Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Function of Muscles Produce movement Maintain posture Stabilize joints Generate heat Skeletal Muscle: Attachments Muscles attach: • Directly—epimysium of muscle is fused to the periosteum of bone or perichondrium of cartilage • Indirectly—connective tissue wrappings extend beyond the muscle as a ropelike tendon or sheetlike aponeurosis The Muscular System Muscles are responsible for all types of body movement Three basic muscle types are found in the body Characteristics of Muscles Muscle cells are elongated (muscle cell = muscle fiber) Contraction of muscles is due to the movement of microfilaments within fiber cells All muscles share some terminology • Prefix myo refers to muscle • Prefix mys refers to muscle • Prefix sarco refers to flesh Table 9.3 Skeletal Muscle Characteristics Most are attached by tendons to bones Cells are multinucleate Striated – have visible banding Voluntary – subject to conscious control Cells are cylindrical Cells are surrounded and bundled by connective tissue Plasma/cell membrane called a sarcolemma Glycosomes for glycogen storage, myoglobin for O2 storage Also contain myofibrils, sarcoplasmic reticulum (modified ER), and T tubules Smooth Muscle Characteristics Has no striations Spindle-shaped cells Single nucleus Involuntary – no conscious control Found mainly in the walls of hollow organs Cardiac Muscle Characteristics Has striations Usually has a single nucleus Joined to another muscle cell at an intercalated disc Involuntary Found only in the heart Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Table 9.1 Fib-Endo-Fas-Per-Ep Nested Structures in a Muscle Epimysium Bone Epimysium Perimysium Endomysium Tendon (b) Perimysium Fascicle Muscle fiber in middle of a fascicle Blood vessel Fascicle (wrapped by perimysium) Endomysium (between individual muscle fibers) Muscle fiber • Whole muscle is surrounded by an epimysium and is composed of wrapped fascicles • Each fascicle is composed of muscle fibers (cells), surrounded by a perimysium Figure 9.1 • Each muscle fiber is surrounded by endomysium ( & then the sarcolemma) • Muscle fibers (cells) contain several myofibrils Sarcolemma Mitochondrion Myofibril Dark A band Light I band Nucleus (b) Diagram of part of a muscle fiber showing the myofibrils. One myofibril is extended afrom the cut end of the fiber. Sarcoplasmic Reticulum is Modified Endoplasmic Reticulum (Storage Depot for Calcium Ions) Part of a skeletal muscle fiber (cell) I band A band I band Z disc H zone Z disc Myofibril M line Sarcolemma Sarcolemma (muscle fiber plasma membrane) Triad: • T tubule • Terminal cisternae of the SR (2) Tubules of the SR Myofibrils Mitochondria T tubules are continuous with the sarcolemma They penetrate the cell’s interior at each A band–I band junction They’re associated with the paired terminal cisternae to form triads that encircle each sarcomere T tubules conduct impulses deep into muscle fiber; contains gated channels that regulate Ca 2+ release Figure 9.5 Patterns Visible in the Sarcomere (Smallest Contractile Unit of a Myofibril) Thin (actin) filament Thick (myosin) filament Z disc I band H zone A band Sarcomere Z disc I band Sarcomere Z disc M line Z disc M line Microscopic Muscle Anatomy (online) Thin (actin) filament Elastic (titin) filaments Thick (myosin) filament Figure 9.2c, d Thin and Thick Filament Composition Longitudinal section of filaments within one sarcomere of a myofibril Thick filament Thin filament In the center of the sarcomere, the thick filaments lack myosin heads. Myosin heads are present only in areas of myosin-actin overlap. Thick filament Thin filament Each thick filament consists of many A thin filament consists of two strands myosin molecules whose heads protrude of actin subunits twisted into a helix at opposite ends of the filament. plus two types of regulatory proteins (troponin and tropomyosin). Portion of a thick filament Portion of a thin filament Myosin head Tropomyosin Troponin Actin Actin-binding sites ATPbinding site Heads Tail Flexible hinge region Myosin molecule Active sites for myosin attachment Actin subunits Actin subunits Tropomyosin and troponin: regulatory proteins bound to actin Figure 9.3 Z Z H A I I 1 Fully relaxed sarcomere of a muscle fiber Z I Z A I 2 Fully contracted sarcomere of a muscle fiber Figure 9.6 Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy How Muscle Contracts: 1) Events at the Neuromuscular Junction Action potential (AP) Myelinated axon of motor neuron Axon terminal of neuromuscular junction Nucleus Sarcolemma of the muscle fiber 1 Action potential arrives at axon terminal of motor neuron. 