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Muscles and Muscle Tissue

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1 Muscles and Muscle Tissue

2 Muscle Overview The three types of muscle tissue are skeletal, cardiac, and smooth Skeletal and smooth muscle cells are elongated and are called muscle fibers Muscle contraction depends on two kinds of myofilaments – actin and myosin

3 Muscle Similarities Muscle terminology is similar
Sarcolemma – muscle plasma membrane Sarcoplasm – cytoplasm of a muscle cell Prefixes – myo, mys, and sarco all refer to muscle

4 Skeletal Muscle Tissue
Packaged in skeletal muscles that attach to and cover the bony skeleton It is striated Is controlled voluntarily Contracts rapidly but tires easily Is responsible for all locomotion, posture, joint stabilization and heat generation

5 Cardiac Muscle Tissue Occurs only in the heart
Is striated like skeletal muscle but involuntary Contracts at a fairly steady rate set by the heart’s pacemaker

6 Smooth Muscle Tissue Found in the walls of hollow visceral organs
Propels food and other substances through organs It is not striated and involuntary

7 Functional Characteristics of Muscle Tissue
Excitability, or irritability – the ability to receive and respond to stimuli Contractility – the ability to shorten forcibly Extensibility – the ability to be stretched or extended Elasticity – the ability to recoil and resume the original resting length

8 Structure and Organization of Skeletal Muscle
Table 9.1a

9 Structure and Organization of Skeletal Muscle

10 Skeletal Muscle Each muscle is a discrete organ composed of muscle tissue, blood vessels, nerve fibers, and connective tissue

11 Skeletal Muscle The three connective tissue sheaths are:
Endomysium – fine sheath of connective tissue surrounding each muscle fiber Perimysium – fibrous connective tissue that surrounds groups of muscle fibers Fascicles

12 Skeletal Muscle Epimysium
A layer of fibrous connective tissue that surrounds the entire muscle

13 Skeletal Muscle Figure 9.2a

14 Skeletal Muscle: Nerve and Blood Supply
Each muscle is served by one nerve, an artery, and one or more veins Each skeletal muscle fiber is supplied with a nerve ending that controls contraction Contracting fibers require continuous delivery of oxygen and nutrients via arteries Wastes must be removed via veins

15 Skeletal Muscle: Attachments
Most skeletal muscles span joints and are attached to bone in at least two places When muscles contract the movable bone, the muscle’s insertion moves toward the immovable bone, the muscle’s origin

16 Skeletal Muscle: Attachments
Muscles attach: Directly – epimysium of the muscle is fused to the periosteum of a bone Indirectly – connective tissue wrappings extend beyond the muscle as a tendon or aponeurosis

17 Microscopic Anatomy of a Skeletal Muscle Fiber
Fibers are up to hundreds of centimeters long Each cell is a syncytium produced by fusion of embryonic cells

18 Microscopic Anatomy of a Skeletal Muscle Fiber
Sarcoplasm has numerous glycosomes and a unique oxygen-binding protein called myoglobin Fibers contain the usual organelles, myofibrils, sarcoplasmic reticulum, and T tubules

19 Skeletal muscle fibers
Sarcoplasmic reticulum (modified ER) T-tubules and myofibrils aid in contraction T-tubules are continuous with the sarcolemma and filled with extracellular fluid

20 Sarcoplasmic Reticulum (SR)
SR - smooth endoplasmic reticulum that mostly runs longitudinally and surrounds each myofibril Terminal cisterna Terminal enlargement of the SR Functions in the regulation of intracellular calcium levels

21 Sarcoplasmic Reticulum (SR)
Figure 9.5

22 T Tubules T tubules are continuous with the sarcolemma
T tubule proteins act as voltage sensors They conduct impulses to the deepest regions of the muscle These impulses signal for the release of Ca2+ from adjacent terminal cisternae

23 T Tubules They penetrate into the cell’s interior at each A band–I band junction Filled with extracellular fluid Triad Pair of terminal cisterna T-tubule

24 Myofibrils Myofibrils are densely packed, rodlike contractile elements
They make up most of the muscle volume The arrangement of myofibrils within a fiber is such that a perfectly aligned repeating series of dark A bands and light I bands is evident

25 Myofibrils Figure 9.3b

26 Sarcomeres The smallest contractile unit of a muscle
It goes from Z disc to Z disc Composed of thin (actin) and thick (myosin) myofilaments

