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Chapter 8 The Muscular System Copyright 2010, John Wiley & Sons, Inc. Chapter 9 Muscular System Muscle Types Skeletal Smooth Cardiac Similarities All muscle cells are elongated = muscle fibers Muscle contraction depends on 2 kinds of myofilaments (actin & myosin) Cell membrane of a muscle cell = sarcolemma – cytoplasm is called sarcoplasm Copyright 2010, John Wiley & Sons, Inc. Types of Muscle and Function Skeletal - 40–50% of total body weight- voluntary Cardiac - involuntary Mostly movement of bone & body parts Stabilizing body positions Heart only Develops pressure for arterial blood flow Smooth- grouped in walls of hollow organs Sphincters regulate flow in tubes Maintain diameter of tubes Move material in GI tract and reproductive organs Copyright 2010, John Wiley & Sons, Inc. Muscle Functions Produce body movements Stabilize body positions Store and move substances Produce heat Copyright 2010, John Wiley & Sons, Inc. Functional Characteristics of Muscle Excitability – to receive & respond to stimuli Contractility – shorten forcibly when stimulated Extensibility – stretched or extended Elasticity – to bounce back to original length Copyright 2010, John Wiley & Sons, Inc. Skeletal Muscle Tissue Muscle includes: muscle fibers, connective tissue, nerves & blood vessels Copyright 2010, John Wiley & Sons, Inc. Connective Tissue Coverings Endomysium – thin delicate layer of CT that wraps each muscle fiber Fascicles – many muscle fibers bundled together into groups Skeletal muscle – many fascicles bundled together Each is wrapped in a 2nd layer of CT made of collagen – perimysium Each is covered by a 3rd layer of dense fibrous CT – epimysium Deep Fascia – each skeletal muscle is then covered by a 4th , very tough fibrous layer of CT May extend past the length of the muscle (tendon or aponeurosis) and attach that muscle to a bone, cartilage or muscle Copyright 2010, John Wiley & Sons, Inc. Structure of a Skeletal Muscle Outside to Inside • epimysium • perimysium • fascicle • endomysium Largest to Smallest • muscle • fascicles • muscle fibers • myofibrils • thick and thin filaments Copyright 2010, John Wiley & Sons, Inc. 9-4 Skeletal Muscle Tissue Copyright 2010, John Wiley & Sons, Inc. Muscle Histology Sarcoplasm contains myoglobin Red pigmented protein related to Hemoglobin that carries oxygen Along entire length are myofibrils Myofibrils made of protein filaments Come in thick and thin filaments Copyright 2010, John Wiley & Sons, Inc. Skeletal Muscle Fibers 2 types of protein filaments (cytoskeletal) Striations are caused by the arrangement of thick & thin filaments Thick – protein myosin Thin – protein actin A-Band – dark area – overlapping of thick & thin I-Band – light area – thin filaments only Length of each myofibril is divided into sarcomeres Sarcomeres meet one another at an area called – Z-line Copyright 2010, John Wiley & Sons, Inc. Sarcomere • I band • A band • H zone • Z line • M line Copyright 2010, John Wiley & Sons, Inc. 9-6 Muscle Histology Copyright 2010, John Wiley & Sons, Inc. Sarcomere Copyright 2010, John Wiley & Sons, Inc. Thick filaments – myosin Rod-like tail (axis) terminates in 2 globular heads or cross bridges Cross bridges interact with active sites on thin filaments Thin filaments – actin Coiled helical structure (resembles twisted strands of pearls) Tropomyosin – rod-shaped protein spiraling around actin backbone to stabilize it Troponin – complex polypeptides One binds to actin One that binds to tropomyosin One that binds to calcium ions Both help control actin’s interaction with myosin during contraction Copyright 2010, John Wiley & Sons, Inc. Myofilaments Copyright 2010, John Wiley & Sons, Inc. 9-7 Sarcomere Copyright 2010, John Wiley & Sons, Inc. Sarcomere Copyright 2010, John Wiley & Sons, Inc. Sarcomere Copyright 2010, John Wiley & Sons, Inc. Functional Structure Tropomyosin blocks myosin binding site when muscle is at rest