2 Voltage-gated Ca2+ channels open and Ca2+ enters the axon terminal. Ca2+ Ca2+ Axon terminal of motor neuron Synaptic vesicle containing ACh Mitochondrion Synaptic cleft Fusing synaptic vesicles Skeletal muscles are stimulated by the axon termini of somatic motor neurons Association of one axon to a particular group of fibers = 1 motor unit Figure 9.8 How Muscle Contracts: 1) Events at the Neuromuscular Junction Myelinated axon of motor neuron Axon terminal of neuromuscular junction Sarcolemma of the muscle fiber Action potential (AP) Nucleus 1 Action potential arrives at axon terminal of motor neuron. 2 Voltage-gated Ca2+ channels Ca2+ Ca2+ open and Ca2+ enters the axon terminal. Axon terminal of motor neuron 3 Ca2+ entry causes some synaptic vesicles to release their contents (acetylcholine) by exocytosis. Synaptic vesicle containing ACh Mitochondrion Synaptic cleft Fusing synaptic vesicles ACh 4 Acetylcholine, a neurotransmitter, diffuses across the synaptic cleft and binds to receptors in the sarcolemma. Junctional folds of sarcolemma Sarcoplasm of muscle fiber 5 ACh binding opens ion channels that allow simultaneous passage of Na+ into the muscle fiber and K+ out of the muscle fiber. 6 ACh effects are terminated by its enzymatic breakdown in the synaptic cleft by acetylcholinesterase. Nerve impulse causes muscular contraction: excitation-contraction (E-C) coupling Na+ Ach– K+ Degraded ACh Na+ Acetylcholinesterase K+ Postsynaptic membrane ion channel opens; ions pass. Events at neuromuscular junction movie Postsynaptic membrane ion channel closed; ions cannot pass. Figure 9.8 How Muscle Contracts: 2) Initiation of an Action Potential Axon terminal Open Na+ Channel Na+ Synaptic cleft Closed K+ Channel ACh ACh Na+ K+ Na+ K+ K+ ++ ++ + + Action potential + + +++ + 2 Generation and propagation of the action potential (AP) Closed Na+Channel 1 Local depolarization: generation of the end plate potential on the sarcolemma Sarcoplasm of muscle fiber Open K+ Channel Na+ K+ 3 Repolarization Figure 9.9 How Muscle Contracts: 3) How the Sarcolemma Resets Depolarization due to Na+ entry Na+ channels close, K+ channels open Repolarization due to K+ exit Na+ channels open Threshold K+ channels close Figure 9.10 Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy How Muscle Contracts: 4) Action Potential Causes Ca++ Release Axon terminal of motor neuron Action potential Synaptic cleft is generated ACh Sarcolemma Terminal cisterna of SR Muscle fiber Ca2+ Triad One sarcomere Figure 9.11, step 1 How Muscle Contracts: 4) Action Potential Causes Ca++ Release 1 Action potential is Steps in E-C Coupling: propagated along the sarcolemma and down the T tubules. Voltage-sensitive tubule protein Sarcolemma T tubule Ca2+ release channel Terminal cisterna of SR 2 Calcium ions are released. Ca2+ Figure 9.11, step 4 How Muscle Contracts: 5) Ca++ Binds to Troponin Actin Ca2+ Troponin Tropomyosin blocking active sites Myosin The aftermath Figure 9.11, step 5 How Muscle Contracts: 5) Ca++ Binds to Troponin Actin Ca2+ Troponin Tropomyosin blocking active sites Myosin 3 Calcium binds to troponin and removes the blocking action of tropomyosin. Active sites exposed and ready for myosin binding The aftermath Figure 9.11, step 6 How Muscle Contracts: 6) Troponin Slides Tropomyosin Off Myosin Binding Sites Actin Ca2+ Troponin Tropomyosin blocking active sites Myosin 3 Calcium binds to troponin and removes the blocking action of tropomyosin. Active sites exposed and ready for myosin binding 4 Contraction begins Myosin cross bridge The aftermath Calcium is sequestered again by the SR, lowering Ca++ levels, and causing muscle to relax as tropomyo. covers binding sites Figure 9.11, step 7 Four Step Power Cycle or “Cross Bridge Cycle” Thin filament Actin Ca2+ Myosin cross bridge ADP Pi Thick filament Myosin Cross bridge formation. 1 ADP ADP Pi Pi ATP hydrolysis 2 The power (working) stroke. 