27 Sarcomeres Figure 9.3c

28 Sarcomere A bands Formed by thin and thick filaments M line
Central part of the thick filament Protein that keeps the thick filaments together H zone On each side of the M line Formed by thick filament Zone of overlap Thin filament between thick filaments

29 Sarcomere I band Formed by thin filaments Z line
actinin proteins that anchors the thin filaments Titin keep the thick and thin filaments aligned and prevents the muscle fiber from overstretching From Z line to Z line

30 Titin Keep the thick and thin filaments aligned
Prevents the muscle fiber from overstretching Helps the sarcomere to restore to resting length after contraction

31 Myofilaments: Banding Pattern
Figure 9.3c,d

32 Ultrastructure of Myofilaments: Thick Filaments
Thick filaments are composed of the protein myosin Each myosin molecule has a tail and two globular heads Tails – two interwoven, heavy polypeptide chains Heads – two smaller, light polypeptide chains called cross bridges

33 Thick Filaments Figure 9.4a,b

34 Thin Filaments Thin filaments are chiefly composed of
G actin - globular subunits The G actin contain the active sites to which myosin heads attach during contraction

35 Thin Filaments Tropomyosin
Double stranded protein that covers the active sites of the G protein Troponin – composed of 3 globular subunits One subunit binds Ca One subunit binds to tropomyosin One subunit binds to G actin

36 Ultrastructure of Myofilaments: Thin Filaments
Figure 9.4c

37 Arrangement of the Filaments in a Sarcomere
Longitudinal section within one sarcomere Figure 9.4d

38 Sliding Filament Model of Contraction
Thin filaments slide past the thick ones so that the actin and myosin filaments overlap to a greater degree In the relaxed state, thin and thick filaments overlap only slightly Upon stimulation, myosin heads bind to actin and sliding begins

39 Sliding Filament Model of Contraction
Each myosin head binds and detaches several times during contraction, acting like a ratchet to generate tension and propel the thin filaments to the center of the sarcomere As this event occurs throughout the sarcomeres, the muscle shortens

40 Skeletal Muscle Contraction
In order to contract, a skeletal muscle must: Be stimulated by a nerve ending Propagate an electrical current, or action potential, along its sarcolemma Have a rise in intracellular Ca2+ levels, the final trigger for contraction

41 Skeletal Muscle Contraction
Linking the electrical signal to the contraction is excitation-contraction coupling

42 Nerve Stimulus of Skeletal Muscle
Skeletal muscles are stimulated by motor neurons of the somatic nervous system Axons of these neurons travel in nerves to muscle cells Axons of motor neurons branch profusely as they enter muscles Each axonal branch forms a neuromuscular junction with a single muscle fiber

43 Neuromuscular Junction
Formed from: Axonal endings whith synaptic vesicles that contain acetylcholine The motor end plate of a muscle, which is a specific part of the sarcolemma that contains ACh receptors and helps form the neuromuscular junction

44 Neuromuscular Junction
Though exceedingly close, axonal ends and muscle fibers are always separated by a space called the synaptic cleft

45 Neuromuscular Junction
Figure 9.7 (a-c)

46 Neuromuscular Junction
When a nerve impulse reaches the end of an axon at the neuromuscular junction: Voltage-regulated calcium channels open and allow Ca2+ to enter the axon Ca2+ inside the axon terminal causes axonal vesicles to fuse with the axonal membrane

47 Neuromuscular Junction
This fusion releases ACh into the synaptic cleft via exocytosis ACh diffuses across the synaptic cleft to ACh receptors on the sarcolemma Binding of ACh to its receptors initiates an action potential in the muscle

48 Role of Acetylcholine (Ach)
ACh binds its receptors at the motor end plate Binding opens chemically (ligand) gated channels that allows Na+ and K+ movements Na+ diffuses in, K+ diffuse out and the interior of the sarcolemma becomes less negative

49 Depolarization Initially, this is a local electrical event called end plate potential Later, it ignites an action potential that spreads in all directions across the sarcolemma

50 Destruction of Acetylcholine
ACh bound to ACh receptors is quickly destroyed by the enzyme acetylcholinesterase This destruction prevents continued muscle fiber contraction in the absence of additional stimuli

51 Muscles and Muscle Tissue

52 Excitation-Contraction Coupling
Once generated, the action potential: Is propagated along the sarcolemma Travels down the T tubules Triggers Ca2+ release from terminal cisternae