Copyright 2010, John Wiley & Sons, Inc. Within sarcoplasm – 2 specialized membranous organelles Sarcoplasmic reticulum (SR) Network of membranous channels that surrounds each myofibril & runs parallel to it Same as ER in other cells SR has high concentrations of Ca ions compared to the sarcoplasm (maintained by active transport – calcium pump) When stimulated by muscle impulse, membranes become more permeable to Ca ions and Ca diffuses out of SR & into sarcoplasm Copyright 2010, John Wiley & Sons, Inc. Within sarcoplasm – 2 specialized membranous organelles Transverse tubules (TT) Set of membranous channels that extends into the sarcoplasm as invaginations continuous with muscle cell membrane (sarcolemma) TT’s are filled with extracellular fluid & extend deep into the cell Each TT runs between 2 enlarged portions of SR – cisternae Form a triad near the region where actin & myosin overlap Copyright 2010, John Wiley & Sons, Inc. Skeletal Muscle Contraction Neuromuscular Junction (NMJ) – site where a motor nerve fiber & a skeletal muscle fiber meet (synapse or synaptic cleft) Fibers must first be stimulated by a motor neuron for a skeletal muscle to contract Motor Unit – 1 motor neuron & many skeletal muscle fibers Motor End Plate – specific part of a skeletal muscle fiber’s sarcolemma directly beneath the NMJ Neurotransmitter – chemical substance released from a motor end fiber, causing stimulation of the sarcolemma of muscle fiber – acetylcholine (ACh) Synaptic Cleft - small space between neuron & muscle Copyright 2010, John Wiley & Sons, Inc. Axon collateral of somatic motor neuron Axon terminal Nerve impulse Synaptic vesicle containing acetylcholine (ACh) Synaptic end bulb Sarcolemma Axon terminal Synaptic end bulb Motor end plate Neuromuscular junction (NMJ) Synaptic cleft (space) Sarcolemma Myofibril (b) Enlarged view of the neuromuscular junction (a) Neuromuscular junction 1 1ACh is released from synaptic vesicle Synaptic end bulb Synaptic cleft (space) 2 2 ACh binds to Ach receptor Motor end plate Na+ Junctional fold 3 Muscle action potential is produced Copyright 2010, John Wiley & Sons, Inc. (c) Binding of acetylcholine to ACh receptors in the motor end plate Stimulus for contraction Begins – motor impulse reaches the end of the motor nerve fiber/ending – membrane depolarized (-70mV to -55mV) Calcium ions rush into motor nerve fiber Neurotransmitter (acetylcholine) released in to NMJ (exocytosis) Acetylcholine diffuses across the NMJ & stimulates/depolarizes the motor end-plate (sarcolemma) 100mV to -70mV The muscle impulse travels over the surface of the skeletal muscle fiber & deep into the muscle fiber by means of the TT Copyright 2010, John Wiley & Sons, Inc. Excitation Contraction Coupling • muscle impulses cause sarcoplasmic reticulum to release calcium ions into sarcoplasm of the muscle fiber • calcium binds to troponin to change its shape, moving tropomyosin and exposing myosin binding sites on actin filament • cross bridges (linkages) form between actin & myosin •Actin filaments are pulled inward by myosin crossbridges •Muscle fiber shortens as contraction occurs Copyright 2010, John Wiley & Sons, Inc. 9-11 Neuromuscular Junction Synaptic end bulbs (at neuron terminal) Release neurotransmitter Copyright 2010, John Wiley & Sons, Inc. Neuromuscular Junction Copyright 2010, John Wiley & Sons, Inc. Neuromuscular Junctions Interactions Animations Neuromuscular Junctions You must be connected to the internet to run this animation. Copyright 2010, John Wiley & Sons, Inc. Contraction Cycle Myosin binds to actin & releases phosphate group (forming crossbridges) Crossbridge swivels releasing ADP and shortening sarcomere (power stroke) ATP binds to Myosin → release of myosin from actin ATP broken down to ADP & Pi → activates myosin head to bind and start again Repeats as long as Ca2+ concentration is high Copyright 2010, John Wiley & Sons, Inc. Contraction Cycle Copyright 2010, John Wiley & Sons, Inc. Sliding