4 Cocking of myosin head. ATP ATP 3 Cross bridge detachment. Figure 9.12 Step One of the Cross Bridge Cycle Actin Ca2+ Myosin cross bridge Thin filament ADP Pi Thick filament Myosin 1 Cross bridge formation. Figure 9.12, step 1 Step Two of the Cross Bridge Cycle ADP Pi 2 The power (working) stroke. Figure 9.12, step 3 Step Three of the Cross Bridge Cycle ATP 3 Cross bridge detachment. Figure 9.12, step 4 Step Four of the Cross Bridge Cycle ADP ATP Pi hydrolysis 4 Cocking of myosin head. Figure 9.12, step 5 Summary of the Cross Bridge Cycle Thin filament Actin Ca2+ Myosin cross bridge ADP Pi Thick filament Myosin Cross bridge formation. 1 ADP ADP Pi Pi ATP hydrolysis 2 The power (working) stroke. 4 Cocking of myosin head. Sliding Filament Theory ATP ATP 3 Cross bridge detachment. Figure 9.12 The Power Cycle A. Masking protein complex (tropomyosin) binds Ca++ released from the SR moves aside to expose head-binding sites B. Steps of the Power Cycle 1. "Cocked" myosin head binds to actin myofilament site 2. Head bends towards the M line (sarcomere center-line), pulling thin filament along and releasing ADP and P (broken ATP) for the power stroke 3. ATP binds to the myosin head, causing it to detach 4. Myosin head “recocks” as ATP broken down to ADP and P Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Principles of Muscle Mechanics 1. The same principles apply to contraction of a single fiber and a whole muscle 2. Contraction produces tension, the force exerted on the load or object to be moved 3. 4. Contraction does not always shorten a muscle: • Isometric contraction: no shortening; muscle tension increases but does not exceed the load • Isotonic contraction: muscle shortens because muscle tension exceeds the load Force and duration of contraction vary in response to stimuli of different frequencies and intensities Muscle Twitch Response of a muscle to a single, brief threshold stimulus Simplest contraction observable in the lab (recorded as a myogram) Three phases of a twitch: • Latent period: events of excitation-contraction coupling • Period of contraction: cross bridge formation; tension increases • Period of relaxation: Ca2+ reentry into the SR; tension declines to zero ExcitationCross bridge contraction formation; tension coupling increases Latent Period of period contraction Ca2+ reentry into the SR; tension declines to zero Period of relaxation Single stimulus (a) Myogram showing the three phases of an isometric twitch Figure 9.14a Graded Muscle Responses Variations in the degree of muscle contraction Required for proper control of skeletal movement Responses are graded by: 1. Changing the frequency of stimulation 2. Changing the strength of the stimulus Response to Change in Stimulus Frequency A single stimulus results in a single contractile response—a muscle twitch Single stimulus single twitch Contraction Relaxation Stimulus A single stimulus is delivered. The muscle contracts and relaxes Response to Change in Stimulus Frequency Increases frequency of stimulus (muscle does not have time to completely relax between stimuli). Distinct peaks are still seen in the myogram. Low stimulation frequency --> unfused (incomplete) tetanus Partial relaxation Stimuli (b) If another stimulus is applied before the muscle relaxes completely, then more tension results. This is temporal (or wave) summation and results in unfused (or incomplete) tetanus. Figure 9.15b Response to Change in Stimulus Frequency Ca2+ release stimulates further contraction temporal (wave) summation Further increase in stimulus frequency unfused (incomplete) tetanus If stimuli are given quickly enough, fused (complete) tetany results High stimulation frequency fused (complete) tetanus Stimuli Figure 9.15c (c) At higher stimulus frequencies, there is no relaxation at all between stimuli. This is fused (complete) tetanus. Response to Change in Stimulus Strength Threshold stimulus: stimulus strength at which the first observable muscle contraction occurs Muscle contracts more vigorously as stimulus strength is increased above threshold Contraction force is precisely controlled by recruitment (multiple motor unit summation), which brings more and more muscle fibers into action Response to Change in Stimulus Strength Stimulus strength Maximal stimulus Threshold stimulus Proportion of motor units excited Strength of muscle contraction Maximal contraction Figure 9.16 Response to Change in Stimulus Strength Size principle: motor units with larger and larger fibers are recruited as stimulus intensity increases Motor Motor Motor unit 1 unit 2 unit 3 Recruitedrecruitedrecruited (small (medium (large fibers) fibers) fibers) Figure 9.17 Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Energy