53 Excitation-Contraction Coupling
Ca2+ binds to troponin and causes Tropomyosin to expose the active site of G protein

54 Excitation-Contraction Coupling
Myosin cross bridges alternately attach and detach Thin filaments move toward the center of the sarcomere Hydrolysis of ATP powers this cycling process Ca2+ is removed into the SR, tropomyosin blockage is restored, and the muscle fiber relaxes

55 Figure 9.10 Synaptic cleft vesicle 1 ACh SR tubules (cut) SR 2 6 3
ADP Pi Net entry of Na+ Initiates an action potential which is propagated along the sarcolemma and down the T tubules. T tubule Sarcolemma SR tubules (cut) Synaptic cleft vesicle Axon terminal ACh Neurotransmitter released diffuses across the synaptic cleft and attaches to ACh receptors on the sarcolemma. Action potential in T tubule activates voltage-sensitive receptors, which in turn trigger Ca2+ release from terminal cisternae of SR into cytosol. Calcium ions bind to troponin; troponin changes shape, removing the blocking action of tropomyosin; actin active sites exposed. Contraction; myosin heads alternately attach to actin and detach, pulling the actin filaments toward the center of the sarcomere; release of energy by ATP hydrolysis powers the cycling process. Removal of Ca2+ by active transport into the SR after the action potential ends. SR Tropomyosin blockage restored, blocking myosin binding sites on actin; contraction ends and muscle fiber relaxes. Ca2+ 1 2 3 4 5 6 Figure 9.10

56 Sequential Events of Contraction
Cross bridge formation – myosin cross bridge attaches to actin filament Working (power) stroke – myosin head pivots and pulls actin filament toward M line 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

57 Figure 9.12 ATP ADP hydrolysis Pi Myosin head (high-energy
configuration) Myosin head attaches to the actin myofilament, forming a cross bridge. Thin filament As ATP is split into ADP and Pi, the myosin head is energized (cocked into the high-energy conformation). Inorganic phosphate (Pi) generated in the previous contraction cycle is released, initiating the power (working) stroke. The myosin head pivots and bends as it pulls on the actin filament, sliding it toward the M line. Then ADP is released. (low-energy As new ATP attaches to the myosin head, the link between myosin and actin weakens, and the cross bridge detaches. Thick filament 1 4 2 3 Figure 9.12

58 Contraction of Skeletal Muscle Fibers
Contraction – refers to the activation of myosin’s cross bridges (force-generating sites) Shortening occurs when the tension generated by the cross bridge exceeds forces opposing shortening Contraction ends when cross bridges become inactive, the tension generated declines, and relaxation is induced

59 Motor Unit: The Nerve-Muscle Functional Unit
A motor unit is a motor neuron and all the muscle fibers it supplies The number of muscle fibers per motor unit can vary from four to several hundred Muscles that control fine movements (fingers, eyes) have small motor units

60 Motor Unit: The Nerve-Muscle Functional Unit
Figure 9.13a

61 Motor Unit: The Nerve-Muscle Functional Unit
Large weight-bearing muscles (thighs, hips) have large motor units Muscle fibers from a motor unit are spread throughout the muscle; therefore, contraction of a single motor unit causes weak contraction of the entire muscle

62 Muscle Twitch Cycle of contraction, relaxation produced by a single brief threshold stimulus There are three phases to a muscle twitch Latent period Period of contraction Period of relaxation

63 Phases of a Muscle Twitch
Latent period – first few msec after stimulus; EC coupling taking place Period of contraction – cross bridges from; muscle shortens Period of relaxation – Ca2+ reabsorbed; muscle tension goes to zero Figure 9.14a

64 Muscle Twitch Comparisons
Figure 9.14b

65 Treppe: The Staircase Effect
Repeated stimulation of the same strength after relaxation phase has been completed It causes increased contraction There is increasing availability of Ca2+ in the sarcoplasm Muscle enzyme systems become more efficient because heat is increased as muscle contracts

66 Treppe: The Staircase Effect
Figure 9.18

67 Graded Muscle Responses
Graded muscle responses are: Variations in the degree of muscle contraction according to the demand imposed on the muscle Responses are graded by: Changing the frequency of stimulation Changing the strength of the stimulus

