Chapter 8 Skeletal Muscle: Structure and Function EXERCISE PHYSIOLOGY Theory and Application to Fitness and Performance, 6th edition Scott K. Powers & Edward T. Howley Skeletal Muscle • Human body contains over 400 skeletal muscles – 40-50% of total body weight • Functions of skeletal muscle – Force production for locomotion and breathing – Force production for postural support – Heat production during cold stress Connective Tissue Covering Skeletal Muscle • Epimysium – Surrounds entire muscle • Perimysium – Surrounds bundles of muscle fibers • Fascicles • Endomysium – Surrounds individual muscle fibers Connective Tissue Surrounding Skeletal Muscle Figure 8.1 Microstructure of Skeletal Muscle • Sarcolemma – Muscle cell membrane • Myofibrils – Threadlike strands within muscle fibers • Actin (thin filament) • Myosin (thick filament) • Sarcomere – Includes Z-line, M-line, H-zone, A-band, I-band • Sarcoplasmic reticulum – Storage sites for calcium • Transverse tubules Microstructure of Skeletal Muscle Figure 8.2 The Sarcoplasmic Reticulum and Transverse Tubules Figure 8.3 The Neuromuscular Junction • Junction between motor neuron and muscle fiber • Motor end plate – Pocket formed around motor neuron by sarcolemma • Neuromuscular cleft – Short gap between neuron and muscle fiber • Acetylcholine is released from the motor neuron – Causes an end-plate potential (EPP) • Depolarization of muscle fiber The Neuromuscular Junction Figure 8.4 Muscular Contraction • The sliding filament model – Muscle shortening occurs due to the movement of the actin filament over the myosin filament – Formation of cross-bridges between actin and myosin filaments • Power stroke – Reduction in the distance between Z-lines of the sarcomere The Sliding Filament Theory Figure 8.5 The Relationships Among Troponin, Tropomyosin, Myosin, and Calcium Figure 8.6 Energy for Muscle Contraction • ATP is required for muscle contraction – Myosin ATPase breaks down ATP as fiber contracts • Sources of ATP – Phosphocreatine (PC) – Glycolysis – Oxidative phosphorylation Sources of ATP for Muscle Contraction Figure 8.7 Excitation-Contraction Coupling • Depolarization of motor end plate (excitation) is coupled to muscular contraction – Action potential travels down T-tubules and causes release of Ca+2 from SR – Ca+2 binds to troponin and causes position change in tropomyosin • Exposing active sites on actin – Strong binding state formed between actin and myosin – Contraction occurs Muscle Excitation, Contraction, and Relaxation Figure 8.9 Steps Leading to Muscular Contraction Figure 8.10 Muscle Fatigue • Decrease in muscle force production – Reduced ability to perform work • Contributing factors: – High-intensity exercise (~60 sec) • Accumulation of lactate, H+, ADP, Pi, and free radicals – Long-duration exercise (2–4 hours) • Muscle factors – Accumulation of free radicals – Electrolyte imbalance – Glycogen depletion • Central Fatigue – Reduced motor drive to muscle from CNS Muscle Fatigue Figure 8.8 Characteristics of Muscle Fiber Types • Biochemical properties – Oxidative capacity • Number of capillaries, mitochondria, and amount of myoglobin – Type of myosin ATPase • Speed of ATP degradation • Contractile properties – Maximal force production • Force per unit of cross-sectional area – Speed of contraction (Vmax) • Myosin ATPase activity – Muscle fiber efficiency Characteristics of Individual Fiber Types • Type IIx fibers – Fast-twitch fibers – Fast-glycolytic fibers • Type IIa fibers – Intermediate fibers – Fast-oxidative glycolytic fibers • Type I fibers – Slow-twitch fibers – Slow-oxidative fibers Characteristics of Muscle Fiber Types Table 8.1 Comparison of Maximal Shortening Velocities Between Fiber Types Figure 8.12 Histochemical Staining of Fiber Type Type I Type IIa Type IIx Figure 8.11 Fiber Types and Performance • Nonathletes – Have about 50% slow and 50% fast fibers • Power athletes – Sprinters – Higher percentage of fast fibers • Endurance athletes – Distance runners – Higher percentage of slow fibers Exercise-Induced Changes in Skeletal Muscles • Strength training – Increase in muscle fiber size (hypertrophy) – Increase in muscle fiber number (hyperplasia) • Endurance training – Increase in oxidative capacity • Alteration in fiber type with training – Fast-to-slow shift • Type IIx IIa • Type IIa I with further training – Seen with endurance and resistance training Effects of Endurance Training on Fiber Type Figure 8.13 Muscle Atrophy Due to Inactivity • Loss of muscle mass and strength – Due to prolonged bed rest, limb immobilization, reduced loading during space flight • Initial atrophy (2 days) – Due to decreased protein synthesis • Further atrophy – Due to reduced protein synthesis • Atrophy is not permanent – Can be reversed by resistance training – During spaceflight, atrophy can be prevented by resistance exercise Age-Related Changes in Skeletal Muscle • Aging is associated with a loss of muscle mass – 10% muscle mass lost between age 25–50 y – Additional 40% lost between age 50–80 y – Also a loss of fast fibers and gain in slow fibers – Also due to reduced physical activity • Regular exercise training can improve strength and endurance – Cannot completely eliminate the age-related loss in muscle mass Types of Muscle Contraction • Isometric – Muscle exerts force without changing length – Pulling against immovable object – Postural muscles • Isotonic (dynamic) – Concentric • Muscle shortens during force production – Eccentric • Muscle produces force but length increases Muscle Actions Type of exercise Muscle Muscle Action Length Change _________________________________________________ Dynamic Concentric Decreases Eccentric Increases Static Isometric No Change Table 8.3 Isometric and Isotonic Contractions Figure 8.14 Speed of Muscle Contraction and Relaxation • Muscle twitch – Contraction as the result of a single stimulus – Latent period • Lasting ~5 ms – Contraction • Tension is developed • 40 ms – Relaxation • 50 ms • Speed of shortening is greater in fast fibers – SR releases Ca+2 at a faster rate – Higher ATPase activity Muscle Twitch Figure 8.15 Force Regulation in Muscle • Force generation depends on: – Types and number of motor units recruited • More motor units = greater force • Fast motor units = greater force • Initial muscle length – “Ideal” length for force generation – Increased cross-bridge formation • Nature of the neural stimulation of motor units – Frequency of stimulation • Simple twitch • Summation • Tetanus Relationship Between Stimulus Strength and Force of Contraction Figure 8.16 LengthTension Relationships in Skeletal Muscle Figure 8.17 Simple Twitch, Summation, and Tetanus Figure 8.18 Force-Velocity Relationship • At any absolute force the speed of movement is greater in muscle with higher percent of fast-twitch fibers • The maximum velocity of shortening is greatest at the lowest force – True for both slow and fast-twitch fibers Muscle Force-Velocity Relationships Figure 8.19 Force-Power Relationship • At any given velocity of movement the power generated is greater in a muscle with a higher percent of fast-twitch fibers • The peak power increases with velocity up to movement speed of 200-300 degrees•second-1 – Power decreases beyond this velocity because force decreases with increasing movement speed Muscle Force-Power Relationships Figure 8.20 Receptors in Muscle • Provide sensory feedback to nervous system – Tension development by muscle – Account of muscle length • Muscle spindle • Golgi tendon organ Muscle Spindle • Responds to changes in muscle length • Consists of: – Intrafusal fibers • Run parallel to normal muscle fibers (extrafusal fibers) – Gamma motor neuron • Stimulate intrafusal fibers to contract with extrafusal fibers (by alpha motor neuron) • Stretch reflex – Stretch on muscle causes reflex contraction • Knee-jerk reflex Muscle Spindles Figure 8.21 Golgi Tendon Organ (GTO) • Monitor tension developed in muscle – Prevents muscle damage during excessive force generation • Stimulation results in reflex relaxation of muscle – Inhibitory neurons send IPSPs to muscle fibers Golgi Tendon Organ Figure 8.22