for Muscle Contraction Short-duration exercise ATP stored in muscles is used first. ATP is formed from creatine Phosphate and ADP. Glycogen stored in muscles is broken down to glucose, which is oxidized to generate ATP. Prolonged-duration exercise ATP is generated by breakdown of several nutrient energy fuels by aerobic pathway. This pathway uses oxygen released from myoglobin or delivered in the blood by hemoglobin. When it ends, the oxygen deficit is paid back. Figure 9.20 Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Isotonic Contractions Muscle changes in length and moves the load Isotonic contractions are either concentric or eccentric: • Concentric contractions—the muscle shortens and does work • Eccentric contractions—the muscle contracts as it lengthens Isometric Contractions The load is greater than the tension the muscle is able to develop Tension increases to the muscle’s capacity, but the muscle neither shortens nor lengthens Summary of Factors Increasing Contractile Force Large number of muscle fibers activated Large muscle fibers High frequency of stimulation Muscle and sarcomere stretched to slightly over 100% of resting length Contractile force Figure 9.21 Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Effects of Exercise on Muscle Results of increased muscle use • Increase in muscle size • Increase in muscle strength • Increase in muscle efficiency • Muscle becomes more fatigue resistant Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Muscle Fiber Types Developmental Aspects Muscular Dystrophy Muscle Fiber Type Fibers classified according to two characteristics: 1. Speed of contraction: slow or fast, according to: • Speed at which myosin ATPases split ATP • Pattern of electrical activity of the motor neurons 2. Metabolic pathways for ATP synthesis: • Slow and fast oxidative fibers—use aerobic pathways (fast oxidative fibers = red meat in birds) • Glycolytic fibers—use anaerobic glycolysis (these fibers are “fast twitch” white meat in birds) Predominance of fast glycolytic (fatigable) fibers: “fast twitch” or “white meat” Contractile velocity Small load Predominance of slow oxidative (fatigue-resistant) fibers: “slow twitch” or dark meat Contractile duration Figure 9.23 Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy Developmental Aspects Cardiac and skeletal muscle become amitotic, but can lengthen and thicken Injured heart muscle is mostly replaced by connective tissue Smooth muscle regenerates throughout life Myoblast-like skeletal muscle satellite cells have limited regenerative ability; are responsible for generating more fibers and in muscle repair Muscular development reflects neuromuscular coordination • Development occurs head to toe, and proximal to distal • Peak natural neural control occurs by midadolescence • Athletics and training can improve neuromuscular control Diseases and Medical Conditions (Myopathies) of the Muscular System • Myasthenia gravis (autoimmumity; destruction of ACh receptors so neuromuscular junctions don’t work) • Poliomyelitis (viral infection of muscle nerves) • Muscle strains cause myalgia (pain) and sometimes myositis (inflammation). Inflamed tendons are fibromyositis. • Fibromyalgia- (muscle pain) causes widespread pain in the muscles accompanied by fatigue and sleep disorders. Thought to be neurologically, blood flow, based. • Cramps (muscle spasms) • Contusion (muscle bruise) • Crush injury (severe trauma releasing myoglobin) • Muscular dystrophy (e.g. Duchenne's; degeneration & atrophy of muscles) Muscular Dystrophy Group of inherited muscle-destroying diseases Muscles enlarge due to fat and connective tissue deposits Muscle fibers atrophy Duchenne muscular dystrophy (DMD): • Most common and severe type • Inherited, sex-linked, carried by females and expressed in males (1/3500) as a lack of dystrophin, a protein that links muscle fibers together • Victims become clumsy and fall frequently; usually die of respiratory failure in their 20s • No cure, but viral gene therapy or infusion of stem cells with correct dystrophin genes show promise Muscle Types and Physiology Types and Characteristics of Muscle Muscle Function and Types Microscopic Anatomy of Muscle Muscular Stimulation Muscular Contraction Mechanism Muscular Response Based on Stimulus Energy Sources for Muscular Contraction Types of Muscular Contractions Effects of Exercise on Muscles Developmental Aspects Muscular Dystrophy