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  • 8/2/2019 Deadrick Chap 9 Handouts Part 1

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    Copyright 2010 PearsonEducation,Inc.

    Three Types of Muscle Tissue

    1. Skeletal muscle tissue: Attached to bones and skin Striated Voluntary (i.e., conscious control) Powerful

    Copyright 2010 PearsonEducation,Inc.

    Three Types of Muscle Tissue

    2. Cardiac muscle tissue: Only in the heart Striated Involuntary More details in Chapter 18

    Copyright 2010 PearsonEducation,Inc.

    Three Types of Muscle Tissue

    3. Smooth muscle tissue: In the walls of hollow organs, e.g.,

    stomach, urinary bladder, and airways

    Not striated Involuntary

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    Copyright 2010 PearsonEducation,Inc. Table 9.3

    Copyright 2010 PearsonEducation,Inc.

    Special Characteristics of Muscle Tissue

    1. Excitability: (responsiveness or irritability):ability to receive and respond to stimuli

    2. Contractility: ability to shorten whenstimulated

    3. Extensibility: ability to be stretched4. Elasticity: ability to recoil to resting length

    Copyright 2010 PearsonEducation,Inc.

    Muscle Functions

    1.Movement of bones or fluids (e.g., blood)2.Maintaining posture and body position3.Stabilizing joints4.Heat generation (especially skeletal muscle)

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    Copyright 2010 PearsonEducation,Inc.

    Skeletal Muscle Anatomy

    A muscle is an organ(In general) Each muscle is served by one

    artery, one nerve, and one or more veins

    Copyright 2010 PearsonEducation,Inc.

    Skeletal Muscle - Anatomy

    Connective tissue sheaths of skeletal muscle: Epimysium: dense regular connective tissue

    surrounding entire muscle

    Perimysium: fibrous connective tissue surroundingfascicles (groups of muscle fibers)

    Endomysium: fine areolar connective tissuesurrounding each muscle fiber

    ***muscle fiber = muscle cell

    Copyright 2010 PearsonEducation,Inc.

    Skeletal Muscle - Anatomy

    Muscle Structure Associated Connective Tissue

    Entire muscle Surrounded by epimysium

    Fascicles (a bundle of muscle cells) Surrounded by perimysium

    Muscle cell (muscle fiber) Surrounded by endomysium

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    Copyright 2010 PearsonEducation,Inc. Figure 9.1

    Bone

    Perimysium Muscle fiber

    Fascicle

    (wrapped by perimysium)

    Tendon

    Epimysium

    Muscle fiber

    in middle ofa fascicle

    Blood vessel

    Endomysium

    Fascicle(a)

    (b)

    Copyright 2010 PearsonEducation,Inc.

    Skeletal Muscle: Attachments

    Muscles attach:

    Directlyepimysium of muscle is fused to theperiosteum of bone orperichondrium of

    cartilage

    Indirectlyconnective tissue wrappingsextend beyond the muscle as a ropelike

    tendon or sheetlike aponeurosis

    Copyright 2010 PearsonEducation,Inc. Table 9.1

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    Copyright 2010 PearsonEducation,Inc.

    Microscopic Anatomy of a Skeletal Muscle

    Fiber

    Cylindrical cell 10 to 100 m in diameter, upto 30 cm long

    Multiple, peripheral nuclei (multinucleate)Many mitochondriaGlycosomes for glycogen storage,

    myoglobin for O2 storage

    Also contain myofibrils, sarcoplasmicreticulum, and T tubules

    Copyright 2010 PearsonEducation,Inc.

    Myofibrils

    Copyright 2010 PearsonEducation,Inc.

    NucleusLight I bandDark A band

    Sarcolemma

    Mitochondrion

    (b) Diagram ofpart of a muscle fibershowing the myofibrils. One

    myofibril is extended from the cut end of the fiber.

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    Copyright 2010 PearsonEducation,Inc.

    Sarcomere

    The organized, functional unit of a musclefiber

    Smallest contractile unit of a muscle fiberThe region of a myofibril between two

    successive Z discs

    Composed ofthick and thinmyofilaments(made of contractile proteins)

    Copyright 2010 PearsonEducation,Inc.