Filament Theory Changes in muscle during contraction H zones and the I bands narrow Regions of overlap widen Z lines move closer together Shortening the sarcomere Copyright 2010, John Wiley & Sons, Inc. Cross-bridge Cycling When calcium ions are present, myosin binding sites on actin are exposed Cross-bridge attaches ATP breakdown provides E to “cock” myosin head “Cocked” myosin attaches to exposed actin binding site Cross-bridge (myosin head) springs from cocked position and pulls on actin filament Cross-bridges break ATP binds to cross-bridge (but is not yet broken down) Myosin heads are released from actin *As long as Ca ions and ATP are present, this walking continues until muscle fiber is fully contracted Copyright 2010, John Wiley & Sons, Inc. 9-13 Relaxation • acetylcholinesterase – enzyme present in NMJ • breaks down acetylcholine, so the motor end-plate is no longer stimulated (cannot cause continuous muscle contraction) • muscle impulse stops • calcium moves back from sarcoplasm into sarcoplasmic reticulum • myosin and actin binding sites are broken •Muscle fiber relaxes Copyright 2010, John Wiley & Sons, Inc. 9-14 Energy Sources for Contraction The E used to power the interaction between actin & myosin comes from ATP ATP stored in skeletal muscle lasts only about six seconds ATP must be regenerated continuously if contraction is to continue 3 pathways in which ATP is regenerated Coupled reaction with Creatine Phosphate (CP) Anaerobic Cellular Respiration Aerobic Cellular Respiration Copyright 2010, John Wiley & Sons, Inc. Energy Sources for Contraction CP + ADP creatine + ATP • creatine phosphate – stores energy that quickly converts ADP to ATP Copyright 2010, John Wiley & Sons, Inc. 9-15 Oxygen Supply and Cellular Respiration • Anaerobic Phase • glycolysis • occur in cytoplasm of cell • produces pyruvic acid & little ATP • Aerobic Phase • citric acid cycle and electron transport chain • occurs in mitochondria • produces CO2 & most ATP • myoglobin stores extra oxygen Copyright 2010, John Wiley & Sons, Inc. 9-16 Muscle Fatigue Physiological inability to contract No O2 is available in muscle cells to complete aerobic respiration Pyruvic acid is converted to lactic acid Muscle fatigue & soreness Results form a relative deficit of ATP and/or accumulation of lactic acid (decreases pH) Copyright 2010, John Wiley & Sons, Inc. Muscle Fatigue •cramp – sustained, involuntary contraction Copyright 2010, John Wiley & Sons, Inc. 9-18 Oxygen Debt Oxygen debt – amount of oxygen needed by liver to convert lactic acid to glucose Copyright 2010, John Wiley & Sons, Inc. 9-17 Heat Production • Almost half of E released during muscle contraction is lost to heat, which helps maintain our body temperature at 37 degrees celsius • by-product of cellular respiration • muscle cells are major source of body heat • blood transports heat throughout body • Heat lost through negative feedback mechanisms – sweating, dilation of superficial blood vessels, increased breathing rate & increased heart rate Copyright 2010, John Wiley & Sons, Inc. 9-19 Muscle Tone Even at rest some motor neuron activity occurs = Muscle Tone If nerves are cut fiber becomes flaccid (very limp) Copyright 2010, John Wiley & Sons, Inc. Metabolism Rapid changes from very low ATP consumption to high levels of consumption Creatine phosphate (high energy store) Fast and good for ~ 15 sec Copyright 2010, John Wiley & Sons, Inc. Nerve impulse 1 Nerve impulse arrives at axon terminal of motor neuron and triggers release of acetylcholine (ACh). 2 ACh diffuses across synaptic cleft, binds to its receptors in the motor end plate, and triggers a muscle action potential (AP). ACh receptor 3 Acetylcholinesterase in Synaptic vesicle synaptic cleft destroys filled with ACh ACh so another muscle action potential does not arise unless more ACh is released from motor neuron. Muscle action potential Transverse tubule 4 Muscle AP travelling along transverse tubule opens Ca2+ release channels in the sarcoplasmic reticulum (SR) membrane, which allows calcium ions to flood into the sarcoplasm. SR Ca2+ 9 Muscle relaxes. 