deadrick chap 9 handouts part 1
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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
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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
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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
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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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Skeletal Muscle Anatomy
A muscle is an organ(In general) Each muscle is served by one
artery, one nerve, and one or more veins
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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
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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)
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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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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
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Myofibrils
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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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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)
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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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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
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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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Sarcoplasmic Reticulum (SR)
Network ofsmooth endoplasmic reticulumsurrounding each myofibril
Pairs ofterminal cisternae formperpendicular cross channels
Functions in the regulation of intracellularCa2+levels
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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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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
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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
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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
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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
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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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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
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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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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
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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
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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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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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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
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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
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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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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