    Features of a Sarcomere

    Thick filaments: run the entire length of an A band Thin filaments: run the length of the I band and

    partway into the A band

    Z disc: coin-shaped sheet of proteins that anchorsthe thin filaments and connects myofibrils to oneanother

    H zone: lighter midregion where filaments do notoverlap

    M line: line of protein myomesin that holds adjacentthick filaments together

    Copyright 2010 PearsonEducation,Inc. Figure 9.2c, d

    I band I bandA bandSarcomere

    H zone

    Thin (actin)filament

    Thick (myosin)filament

    Z disc Z disc

    M line

    (c)Small part of one myofibril enlarged to show the myofilamentsresponsible for the banding pattern. Each sarcomereextends fromone Z disc to the next.

    Z disc Z discM line

    Sarcomere

    Thin (actin)filament

    Thick(myosin)filament

    Elastic (titin)filaments

    (d)Enlargement of one sarcomere (sectioned lengthwise). Notice themyosin heads on the thick filaments.

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    Copyright 2010 PearsonEducation,Inc.

    Ultrastructure of Thick Filament

    Composed of the protein myosin

    Myosin tails contain: 2 interwoven, heavy polypeptide chains

    Myosin heads contain: 2 smaller, light polypeptide chains that act ascross

    bridges during contraction

    Binding sites for actin of thin filaments Binding sites for ATP ATPase enzymes

    Copyright 2010 PearsonEducation,Inc.

    Ultrastructure of Thin Filament

    Twisted double strand of fibrous protein Factin

    F actin consists of G (globular) actin subunitsG actin bears active sites for myosin head

    attachment during contraction

    Tropomyosin and troponin: regulatoryproteins bound to actin

    Copyright 2010 PearsonEducation,Inc. Figure 9.3

    Flexible hinge region

    Tail

    Tropomyosin Troponin ActinMyosin head

    ATP-bindingsite

    Heads Active sitesfor myosinattachment

    Actinsubunits

    Actin-binding sites

    Thick filamentEach thick filament consists of manymyosin molecules whose heads protrude

    at opposite ends of the filament.

    Thin filamentA thin filament consists of two strandsof actin subunits twisted into a helix

    plus two types of regulatory proteins(troponin and tropomyosin).

    Thin filamentThick filament

    In the center of the sarcomere, the thickfilaments lack myosin heads. Myosin headsare present only in areas of myosin-actin overlap.

    Longitudinal section of filamentswithin one sarcomere of a myofibril

    Portion of a thick filamentPortion of a thin filament

    Myosin molecule Actin subunits

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    Copyright 2010 PearsonEducation,Inc.

    Sarcoplasmic Reticulum (SR)

    Network ofsmooth endoplasmic reticulumsurrounding each myofibril

    Pairs ofterminal cisternae formperpendicular cross channels

    Functions in the regulation of intracellularCa2+levels

    Copyright 2010 PearsonEducation,Inc.

    T Tubules

    Continuous with the sarcolemmaPenetrate the cells interiorat each A band

    I band junction

    Associate with the paired terminal cisternaeto form triads that encircle each sarcomere

    Carry the signal for contraction deep intothe sarcoplasm

    Copyright 2010 PearsonEducation,Inc. Figure 9.5

    Myofibril

    Myofibrils

    Triad:

    Tubules of

    the SR

    Sarcolemma

    Sarcolemma

    Mitochondria

    I band I bandA band

    H zone Z discZ disc

    Part of a skeletal

    muscle fiber (cell)

    T tubule Terminal

    cisternaeof the SR (2)

    M line

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    Copyright 2010 PearsonEducation,Inc.

    Triad Relationships

    T tubules conduct impulses deep intomuscle fiber

    Integral proteins protrude into theintermembrane space from T tubule and SR

    cisternae membranes

    T tubule proteins: voltage sensorsSR foot proteins: gated channels that

    regulate Ca2+ release from the SR cisternae

    Copyright 2010 PearsonEducation,Inc.

    Contraction

    The generation of forceDoes not necessarily cause shortening of the

    fiber

    Shortening occurs when tension generated bycross bridges on the thin filaments exceeds

    forces opposing shortening

    Copyright 2010 PearsonEducation,Inc.