8 Troponin–tropomyosin complex slides back into position where it blocks the myosin binding sites on actin. 5 Ca2+ binds to troponin on the thin filament, exposing the binding sites for myosin. Elevated Ca2+ Ca2+ active transport pumps 7 Ca2+ release channels in SR close and Ca2+ active transport pumps use ATP to restore low level of Ca2+ in sarcoplasm. 6 Contraction: power strokes use ATP; myosin heads bind to actin, swivel, and release; thin filaments are pulled toward center of sarcomere. Copyright 2010, John Wiley & Sons, Inc. Production of ATP for Muscle Contraction Copyright 2010, John Wiley & Sons, Inc. Glycolysis Break down glucose to 2 pyruvates getting 2 ATPs If insufficient mitochondria or oxygen, pyruvate → lactic acid Get about 30–40 seconds more activity maximally Copyright 2010, John Wiley & Sons, Inc. Production of ATP for Muscle Contraction Copyright 2010, John Wiley & Sons, Inc. Aerobic Cellular Respiration Production of ATP in mitochondria Requires oxygen and carbon substrate Produces CO2 and H2O and heat. Copyright 2010, John Wiley & Sons, Inc. Fatigue Inability to contract forcefully after prolonged activity Limiting factors can include: Ca2+ Creatine Phosphate Oxygen Build up of acid Neuronal failure Copyright 2010, John Wiley & Sons, Inc. Control of Muscle Contraction Single action potential(AP) → twitch Total tension of fiber depends on frequency of APs (number/second) Smaller than maximum muscle force Require wave summation Maximum = tetanus Total tension of muscle depends on number of fibers contracting in unison Increasing numbers = Motor unit recruitment Copyright 2010, John Wiley & Sons, Inc. Muscular Responses Threshold Stimulus • minimal strength required to cause contraction Copyright 2010, John Wiley & Sons, Inc. 9-20 Recording a Muscle Contraction Myogram – recording of a muscle contraction Twitch- single contraction that lasts a fraction of a second, followed by relaxation latent period – delay between stimulation & contraction refractory period – muscle fiber must return to its resting state (-100mV) before it can be stimulated again all-or-none response If a muscle fiber is brought to threshold or above, it responds with a complete twitch If the stimulus is sub-threshold, the muscle fiber will not respond Copyright 2010, John Wiley & Sons, Inc. Myogram Copyright 2010, John Wiley & Sons, Inc. Myogram Copyright 2010, John Wiley & Sons, Inc. Staircase Effect (treppe) Most muscle fiber contraction is “all or nothing” A muscle fiber that has been inactive can be subjected to a series of stimuli & The fiber undergoes a series of twitches with relaxation between & The strength of each successive contraction increases slightly Phenomenon is small & brief & involves excess calcium in sarcoplasm Copyright 2010, John Wiley & Sons, Inc. Summation •When several stimuli are delivered in succession to a muscle fiber, it cannot completely relax between contractions •individual twitches combine and muscle contraction becomes sustained •When resulting sustained contraction lacks even slight relaxation, it is called tetanic contractions Copyright 2010, John Wiley & Sons, Inc. 9-21 Motor Units Motor neuron & the many skeletal muscle fibers it stimulates Motor neuron branches into several motor nerve endings, it can stimulate many skeletal muscles fibers simultaneously, which then contract simultaneously The # of muscle fibers in a motor unit varies from 10 - hundreds Copyright 2010, John Wiley & Sons, Inc. Recruitment of Motor Units • recruitment - increase in the number of motor units activated • whole muscle composed of many motor units, controlled by many different motor neurons, simultaneous contraction of all units does not necessarily occur • as intensity of stimulation increases, recruitment of motor units continues until all motor units