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Copyright © 2010 Pearson Education, Inc. Muscle Functions 1. Movement of bones or fluids (e.g., blood) 2. Maintaining posture and body position 3. Stabilizing joints 4. Heat generation (esp. skeletal muscle) Copyright © 2010 Pearson Education, Inc. Skeletal Muscle • Connective tissue sheaths of skeletal muscle: • Epimysium: fibrous CT surrounding entire muscle • Perimysium: fibrous CT surrounding fascicles (groups of muscle fibers) • Endomysium: delicate CT surrounding each muscle fiber Copyright © 2010 Pearson Education, Inc. Epimysium Epimysium Tendon Fascicle Perimysium Endomysium Muscle fiber in middle of a fascicle (wrapped by perimysium) Endomysium Perimysium Fascicle Copyright © 2010 Pearson Education, Inc. Muscle fiber Skeletal Muscle: Attachments • Muscles attach to bone by an origin and insertion • Origin —is fixed and on the immovable bone • Insertion—is on the movable bone. • As a contraction occurs the insertion moves towards the origin Copyright © 2010 Pearson Education, Inc. Microscopic Anatomy of a Skeletal Muscle Fiber • Multiple peripheral nuclei, many mitochondria • Also contain sarcolemma, myofibrils, sarcoplasmic reticulum, T tubules Sarcolemma Mitochondrion Myofibril Dark A band Light I band Nucleus Copyright © 2010 Pearson Education, Inc. I band A band I band Z disc H zone Z disc Myofibril M line Sarcolemma Triad: tubule •T • Terminal cisternae of the SR (2) Tubules of the SR Myofibrils Mitochondria Copyright © 2010 Pearson Education, Inc. Myofibrils • Densely packed, rodlike organelles • ~80% of cell volume • Composed of sarcomeres • Exhibit striations:perfectly aligned repeating series of dark A bands and light I bands Myofibril Copyright © 2010 Pearson Education, Inc. Sarcomere • Smallest contractile unit (functional unit) of a muscle fiber • The region of a myofibril between two successive Z discs • Composed of myofilaments: Thick (myosin) and thin (actin) Copyright © 2010 Pearson Education, Inc. Regions of a Sarcomere • A band (Dark Band)- Contains thin & thick filaments • H zone: lighter midregion where filaments do not overlap • M line: proteins that hold adjacent thick filaments together; center of sarcomere • I band (Light Band)- Contains only thin filaments • Z disc: proteins that anchor thin filaments; mark start and end of one sarcomere Copyright © 2010 Pearson Education, Inc. Thin (actin) filament Thick (myosin) filament (c)Part Z disc H zone I band A band Sarcomere Z disc I band M line of one myofibril Sarcomere Z disc M line Z disc Thin (actin) filament Elastic (titin) filaments Thick (myosin) filament (d) Copyright © 2010 Pearson Education, Inc. Myofilaments of a Sarcomere • Thick Filament • Thin Filament • Composed of many myosin proteins • Twisted double strand of actin protein • A single myosin protein has a “tail” and a “head” that can: • Has active sites for the myosin head • Bind actin and pull it during a contraction • Hydrolyze ATP to release energy Copyright © 2010 Pearson Education, Inc. Longitudinal section of filaments within one sarcomere of a myofibril Thick filament Thin filament Thick filament Thin filament Portion of a thick filament Myosin head Portion of a thin filament Tropomyosin Troponin Actin Actin-binding sites ATPbinding site Heads Tail Flexible hinge region Copyright © 2010 Pearson Education, Inc. Myosin molecule Active sites for myosin attachment Actin subunits Actin subunits •In the relaxed state, thin and thick filaments slightly overlap •During contraction, myosin heads bind to actin, detach, and bind again, propelling thin filaments toward M line •As H zones shorten and disappear, sarcomeres shorten, muscle cells shorten, and whole muscle shortens Z I 1 Copyright © 2010 Pearson Education, Inc. I Fully relaxed Z 2 Z H A I Z A Fully contracted I The Neuromuscular Junction • Defined as: • Axons of motor neurons: travel from the brain/spinal cord via nerves to skeletal muscles • Each axon: branches into a number of axon terminals as it enters a muscle • Each axon ending forms: a neuromuscular junction with a single muscle fiber Copyright © 2010 Pearson Education, Inc. 