac myocytes follo"in! heart attacks, it does not produce si!nificant re!enerated muscle. 6. #ardiac muscle contains a uilt7in pacemaker that rhythmically sets off a "ave of electrical e'citation. a. #ardiac muscle is said to e autorhythmic ecause of this a ility. . 3he heart does receive fi ers from the autonomic nervous system that can increase or decrease heart rate and contraction stren!th. c. #ardiac muscle does not e'hi it *uick t"itches ut maintains tension for (00/ (-0 msec. 9. #ardiac muscle uses aero ic respiration almost e'clusively. a. It is rich in myo!lo in and !lyco!en and has lar!e mitochondria that fill (-M of the cell. . It is adapta le "ith respect to fuel, ut vulnera le to lack of o'y!en. c. Because little anaero ic fermentation takes place, cardiac muscle is hi!hly resistant to fati!ue. 8. Smooth muscle is composed of myocytes "ith a fusiform shape, a out )0/(00 2m lon!, -/%0 2m "ide at the middle, and taperin! to a point at each end, there is only one nucleus. (pp. 4)%/ 4)4) %. 3hick and thin filaments are oth present, ut they are not ali!ned and produce no visi le striations. (. > discs are a sent and in their place are protein pla*ues on the inner plasma mem rane and protein masses called dense odies in arrays in the cytoplasm. ). 3heir cytoplasm contains an e'tensive cytoskeleton of intermediate fi ers, "hich are also attached to the plasma mem rane. (0i!. %%.(4) 4. 3he sarcoplasmic reticulum is scanty, and there are no 3 tu ules. a. #a(B for contraction comes mainly from the +#0 y "ay of channels in the sarcolemma. . 8urin! rela'ation, #a(B is pumped ack out of the cell. -. Some smooth muscle has not nerve supply, ut "hen nerve fi ers are present, they are autonomic, not somatic. 6. Smooth muscle is capa le of mitosis and hyperplasia.

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5a!e %9

9. 3here are t"o functional types of smooth muscleF multiunit and sin!le7unit. (0i!. %%.(%) a. $ultiunit smooth muscle occurs in some lar!e arteries and pulmonary air passa!es, in piloerector muscles, and in the iris. i. Its autonomic nerve innervation is similar to the motor nerve innervation of skeletal muscle, "ith motor units that contract independently. . Sin!le7unit smooth muscle is more "idespread, occurrin! in most lood vessels and in the di!estive, respiratory, urinary, and reproductive tracts. i. It is also called visceral muscle. ii. In hollo" or!ans, it forms t"o or more layers, and inner circular layer and an outer lon!itudinal layer. (0i!. %%.(() iii. Its name refers to the fact that myocytes are electrically coupled to each other y !ap 1unctions, so that cells directly stimulate each other and can contract as a unit. G. Smooth muscle contraction is involuntary and does not re*uire nerve stimulation. a. Some smooth muscle contracts in response to chemical stimuli and in response to stretch. . Some sin!le7unit smooth muscle, especially in the stomach and intestines, has pacemaker cells that set off "aves of contraction, thus it is autorhythmic. c. $ost smooth muscle is innervated y autonomic nerve fi ers that can tri!!er or modify its contraction. i. 3hese nerve fi ers stimulate smooth muscle "ith either acetylcholine or norepinephrine. ii. 3he nerves have contrastin! effects in different locations, such as rela'in! smooth muscle in the arteries "hile contractin! smooth muscle in the ronchioles. d. In sin!le7unit smooth muscle, each autonomic nerve fi er has up to (0,000 eadlike s"ellin!s called varicosities alon! its len!th consistin! of synaptic vesicles and a fe" mitochondria. (0i!s. %%.(%, %%.()) e. Instead of approachin! any one myocyte, the nerve fi er passes amon! several myocytes and stimulates all of them "hen it releases neurotransmitter. i. 3he muscle cells do not have motor end plates, ut instead have receptor sites on their surface. f. 3hese nerve/muscle relationships are called diffuse 1unctions ecause there is no one7to7one relationship et"een nerve fi er and myocyte.