are activated Copyright 2010, John Wiley & Sons, Inc. 9-22 Sustained Contractions •Even when a muscle is at rest, a certain amount of sustained contraction is occurring in its fibers - muscle tone • very important in maintaining posture Copyright 2010, John Wiley & Sons, Inc. 9-23 Types of Contractions • concentric – shortening contraction • eccentric – lengthening contraction • isometric – muscle contracts but does not change length – attachments do not move, tensing a muscle Copyright 2010, John Wiley & Sons, Inc. 9-24 Fiber Types Slow oxidative (SO)- small diameter and red Fast oxidative- glycolytic (FOG) Large amounts of myoglobin and mitochondria ATP production primarily oxidative Fatigue resistant Large diameter = many myofibrils Many mitochondria and high glycolytic capacity Fast glycolytic fibers (FG) White, fast & powerful and fast fatiguing For strong, short term use Copyright 2010, John Wiley & Sons, Inc. Recruitment Recruited in order: SO → FOG → FG Copyright 2010, John Wiley & Sons, Inc. Fast and Slow Twitch Muscle Fibers Fast-twitch fatigueSlow-twitch fibers (type I) resistant fibers (type IIb) • always oxidative • intermediate fibers • resistant to fatigue • oxidative • red fibers • oxygen containing pigment - myoglobin • intermediate amount of myoglobin • good blood supply • pink to red in color • contain many mitochondria Fast-twitch glycolytic fibers (type II) • white fibers (less myoglobin) • fewer mitochondria • contain extensive sarcoplasmic reticulum to store and reabsorb calcium • poorer blood supply • contract rapidly, but fatigue easily due to lactic acid accumulation Copyright 2010, John Wiley & Sons, Inc. 9-25 Effects of Exercise SO/FG fiber ratio genetically determined Endurance exercise gives FG → FOG High FG → sprinters High SO → marathoners Increased diameter and numbers of mitochondria Strength exercise increases size and strength of FG fibers Copyright 2010, John Wiley & Sons, Inc. Cardiac Muscle Location • only in the heart Anatomy •Arrangement of actin & myosin not as organized as skeletal Physiology • Self-exciting tissue (pacemaker) • Involuntary control – all or nothing contractions •Function as a “syncytium” (all or nothing) • rhythmic contractions (60-100 beats/minute) • longer refractory period than skeletal muscle •Pumps blood to •Lungs for oxygenation •Body for distribution of oxygen & nutrients Copyright 2010, John Wiley & Sons, Inc. 9-29 Cardiac Muscle Involuntary muscle found only in heart wall Striated, branched short fibers with single, central nucleus in each fiber Fibers connected by: Intercalated discs (thickened cell membranes) Gap junctions that allow spread of action potentials ATP generated by abundant mitochondria and by lactic acid when cells lack oxygen Copyright 2010, John Wiley & Sons, Inc. Cardiac Muscle Does not require nerve stimulation nerve Has its own intrinsic pacemaker (and conduction system within cardiac muscle) that initiates cardiac contraction Known as auto-rhythmicity Ca2+released from S.R. and extracellular spaces Intercalated discs with gap junctions transmit action potentials from ne muscle cell to the next Copyright 2010, John Wiley & Sons, Inc. Cardiac Muscle Copyright 2010, John Wiley & Sons, Inc. Smooth Muscle Fibers Compared to skeletal muscle fibers • shorter • single nucleus • elongated with tapering ends • myofilaments randomly organized • no striations • lack transverse tubules • sarcoplasmic reticula are reduced • walls of hollow visceral organs & blood vessels • Involuntary control • Contractions are slow & sustained • troponin is absent Copyright 2010, John Wiley & Sons, Inc. 9-26 Types of Smooth Muscle Visceral Smooth Muscle • single-unit smooth muscle • sheets of muscle fibers • fibers held together by gap junctions • exhibit rhythmicity – pattern of repeated contractions • exhibit peristalsis – wave-like motion that helps push substances through passageways • walls of most hollow organs • contraction is slow & sustained •Structure •Random arrangement of actin & myosin filaments •2 layers of muscle surround passageway •Inner circular layer •Outer longitudinal layer Multiunit Smooth Muscle • fibers function separately • irises of eye • walls of blood vessels • contraction is rapid & vigorous (similar to skeletal muscle tissue Copyright 2010, John Wiley & Sons, Inc. 9-27 Smooth Muscle Contraction Resembles skeletal muscle contraction • interaction between actin and myosin • both use calcium and ATP • both depend on impulses Different from skeletal muscle contraction • A protein, calmodulin binds to calcium ions (no troponin) and activates the contraction mechanism • most calcium diffuses in to smooth muscle cells from the extracellular fluid (reduced SR) • two neurotransmitters affect smooth muscle • acetlycholine and norepinephrine • hormones affect smooth muscle • stretching can trigger smooth muscle contraction • contraction is slow & sustained • smooth muscle more resistant to fatigue 9-28 Copyright 2010, John Wiley & Sons, Inc. Smooth Muscle Involuntary Found in internal organs such as stomach, bladder, walls of arteries Structure Tapered cells each with single nucleus Filaments not regular so tissue does not appear striated Copyright 2010, John Wiley & Sons, Inc. Smooth Muscle Types Visceral (single unit) type or Multi-unit type Form sheets and are auto-rhythmic Contract as a unit Each has own nerve and can contract independently Graded contractions and slow responses Often sustain long term tone Often triggered by autonomic nerves Modulated chemically, by nerves, by mechanical events (stretching) Copyright 2010, John Wiley & Sons, Inc. Smooth Muscle Copyright 2010, John Wiley & Sons, Inc. Life-Span Changes • myoglobin, ATP, and creatine phosphate decline in ones forties • by age 80, half of muscle mass has atrophied – replaced by connective & adipose tissue •Reflexes are reduced • exercise helps to maintain muscle mass and function Copyright 2010, John Wiley & Sons, Inc. 9-65 Aging As with bone there is a slow progressive loss of skeletal muscle mass Relative number of SO fibers tends to increase Copyright 2010, John Wiley & Sons, Inc. Homeostatic Imbalances/Disorders Tendinitis Compartment Syndrome Poliomyelitis Myasthenia Gravis Duchenne Muscular Dystrophy Rigor Mortis Botulism TMJ Parkinson’s Disease Copyright 2010, John Wiley & Sons, Inc. Clinical Application Myasthenia Gravis • autoimmune disorder • receptors for acetylcholine on muscle cells are attacked • weak and easily fatigued muscles result • difficulty swallowing and chewing • ventilator needed if respiratory muscles are affected • treatments include • drugs that boost acetylcholine • removing thymus gland • immunosuppressant drugs • antibodies Copyright 2010, John Wiley & Sons, Inc. 9-66 Movement Muscles move one bone relative to another around one or more joint(s) Origin → most stationary end Insertion → most mobile end Location where the tendon attaches Location where tendon inserts Action → the motion or function of the muscle Copyright 2010, John Wiley & Sons, Inc. Movement Generally arranged in opposing pairs Flexors - extensors; abductors - adductors The major actor: prime mover or agonist Muscle with opposite effect: antagonist Synergists - help prime mover Fixators - stabilize origin of prime mover Role of muscle varies with motion Copyright 2010, John Wiley & Sons, Inc. Basis of Muscle Names: Table 8.2 Direction of fibers relative to body axes Size of muscle Examples: lateralis, medialis (medius), intermedius, rectus Examples: alba, brevis, longus, magnus, vastus Shape of muscle Examples: deltoid, orbicularis, serratus, trapezius Copyright 2010, John Wiley & Sons, Inc. Basis of Muscle Names: Table 8.2 Action of muscle Number of tendons (heads) of origin Examples: abductor, adductor, flexor, extensor Examples: biceps, triceps, quadriceps Location of muscle Examples: abdominus, brachialis, cleido, oculo-, uro-, Copyright 2010, John Wiley & Sons, Inc.