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 Fusing synaptic vesicles Copyright © 2010 Pearson Education, Inc. Synaptic vesicle containing ACh Mitochondrion Synaptic cleft The Neuromuscular Junction • Axon terminal and muscle fiber are: separated by space called the synaptic cleft • Synaptic vesicles within axon terminal contain: the neurotransmitter acetylcholine (ACh) • Junctional folds of the sarcolemma contain: ACh receptors (chemically-gated channels) Copyright © 2010 Pearson Education, Inc. Events at the Neuromuscular Junction 1) A nerve impulse: arrives at the axon terminal 2) Ca2+ floods into axon terminal 3) Ca2+ entry causes synaptic vesicles to release Ach 4) ACh diffuses across the synaptic cleft and binds to receptors on the sarcolemma 5) Ach binding opens channels 6) Na+ floods into muscle fiber and K+ floods out making the interior of cell less negative 7) Once threshold is reached an AP is generated Copyright © 2010 Pearson Education, Inc. The Action Potential • The AP is an unstoppable, electrical event that travels along the entire sarcolemma conducting the electrical impulse from one end of cell to the other • Repolarization :The muscle cell returns to its resting state mainly by the exit of K+ Copyright © 2010 Pearson Education, Inc. 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 open and Ca2+ enters the axon terminal. Ca2+ Ca2+ Axon terminal of motor neuron 3 Ca2+ entry causes some Fusing synaptic vesicles synaptic vesicles to release their contents (acetylcholine) by exocytosis. ACh 4 Acetylcholine, a neurotransmitter, diffuses across the synaptic cleft and binds to receptors in the sarcolemma. Na+ K+ channels that allow simultaneous passage of Na+ into the muscle fiber and K+ out of the muscle fiber. by its enzymatic breakdown in the synaptic cleft by acetylcholinesterase. Copyright © 2010 Pearson Education, Inc. Junctional folds of sarcolemma Sarcoplasm of muscle fiber 5 ACh binding opens ion 6 ACh effects are terminated Synaptic vesicle containing ACh Mitochondrion Synaptic cleft Ach– Degraded ACh Na+ Acetylcholinesterase Postsynaptic membrane ion channel opens; ions pass. Postsynaptic membrane ion channel closed; ions cannot pass. K+ Figure 9.8 Destruction of Acetylcholine • ACh effects are quickly terminated by the enzyme acetylcholinesterase • Prevents continued muscle fiber contraction in the absence of additional stimulation Copyright © 2010 Pearson Education, Inc. 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) 1 Local depolarization: Sarcoplasm of muscle fiber Copyright © 2010 Pearson Education, Inc. Closed Na+ Open K+ Channel Channel Na+ K+ 3 Repolarization Setting the stage Axon terminal of motor neuron Action potential Synaptic cleft is generated ACh Sarcolemma Terminal cisterna of SR Muscle fiber Ca2+ Triad One sarcomere Copyright © 2010 Pearson Education, Inc. 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 Ca2+ Copyright © 2010 Pearson Education, Inc. 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 Ca2+ Copyright © 2010 Pearson Education, Inc. 2 Calcium ions are released. Role of Calcium (Ca2+) in Contraction • At low intracellular Ca2+ concentration: • Active sites on actin are blocked • Myosin heads cannot attach to actin • Muscle fiber relaxes Copyright © 2010 Pearson Education, Inc. 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 Copyright © 2010 Pearson Education, Inc. Role of Calcium (Ca2+) in Contraction • At higher intracellular Ca2+ concentrations: • Ca2+ causes binding sites on actin to be exposed • Events of the cross bridge cycle occur • When nervous stimulation ceases, Ca2+ is pumped back into the SR and contraction ends Copyright © 2010 Pearson Education, Inc. 