    Sliding Filament Model of Contraction

    In the relaxed state, thin and thick filamentsoverlap only slightly

    During contraction, myosin heads bind toactin, detach, and bind again, to propel the

    thin filaments toward the M line

    As H zones shorten and disappear,sarcomeres shorten, muscle cells shorten,

    and the whole muscle shortens

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    Copyright 2010 PearsonEducation,Inc. Figure 9.6

    IFully relaxed sarcomere of a muscle fiber

    Fully contracted sarcomere of a muscle fiber

    IAZ ZH

    I IAZ Z

    1

    2

    Copyright 2010 PearsonEducation,Inc.

    Requirements for Skeletal Muscle

    Contraction

    1. Activation: neural stimulation at aneuromuscular junction

    2. Excitation-contraction coupling: Generation and propagation of an action

    potential along the sarcolemma

    Final trigger: a brief rise in intracellular Ca2+levels

    Copyright 2010 PearsonEducation,Inc.

    Events at the Neuromuscular Junction

    Skeletal muscles are stimulated by somaticmotor neurons (somatic efferents)

    Axons of motor neurons travel from thecentral nervous system via nerves to skeletal

    musclesEach axon forms several branches as it

    enters a muscle

    Each axon ending forms a neuromuscularjunction with a single muscle fiber

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    Copyright 2010 PearsonEducation,Inc.

    Neuromuscular Junction

    Situated midway along the length of a musclefiber

    Axon terminal and muscle fiber are separatedby a gel-filled space called the synaptic cleft

    Synaptic vesicles of axon terminal contain theneurotransmitteracetylcholine (ACh)

    Junctional folds of the sarcolemma containACh receptors

    Copyright 2010 PearsonEducation,Inc.

    Events at the Neuromuscular Junction

    Nerve impulse arrives at axon terminalACh is released and binds with receptors on

    the sarcolemma

    Electrical events lead to the generation of anaction potential

    Copyright 2010 PearsonEducation,Inc. Figure 9.8

    Nucleus

    Action

    potential (AP)Myelinated axon

    of motor neuronAxon terminal of

    neuromuscularjunction

    Sarcolemma ofthe muscle fiber

    Ca2+ Ca2+

    Axon terminalof motorneuron

    Synaptic vesicle

    containing AChMitochondrion

    Synapticcleft

    Junctional

    folds ofsarcolemma

    Fusing synaptic vesicles

    ACh

    Sarcoplasm of

    muscle fiberPostsynaptic membrane

    ion channel opens;ions pass.

    Na+ K+

    AchNa+

    K+

    DegradedACh

    Acetyl-cholinesterase

    Postsynaptic membraneion channel closed;ions cannot pass.

    1 Action potential arrives ataxon terminal of motor neuron.

    2 Voltage-gated Ca2+ channelsopen and Ca2+ enters the axonterminal.

    3Ca2+ entry causes somesynaptic vesicles to release

    their contents (acetylcholine)by exocytosis.4 Acetylcholine, aneurotransmitter, diffuses acrossthe synaptic cleft and binds toreceptors in the sarcolemma.

    5 ACh binding opens ionchannels that allow simultaneous

    passage of Na+ into the musclefiber and K+ out of the muscle

    fiber.

    6 ACh effects are terminatedby its enzymatic breakdown inthe synaptic cleft byacetylcholinesterase.

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    Copyright 2010 PearsonEducation,Inc.

    Destruction of Acetylcholine

    ACh effects are quickly terminated by theenzyme acetylcholinesterase

    Prevents continued muscle fiber contraction inthe absence of additional stimulation

    Copyright 2010 PearsonEducation,Inc.

    Events in Generation of an Action Potential

    1. Local depolarization (end plate potential): ACh binding opens chemically (ligand)

    gated ion channels

    Simultaneous diffusion of Na+ (inward) andK+ (outward)

    More Na+ diffuses, so the interior of thesarcolemma becomes less negative

    Local depolarization = end plate potential

    Copyright 2010 PearsonEducation,Inc.

