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SKELETAL MUSCLE PHYSIOLOGY Abraham D. Lee, Ph.D.,P.T. Department of Physical Therapy Office: Collier Building # 4206 Phone #: 419-383-3437 Email: [email protected] Contents 1. Muscle structure & organization 2. Muscle fiber type 3. Muscle action 4. Muscle mechanics 5. Motor unit and its recruitment 6. Local muscle control 7. Muscle plasticity 8. Summary Muscle organization • Epimysium: wraps an entire muscle • Perimysium: wraps a bundle of muscle fibers. This bundle is called fascicle or fasciculus • Endomysium: wraps an individual muscle fiber • Sarcolemma: muscle membrane • Myofibrils: contractile filaments Myofibrils • Thin filament – Actin filaments – Troponin – Tropomyosin • Thick filament – Myosin: 4 light chains and 2 heavy chains – Heavy chains • Myosin head region: heavy meromyosin • Myosin tail region: light memromysin Muscle pennation • Longitudinal (non-pennated) architecture: muscle fibers in parallel to the muscle force generating axis – – • Example: biceps brachii, sartorius muscle In these muscles fibers are said to be fusiform or spindle shaped. “Pennate” architecture: muscle fibers are oriented at an angle or multiple angles relative to forcegenerating axis. 1) Unipennate: soleus-25 degree; vastus medialis-5 degree 2) Bipennate: gastrocnemius, rectus femoris 3) Multipennate: deltoid Effect of pennation Force loss Space saving Comparison b/n non-pennated & pennated muscle Non-pennated Pennated •Contraction •Fiber packing w/ given volume •Force loss due to pennation Fast Less Slow More No Yes •# fiber Less More •Muscle force Production Less Greater •CSA Less Greater Muscle Fiber Type Muscle fiber type • Type I, – Slow-oxidative (SO) fibers • Type IIa, – Fast-oxidative-glycolytic (FOG) fibers • Type IIb, – Fast-glycolytic (FG) fibers Characteristics of different fibers #Mitochondria Resistance to fatigue Energy ATPase activity Vmax Efficiency • • • • • • • Type I H H A L L H L: low H: high M: moderate A: aerobic AN: anaerobic HH: highest Type IIa H/M H/M A+AN H H M Type IIb L L AN HH HH L Muscle composition in athletes % Type I %Type IIa &IIb Distance runners 70-80 20-30 Track sprinters 25-30 70-75 Weight lifters 45-55 45-55 Non-athletes 47-53 47-53 Will fiber type change with training? Muscle Action • Excitation-contraction coupling • Type of muscle action Excitation-Contraction Coupling •Nerve impulse generation and propagation •Neuromuscular junction transmission •Muscle action potential propagation •Ca2+ release from SR •Ca2+ binding to troponin •Interaction of myosin head and actin •Cross bridge moves: tension development •Ca2+ taken up to SR •Ca2+ removal from troponin •Relaxation E-C coupling DHPR: dihydropyridine receptors RyR: ryanodine receptor Other possible mechanism: Inositol 1,4,5-triphosphate (InsP3) InsP3 receptor activation Ca2+ release from SR May play a role in slow twitch muscle in developmental stage (Talon et al., Am. J. Physiol 282: R1164-R1173, 2002) E-C coupling Sliding Filament Theory Changes during shortening muscle action • Sarcomere length (distance between two adjacent Z lines): shortens • A band: no change • I band: shortens • H zone: shortens Different type of muscle action (contraction) • Isometric action • Isotonic action (dynamic action) – Concentric action – Eccentric action Muscle mechanics It deals with how muscle force is generated and regulated. Factors that affect muscle force generation • • • • • • Rate of muscle stimulation Muscle length Joint angle Speed of action (speed of contraction) Muscle fiber type # of MU recruitment Rate of Muscle Stimulation • Twitch: • Tetanus: Muscle twitch Muscle tetanus Effect of Muscle Length Force-Length Relationship • Isolated muscle • In vivo human muscles Force-Length Relationship • Isolated muscle Force-Length Relationship • In vivo human muscles – Two things are considered: muscle length and joint angle – In general, a group of muscles produces more force (torque) when muscles are lengthened before contraction. But some muscles do not follow this rule. Shoulder muscles Shoulder flexors (anterior deltoid): causes to flex shoulder joint 180° 135° Shoulder extensors(posterior deltoid): causes to extend 90° shoulder joint 45° 40° 0° Knee flexors • A person is lying on the stomach (prone position) 90° 120° 45° 0° Lower leg Thigh Knee joint Trunk Hip joint Knee flexors (hamstring muscles): causes to flex knee joint Hip flexors • A person is lying on the back (supine position) 90° 120° 45° 0° Lower leg Thigh Trunk Knee joint hip joint Hip flexors (iliopsoas, sartorius): causes to flex hip joint Knee extensors • A person is sitting on the bench 120° 90° 45° 0° Knee extensors (quadriceps muscles): causes to extend knee joint Elbow flexors Elbow flexors (biceps brachii): causes to flex elbow joint Force arm distance Effect of Velocity (Speed of Action) Force-velocity curve Effect of Muscle Fiber Effect of muscle fiber type on force Muscle Power Need to consider two factors: 1. Muscle force 2. Speed of action Power-Velocity Relationship Power = work/time = (force x distance)/time = force x speed Effect of muscle fiber type on power Factors that affect muscle force/power generation • • • • • • Rate of muscle stimulation Muscle length Joint angle Speed of action Muscle fiber type # of MU recruitment Motor Unit How does an individual generate appropriate force for a given task? Motor Unit (MU) Functional unit of movement Motor Unit (MU) •MU consists of •Single -motor neuron •Muscle fibers innervated by the -motor neuron Motor Unit (MU) •Fast fatigable MU (FF) •High twitch tension •High fatigue index •Fast fatigue resistant MU (FR) •Intermediate twitch tension •Intermediate fatigue index •Slow MU (S) •Low twitch tension •Low fatigue index Reasons for different twitch tension in different MU •Depends on number of muscle fibers and fiber size # muscle fiber: FF>FR> S Size of fiber: FF>FR>S Relationship b/n MU & Fiber type MU Fiber type FF FR S Fast glycolytic Fast oxidative Slow oxidative Motor Unit Muscle • Biceps brachii • Gastrocnemius • First lumbrical # neuron 774 580 98 # fibers/MU 750 1720 110 Motor Unit Recruitment • Follows the size principle – Small neuron cell body and axon activated first – Larger cell body and axon recruited later • Example: S MU FR MU FF MU 100% 0% % of effort level (Intensity of exercise) Gradation of Muscle Strength • By increasing # of MU recruited • By increasing frequency of stimulation Local Control of Muscle Action • Muscle spindle: muscle length monitor – Consists of 1) afferent nerves, 2) intrafusal fibers & 3) γ(gamma)-motor neurons • Golgi Tendon Organ: muscle tension monitor Structure of muscle spindle Action of muscle spindle Nerve impulse pattern of afferent nerves Rest Stretch Contraction Return to rest Speed of stretch on impulse discharge pattern Clinical implications for individuals with spastic muscle? Golgi Tendon Organ •GTO#<spindle # in given muscle •Composed of network of unmyelinated nerve fibers enclosed by fine capsule •Activated by either muscle stretch or muscle contraction. More sensitive to muscle contraction. •Activates inhibitory interneuron in spinal cord, which, in turn, inhibits -motor neuron of contracting muscle (agonists). Impulse discharge pattern of GTO during stretch and contraction Plasticity of Muscle • Metabolic and morphological changes to changes in stimulus – Increased stimulus: exercise training – Decreased stimulus: non-weight bearing, bed rest and aging Endurance training • Mode: jogging, running, cycling, swimming, etc • Adaptations – # of mitochondria – size of mitochondria – Oxidative enzyme activities • Krebs cycle, beta-oxidation, ETS – Some glycolytic enzymes – Capillary density Resistance training • Strength – Neural factor – Muscle fiber enlargement (hypertrophy) 33% 27% 38% 31% 5-6 month resistance training using triceps brachii MacDougall et al, EJAP 43:25-34 6 wk Resistance Training Limb suspension (Non-weight bearing) Limb unloading on muscle strength and X-area X-area Knee extensor Strength Dudley et al, in ACSM’s Resource Manual, p.201 Selective muscle atrophy with non-weight bearing Dudley et al, in ACSM’s Resource Manual, p.201 Bed Rest Muscle strength change with bed rest (soleus and gastroc.).) Dudley et al, in ACSM’s Resource Manual, p.203 Changes in skeletal muscles with aging • • • • # of muscle fibers Muscle area Fiber type distribution Muscle strength McArdle et al, in Exercise Physiology, p639 Muscle fiber distribution with aging Fiber Area Age %Type II Type I Type II 26.1 59.5 2944 3663 35.3 63.2 2854 3509 42.6 51.8 3133 3361 54.5 48.3 2877 2802 61.6 45.0 2264 2120 Muscle strength with aging • A decline in muscle strength is associated with a decrease in muscle mass. • A decline in lower extremity muscle strength is related to poor functional performance – walking ability, balance, stair-climbing ability, falls Trainability of skeletal muscles with aging Frontera et al, J. Appl. Physiol., 64:1038-1044, 1988 •Untrained old men (60-72 yrs) •8 reps/set, 3 sets/day, 3days/week at 80% of 1 RM for 12 weeks training •Thigh muscle X-area •Knee extension and flexion strength. Leg strength Frontera et al 1 RM max X-area of quadriceps Frontera et al Physician’s Role for Physical Activity Summary Know followings: names & functions •Muscle structure: connective tissues, pennation, myofibrils •Muscle fiber characteristics •Muscle action: E-C coupling •Sliding filament theory: changes during contraction •Muscle mechanics: force & power-length-velocity •MU: elements, function, characteristics. •Muscle action monitor: muscle spindle and GTO •Changes in muscle in training •Changes in muscle w/ suspension, bed rest aging The End