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 Copyright © 2010 Pearson Education, Inc. Cross Bridge Cycle • Continues as long as the Ca2+ signal and adequate ATP are present • Cross bridge formation: high-energy myosin head attaches to thin filament • Power stroke: myosin head pivots and pulls thin filament toward M line Copyright © 2010 Pearson Education, Inc. Cross Bridge Cycle • Cross bridge detachment: ATP attaches to myosin head and the cross bridge detaches • “Cocking” of the myosin head: energy from hydrolysis of ATP cocks the myosin head into the high-energy state Copyright © 2010 Pearson Education, Inc. 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. Copyright © 2010 Pearson Education, Inc. Spinal cord Motor Motor unit 1 unit 2 Axon terminals at neuromuscular junctions Nerve Motor neuron cell body Motor Muscle neuron axon Muscle fibers • Motor unit = a motor neuron and all (four to several hundred) muscle fibers it supplies Copyright © 2010 Pearson Education, Inc. Graded Muscle Responses • Defined: Variations in the degree of muscle contraction Responses are graded by: 1. Changing the frequency of stimulation 2. Changing the number of muscle cells being stimulated at one time (by changing strength of stimulus) Copyright © 2010 Pearson Education, Inc. Response to Change in Stimulus Frequency Single stimulus single twitch Contraction Relaxation Stimulus A single stimulus results in a single contractile response called a muscle twitch Copyright © 2010 Pearson Education, Inc. Figure 9.15a Response to Change in Stimulus Frequency • Increase frequency of stimulus muscle doesn’t have time to completely relax (btwn. stimuli) • Ca2+ release stimulates further contraction temporal (wave) summation • Further increase in stimulus frequency unfused (incomplete) tetanus • If stimuli are given quickly enough, fused (complete) tetanus results Copyright © 2010 Pearson Education, Inc. Low stimulation frequency unfused (incomplete) tetanus Partial relaxation Stimuli (b) If another stimulus is applied before the muscle relaxes completely, then more tension results. Copyright © 2010 Pearson Education, Inc. Figure 9.15b High stimulation frequency fused (complete) tetanus Stimuli (c) At higher stimulus frequencies, there is no relaxation at all between stimuli. This is fused (complete) tetanus. Copyright © 2010 Pearson Education, Inc. Figure 9.15c Muscle Metabolism: Energy for Contraction • ATP is the only source used directly for contractile activities • Available stores of ATP are depleted in 4–6 seconds Copyright © 2010 Pearson Education, Inc. Muscle Metabolism: Energy for Contraction • ATP is regenerated by: • Direct phosphorylation of ADP by creatine phosphate (CP) • Anaerobic pathway • Aerobic pathway Copyright © 2010 Pearson Education, Inc. Muscle Metabolism: Energy for Contraction • Direct phosphorylation of ADP by creatine phosphate (CP) • CP is more concentrated in muscle fibers than ATP (~4 X more) • When ATP stores are depleted: muscle fibers use CP to regenerate ATP • Products are: 1 ATP/ CP • Provides energy for: ~ 15 seconds of activity Copyright © 2010 Pearson Education, Inc. (a) Direct phosphorylation Coupled reaction of creatine phosphate (CP) and ADP Energy source: CP CP ADP Creatine kinase Creatine ATP Oxygen use: None Products: 1 ATP per CP, creatine Duration of energy provision: 15 seconds Copyright © 2010 Pearson Education, Inc. Figure 9.19a Anaerobic Pathway • Under intense muscle activity or when oxygen delivery is impaired: the body switches to the anaerobic pathway • Begins just like aerobic pathway (Glucose breakdown) but pyruvic acid is converted into lactic acid • Products are: 2ATP/glucose • Provides energy for : 60 seconds of activity Copyright © 2010 Pearson Education, Inc. (b) Anaerobic pathway Glycolysis and lactic acid formation Energy source: glucose Glucose (from glycogen breakdown or delivered from blood) Glycolysis in cytosol 2 O2 ATP Pyruvic acid net gain O2 Released to blood Lactic acid Oxygen use: None Products: 2 ATP per glucose, lactic acid Duration of energy provision: 60 seconds, or slightly more Copyright © 2010 Pearson Education, Inc. Figure 9.19b Aerobic Pathway • Produces 95% of ATP during rest and light to moderate exercise • Fuels: stored glycogen, then bloodborne glucose, pyruvic acid from glycolysis, and free