    Events in Generation of an Action Potential

    2. Generation and propagation of an actionpotential:

    End plate potential spreads to adjacentmembrane areas

    Voltage-gated Na+ channels open Na+ influx decreases the membrane voltage

    toward a critical threshold

    If threshold is reached, an actionpotential is generated

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    Copyright 2010 PearsonEducation,Inc. Figure 9.9, step 1

    Na+

    Na+

    Open Na+

    Channel

    Closed K+

    Channel

    K+

    Na+ K+Action potential++++

    ++++++++

    Axon terminal

    Synaptic

    cleft

    ACh

    ACh

    Sarcoplasm of muscle fiber

    K+

    1 Local depolarization: generation of theend plate potential on the sarcolemma1

    Waveofdepolarization

    Copyright 2010 PearsonEducation,Inc. Figure 9.9, step 2

    Na+

    Na+

    Open Na+

    Channel

    Closed K+

    Channel

    K+

    Na+ K+Action potential++++

    ++++++++

    Axon terminal

    Synapticcleft

    ACh

    ACh

    Sarcoplasm of muscle fiber

    K+

    Generation and propagation of theaction potential (AP)

    1 Local depolarization: generation of theend plate potential on the sarcolemma

    2

    1

    Waveofdepolarization

    Copyright 2010 PearsonEducation,Inc. Figure 9.9, step 3

    Na+

    Closed Na+

    Channel

    Open K+

    Channel

    K+

    Repolarization3

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    Copyright 2010 PearsonEducation,Inc. Figure 9.9

    Na+

    Na+

    Open Na+

    Channel

    Closed K+

    Channel

    Action potential++++++

    ++++++

    Axon terminal

    Synaptic

    cleft

    ACh

    ACh

    Sarcoplasm of muscle fiber

    K+

    2Generation and propagation ofthe action potential (AP)

    3 Repolarization

    1 Local depolarization:generation of the endplate potential on thesarcolemma

    K+

    K+Na+

    K+Na+

    Wave

    of

    dep

    ola

    riza

    tion

    Closed Na+

    Channel

    Open K+

    Channel

    Copyright 2010 PearsonEducation,Inc. Figure 9.10

    Na+ channels

    close, K+ channels

    open

    K+ channels

    close

    Repolarization

    due to K+ exit

    Threshold

    Na+

    channels

    open

    Depolarization

    due to Na+ entry

    Copyright 2010 PearsonEducation,Inc.

    Excitation-Contraction (E-C) Coupling

    Sequence of events by which transmission ofan AP along the sarcolemma leads to sliding

    of the myofilaments

    Latent period: Time when E-C coupling events occur Time between AP initiation and the

    beginning of contraction

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    Copyright 2010 PearsonEducation,Inc.

    Events of Excitation-Contraction (E-C)

    Coupling

    AP is propagated along sarcomere to Ttubules

    Voltage-sensitive proteins stimulate Ca2+release from SR

    Ca2+ is NECESSARY for contraction

    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 1

    Axon terminal

    of motor neuron

    Muscle fiberTriad

    One sarcomere

    Synaptic cleft

    Setting the stage

    Sarcolemma

    Action potential

    is generated

    Terminal cisterna of SR

    ACh

    Ca2+

    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 2

    Action potentialis propagatedalongthe sarcolemma anddown the Ttubules.

    Steps in E-C Coupling:

    Troponin Tropomyosinblockingactivesites

    Myosin

    Actin

    Activesitesexposedandreadyformyosin binding

    Ca2+

    TerminalcisternaofSR

    Voltage-sensitivetubuleprotein Ttubule

    Ca2+releasechannel

    Myosincrossbridge

    Ca2+

    Sarcolemma

    Calcium ionsare released.

    Calcium bindsto troponin andremovesthe blocking action oftropomyosin.

    Contraction begins

    The aftermath

    1

    2

    3

    4

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    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 3

    Steps inE-C Coupling:

    Terminalcisternaof SR

    Voltage-sensitivetubule protein

    T tubule

    Ca2+releasechannel

    Ca2+

    Sarcolemma

    Action potential ispropagated along thesarcolemma and down

    the T tubules.

    1

    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 4

    Steps inE-C Coupling:

    Terminalcisternaof SR

    Voltage-sensitivetubule protein

    T tubule

    Ca2+releasechannel

    Ca2+

    Sarcolemma

    Action potential ispropagated along thesarcolemma and downthe T tubules.

    Calciumions arereleased.