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5a!e %G

D. #ontraction in smooth muscle is tri!!ered y #a(B, ener!ized y A35, and involves slidin! of thin filaments over thick filaments, ut the mechanism of e'citation/ contraction couplin! is very different. a. $ost #a(B comes from the e'tracellular fluid, not from the sarcoplasmic reticulum. . #alcium channels in the sarcolemma admit the #a(B, ut they are of different types, includin! volta!e re!ulated, li!and7re!ulated, and mechanically re!ulated (respond to stretch). c. Smooth muscle has no troponin, instead, calcium inds to a protein called calmodulin, associated "ith thick filaments. i. #almodulin activates the enzyme myosin li!ht7chain kinase, "hich adds a phosphate to a protein on the myosin head. ii. 3his phosphate addition activates myosin A35ase, so that myosin can ind to actin and hydrolyze A35. d. 3hick filaments pull on thin filaments, "hich in turn pull on the dense odies and mem rane pla*ues, transferrin! force to the plasma mem rane and shortenin! the entire cell. e. .hen a smooth muscle cell contracts, it puckers and t"ists some"hat like "rin!in! out a "et to"el. (0i!. %%.(4) f. #ompared to skeletal muscle, smooth muscle is very slo" to contract and rela'. i. 3he latent period et"een stimulation and contraction in smooth muscle is -0/%00 msec, compared "ith ( msec in skeletal muscle. ii. 3ension in smooth muscle peaks a out 0.- sec after stimulus and then declines over a period of % to ( sec. iii. Smooth muscleJs myosin A35ase is a slo" enzyme, and the pumps that remove #a(B from the cell are also slo". !. As #a(B falls, myosinJs a ility to hydrolyze A35 drops, ut it does not detach from actin immediately, its latch rid!e mechanism ena les it to remain attached to actin for a prolon!ed time "ithout consumin! A35. h. Smooth muscle often e'hi its tetanus and is hi!hly resistant to fati!ue. i. It makes A35 aero ically, ut its A35 re*uirement is small. ii. It re*uires %0 to )00 times less A35 as does skeletal muscle to maintain the same amount of tension. i. Smooth muscleJs fati!ue resistance and latch7 rid!e mechanism allo" maintenance of smooth muscle tone (tonic contraction) that keeps arteries in a state of partial constriction called vasomotor tone.

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5a!e %D
i. A loss of vasomotor tone can cause a dan!erous drop in lood pressure. 1. Smooth muscle tone also keep the intestines partially contracted.

%0. Stretch alone sometimes causes smooth muscle to contract. a. 8istension of the esopha!us "ith food or the colon "ith feces causes a "ave of contraction called peristalsis. . Smooth muscle e'hi its a reaction called the stress7rela'ation response or receptive rela'ation response. .hen stretched, it riefly contracts and resists, ut then rela'es. i. 3his response allo"s hollo" or!ans such as the urinary ladder to fill, "ithout the smooth muscle in the "alls e'pellin! the contents immediately. ii. Smooth muscle is also not su 1ect to the limitations of the len!th/ tension relationships as is skeletal muscle, so that even "hen stretched, smooth muscle can contract forcefully, such as "hen emptyin! a full urinary ladder. c. 3hree features contri ute to smooth muscleJs a ility to stretch e'tensively and then contract po"erfullyF i. It has no > discs, so thick filaments do not utt up a!ainst them and stop contractin!. ii. 3he thick and thin filaments are not arran!ed in orderly sarcomeres, so havin! too little overlap for cross7 rid!in! does not occur. iii. 3he thick filaments have myosin heads alon! their entire len!th, "ith no are zone, so cross rid!es can form any"here. d. Smooth muscle also e'hi its plasticityIthe a ility to ad1ust its tension to the de!ree of stretch, an or!an such as the ladder can e !reatly stretched yet not ecome fla y "hen empty. +. 3he muscular system suffers fe" diseases ut does have some common dysfunctions. (3a le %%.6) Insight 11.5 $uscular 8ystrophy and $yasthenia =ravis(0i!. %%.(-)

#ross &eferences
Additional information on topics mentioned in #hapter %% can e found in the chapters listed elo". #hapter (F Anaero ic and aero ic A35 synthesis #hapter )F 3he sodium/potassium pump #hapter -F 3he three types of muscular tissue

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#hapter %0F #omposition of a skeletal muscle #hapter %(F ?eurotransmitters #hapter %(F +lectrophysiolo!y #hapter %(F Action potentials #hapter %(F All7or7none la" #hapter %DF #ardiac muscle #hapter (DF +ffects of a!in! on the muscular system

5a!e (0