fatty acids • Products are: 32 ATP/glucose, CO2 and H2O • Provides energy for: hours (endurance activities) Copyright © 2010 Pearson Education, Inc. (c) Aerobic pathway Aerobic cellular respiration Energy source: glucose; pyruvic acid; free fatty acids from adipose tissue; amino acids from protein catabolism Glucose (from glycogen breakdown or delivered from blood) O2 Pyruvic acid Fatty acids O2 Aerobic respiration Aerobic respiration in mitochondria mitochondria Amino acids 32 CO2 H2O ATP net gain per glucose Oxygen use: Required Products: 32 ATP per glucose, CO2, H2O Duration of energy provision: Hours Copyright © 2010 Pearson Education, Inc. Figure 9.19c Short-duration exercise ATP stored in muscles is used first. ATP is formed from creatine Phosphate and ADP. Copyright © 2010 Pearson Education, Inc. 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. MUSCLE IDENTIFICATION AND NAMING Copyright © 2010 Pearson Education, Inc. Naming Skeletal Muscles • Location—bone or body region associated with the muscle • Shape—e.g., deltoid muscle (deltoid = triangle) • Relative size—e.g., maximus (largest), minimus (smallest), longus (long) • Direction of fibers or fascicles—e.g., rectus (fibers run straight), transversus, and oblique (fibers run at angles to an imaginary defined axis) Copyright © 2010 Pearson Education, Inc. Naming Skeletal Muscles • Number of origins—e.g., biceps (2 origins) and triceps (3 origins) • Location of attachments—named according to point of origin or insertion • Action—e.g., flexor or extensor, muscles that flex or extend, respectively Copyright © 2010 Pearson Education, Inc. Head Temporalis Masseter Shoulder Trapezius Deltoid Arm Triceps brachii Biceps brachii Brachialis Forearm Hand, wrist and finger flexors Pelvis/thigh Iliopsoas Pectineus Thigh Rectus femoris Vastus lateralis Vastus medialis Leg Fibularis longus Extensor digitorum longus Tibialis anterior Copyright © 2010 Pearson Education, Inc. Facial Epicranius, frontal belly Orbicularis oculi Zygomaticus Orbicularis oris Neck Platysma Sternocleidomastoid Thorax Pectoralis major Abdomen Rectus abdominis Internal oblique Transversus abdominis External oblique Thigh Sartorius Adductors Gracilis Leg Gastrocnemius Soleus Muscles of Facial Expression • Epicranius (Frontal belly and Occipital belly) • Raises eyebrows, wrinkles forehead (frontal belly) • Pulls scalp posteriorly (occipital belly) • Orbicularis Oculi • Closes eyes, squinting, blinking • Orbicularis Oris • Closes mouth, protrudes lips • Buccinator • Flattens cheek (as in whistling) • Zygomaticus • Pulls corners of mouth superiorly (as in smiling) • Platysma • Pulls corners of mouth inferiorly Copyright © 2010 Pearson Education, Inc. Orbicularis oculi Zygomaticus Buccinator Orbicularis oris Platysma Copyright © 2010 Pearson Education, Inc. Temporalis Masseter Sternocleidomastoid Trapezius Muscles of Mastication • Temporalis and Masseter • Elevate the mandible (closing jaw) Copyright © 2010 Pearson Education, Inc. Temporalis Orbicularis oris Buccinator (a) Copyright © 2010 Pearson Education, Inc. Masseter Muscles of the Neck • Sternocleidomastoid—major head flexor (also rotates the head) Copyright © 2010 Pearson Education, Inc. 1st cervical vertebra Sternocleidomastoid (a) Anterior Copyright © 2010 Pearson Education, Inc. Base of occipital bone Mastoid process AnteriorTrunk Muscles • Pectoralis Major • Adducts and flexes the arm • Rectus Abdominis • Flexes vertebral column • External and Internal obliques • Flex vertebral column • Transversus Abdominis • Compresses abdomen Copyright © 2010 Pearson Education, Inc. Subscapularis Pectoralis minor Sternum (a) Copyright © 2010 Pearson Education, Inc. Serratus anterior Pectoralis major Linea alba Tendinous intersection External oblique (a) Copyright © 2010 Pearson Education, Inc. Rectus abdominis Posterior Trunk Muscles • Trapezius • Extends head; elevates, depresses scapula • Latissimus Dorsi • Adducts and extends humerus • Erector Spinae • Extends vertebral column Copyright © 2010 Pearson Education, Inc. Levator scapulae