    1

    2

    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 5

    Troponin Tropomyosinblocking active sitesMyosin

    Actin

    Ca2+

    The aftermath

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    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 6

    Troponin Tropomyosin

    blocking active sitesMyosin

    Actin

    Active sites exposed andready for myosin binding

    Ca2+

    Calcium binds totroponin and removesthe blocking action oftropomyosin.

    The aftermath

    3

    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 7

    Troponin Tropomyosinblocking active sitesMyosin

    Actin

    Active sites exposed andready for myosin binding

    Ca2+

    Myosincrossbridge

    Calcium binds totroponin and removesthe blocking action oftropomyosin.

    Contraction begins

    The aftermath

    3

    4

    Copyright 2010 PearsonEducation,Inc. Figure 9.11, step 8

    Action potentialis propagatedalongthe sarcolemma anddown the Ttubules.

    Steps in E-C Coupling:

    Troponin Tropomyosinblockingactivesites

    Myosin

    Actin

    Activesitesexposedandreadyformyosin binding

    Ca2+

    TerminalcisternaofSR

    Voltage-sensitivetubuleprotein Ttubule

    Ca2+releasechannel

    Myosincrossbridge

    Ca2+

    Sarcolemma

    Calcium ionsare released.

    Calcium bindsto troponin andremovesthe blocking action oftropomyosin.

    Contraction begins

    The aftermath

    1

    2

    3

    4

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    Copyright 2010 PearsonEducation,Inc.

    Role of Calcium (Ca2+) in Contraction

    At low intracellular Ca2+ concentration: Tropomyosin blocks the active sites on actin Myosin heads cannot attach to actin Muscle fiber relaxes

    Copyright 2010 PearsonEducation,Inc.

    Role of Calcium (Ca2+) in Contraction

    At higher intracellular Ca2+ concentrations: Ca2+ binds to troponin Troponin changes shape and moves

    tropomyosin away from active sites

    Events of the cross bridge cycle occur When nervous stimulation ceases, Ca2+ is

    pumped back into the SR and contraction

    ends

    Copyright 2010 PearsonEducation,Inc.

    Cross Bridge Cycle

    Continues as long as the Ca2+ signal andadequate ATP are present

    Cross bridge formationhigh-energymyosin head attaches to thin filament

    Working (power) strokemyosin headpivots and pulls thin filament toward M line

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    Copyright 2010 PearsonEducation,Inc.

    Cross Bridge Cycle

    Cross bridge detachmentATP attaches tomyosin head and the cross bridge detaches

    Cocking of the myosin headenergy fromhydrolysis of ATP cocks the myosin head into

    the high-energy state

    Copyright 2010 PearsonEducation,Inc. Figure 9.12

    1

    Actin

    Cross bridge formation.

    Cocking of myosin head. The power (working)stroke.

    Cross bridge

    detachment.

    Ca2+

    Myosin

    cross bridge

    Thick

    filament

    Thin filament

    ADP

    Myosin

    Pi

    ATP

    hydrolysis

    ATP

    ATP

    24

    3

    ADP

    PiADP

    Pi

    Copyright 2010 PearsonEducation,Inc. Figure 9.12, step 1

    Actin

    Cross bridge formation.

    Ca2+

    Myosin

    cross bridgeThick filament

    Thin filament

    ADP

    Myosin

    Pi

    1

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    Copyright 2010 PearsonEducation,Inc. Figure 9.12, step 3

    The power (working) stroke.

    ADP

    Pi

    2

    Copyright 2010 PearsonEducation,Inc. Figure 9.12, step 4

    Cross bridge detachment.

    ATP

    3

    Copyright 2010 PearsonEducation,Inc. Figure 9.12, step 5

    Cocking of myosin head.

    ATP

    hydrolysis

    ADP

    Pi

    4

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    Copyright 2010 PearsonEducation,Inc. Figure 9.12

    1

    Actin

    Cross bridge formation.

    Cocking of myosin head. The power (working)stroke.

    Cross bridge

    detachment.

    Ca2+

    Myosin

    cross bridgeThick

    filament

    Thin filament

    ADP

    Myosin

    Pi

    ATP

    hydrolysis

    ATP

    ATP

    24

    3

    ADP

    PiADPPi