Trapezius Rhomboid minor Rhomboid major (c) Copyright © 2010 Pearson Education, Inc. Arm Muscles • Anterior Flexor Muscles • Brachialis, Biceps brachii, Brachioradialis • Forearm flexors • Posterior extensor muscles • Triceps brachii • Extend forearm Copyright © 2010 Pearson Education, Inc. Biceps brachii Brachialis Brachioradialis (a) Anterior view Copyright © 2010 Pearson Education, Inc. Shoulder Muscles • Deltoid • Abducts arm, but can do all angular movements • Rotator Cuff Muscles • Supraspinatus • Infraspinatus • Subscapularis • Teres Minor Copyright © 2010 Pearson Education, Inc. Supraspinatus Infraspinatus Teres minor Triceps brachii: Lateral head Long head (b) Posterior view Copyright © 2010 Pearson Education, Inc. Muscles of the Forearm • Actions: movements of the wrist, hand, and fingers • Most anterior muscles are flexors and insert via the flexor retinaculum • Most posterior muscles are extensors and insert via the extensor retinaculum Copyright © 2010 Pearson Education, Inc. Flexor retinaculum Brachioradialis Medial head of triceps brachii Copyright © 2010 Pearson Education, Inc. (a) Hip/Thigh Muscles • Iliopsoas and Sartorius • Hip/thigh flexion Copyright © 2010 Pearson Education, Inc. Iliopsoas Psoas major Iliacus Sartorius (a) Copyright © 2010 Pearson Education, Inc. 5th lumbar vertebra Hip/Thigh Muscles • Gluteus Maximus • Lateral thigh • Extends leg at hip • Gluteus Medius • Abducts thigh • Adductor Muscles • Medial thigh • Adductor thigh Copyright © 2010 Pearson Education, Inc. Gluteus medius (cut) Gluteus minimus Gluteus maximus (cut) (c) Copyright © 2010 Pearson Education, Inc. Pectineus (cut) Adductor brevis Adductor longus Adductor magnus Femur (b) Copyright © 2010 Pearson Education, Inc. O = origin I = insertion Thigh Muscles • Hamstring muscles (Biceps femoris, Semitendinosus, Semimembranosus ) • Posterior thigh • All three flex leg at knee and extend hip Copyright © 2010 Pearson Education, Inc. Long head Biceps Short head femoris Semitendinosus Semimembranosus (a) Copyright © 2010 Pearson Education, Inc. Hamstrings Muscles of the Thigh that Move the Knee Joint • Quadriceps femoris (Vastus Medialis, Lateralis, intermedius and rectus femoris) • Anterior Thigh • All extend the knee Copyright © 2010 Pearson Education, Inc. Quadriceps femoris • Rectus femoris (superficial to vastus intermedius) • Vastus lateralis • Vastus medialis Tendon of quadriceps femoris Patella (a) Copyright © 2010 Pearson Education, Inc. Arm Triceps brachii Brachialis Forearm Brachioradialis Extensors of forearm Extensor carpi ulnaris Extensor digitorum Iliotibial tract Leg Gastrocnemius Soleus Fibularis longus Calcaneal (Achilles) tendon Copyright © 2010 Pearson Education, Inc. Neck Epicranius, occipital belly Sternocleidomastoid Trapezius Shoulder Deltoid Infraspinatus Teres major Rhomboid major Latissimus dorsi Hip Gluteus medius Gluteus maximus Thigh Adductor magnus Hamstrings: Biceps femoris Semitendinosus Semimembranosus Figure 10.5 Gluteus medius Gluteus maximus Adductor magnus Gracilis Iliotibial tract Biceps femoris Semitendinosus Semimembranosus (a) Copyright © 2010 Pearson Education, Inc. Hamstrings Muscles of the Anterior Compartment of the Leg • Tibialis anterior & Extensor digitorum longus • Primary toe extensors and ankle dorsiflexors Copyright © 2010 Pearson Education, Inc. Tibialis anterior Extensor digitorum longus (a) Copyright © 2010 Pearson Education, Inc. Muscles of the Lateral Compartment of the Leg • Fibularis longus • Plantar flexion and eversion of the foot Copyright © 2010 Pearson Education, Inc. Head of fibula Fibularis longus Lateral malleolus (a) Copyright © 2010 Pearson Education, Inc. Muscles of the Posterior Compartment of the Leg • Gastrocnemius and Soleus • Both do plantar flexion of foot • Gastrocnemius also does flexion at knee Copyright © 2010 Pearson Education, Inc. Gastrocnemius Medial head Lateral head Tendon of gastrocnemius Calcaneal tendon Medial malleolus Lateral malleolus Calcaneus (a) Superficial view of the posterior leg. Copyright © 2010 Pearson Education, Inc.