68 Changing the Frequency of Stimulation
Repeated stimulation before relaxation phase has been completed will cause: Wave summation Frequently delivered stimuli where muscle does not have time to completely relax It increases contractile force

69 Changing the Frequency of Stimulation
Incomplete tetanus More rapidly delivered stimuli than in wave summation. Only some muscle relaxation allowed Complete tetanus If stimuli are given even more rapidly than in incomplete tetanus. No muscle relaxation allowed Figure 9.15

70 Frequency of Stimulation

71 Changing the Strength of Stimulation
Threshold stimulus – the stimulus strength at which the first muscle contraction occurs Multiple motor unit summation or recruitment Muscle contracts more vigorously as stimulus strength is increased It brings more and more muscle fibers into play

72 Threshold and Recruitment
Figure 9.16

73 Size Principle Figure 9.17

74 Muscle Tone Muscle tone:
Is the constant, slightly contracted state of all muscles, which does not produce active movements Keeps the muscles firm, healthy, and ready to respond to stimulus Keeps posture Stabilizes bones and joints

75 Muscle Tone Spinal reflexes account for muscle tone by:
Activating one motor unit and then another Responding to activation of stretch receptors in muscles and tendons

76 Isotonic Contractions
The muscle changes in length Once tension is sufficient to move the load it will stay the same Types of isotonic contractions Concentric contractions – the muscle shortens because the muscle tension is greater than the load

77 Isotonic Contractions
Eccentric contractions – the muscle elongates because the peak tension developed is less than the load The muscle elongates because the contraction of opposing muscles or the gravity

78 Isotonic Contractions
Figure 9.19a

79 Isometric Contractions
Tension increases to the muscle’s capacity, but the muscle neither shortens nor lengthens Occurs if the load is greater than the tension the muscle is able to develop

80 Isometric Contractions
Figure 9.19b

81 Muscle Metabolism: Energy for Contraction
ATP is the only source used directly for contractile activity As soon as available stores of ATP are hydrolyzed (4-6 seconds), they are regenerated by: The interaction of ADP with creatine phosphate (CP) Aerobic respiration Anaerobic glycolysis

82 Muscle Contraction requires large amounts of energy
Immediate energy ATP + Creatine = ADP + creatine phosphate Creatine phosphate releases stored energy to convert ADP to ATP Aerobic metabolism provides most ATP needed for contraction Long-term energy Krebs cycle in mitochondria

83 Muscle Metabolism: Energy for Contraction
Figure 9.20

84 Muscle Contraction requires large amounts of energy
Anaerobic metabolism When muscle contractile activity reaches 70% of maximum: Bulging muscles compress blood vessels Oxygen delivery is impaired Pyruvic acid is converted into lactic acid

85 Muscle Contraction requires large amounts of energy
The lactic acid: Diffuses into the bloodstream Is picked up and used as fuel by the liver, kidneys, and heart Is converted back into pyruvic acid by the liver Short-term energy

86 Muscle Fatigue The muscle is in a state of physiological inability to contract Muscle fatigue occurs when: ATP production fails to keep pace with ATP use There is a relative deficit of ATP, causing contractures Lactic acid accumulates in the muscle Ionic imbalances are present

87 Muscle Fatigue Intense exercise produces rapid muscle fatigue (with rapid recovery) Na+-K+ pumps cannot restore ionic balances quickly enough Low-intensity exercise produces slow-developing fatigue SR is damaged and Ca2+ regulation is disrupted

88 Muscles and Muscle Tissue

89 Oxygen Debt Oxygen debt – the extra amount of O2 needed for the above restorative processes

90 Oxygen Debt Vigorous exercise causes dramatic changes in muscle chemistry For a muscle to return to a resting state: Oxygen reserves must be replenished Lactic acid must be converted to pyruvic acid Glycogen stores must be replaced ATP and CP reserves must be resynthesized

91 Heat Production During Muscle Activity
Only 40% of the energy released in muscle activity is useful as work The remaining 60% is given off as heat Dangerous heat levels are prevented by radiation of heat from the skin and sweating

92 Force of Muscle Contraction
The force of contraction is affected by: The number of muscle fibers contracting – the more motor fibers in a muscle, the stronger the contraction The relative size of the muscle – the bulkier the muscle, the greater its strength

93 Force of Muscle Contraction
Degree of muscle stretch – muscles contract strongest when muscle fibers are % of their normal resting length

94 Force of Muscle Contraction
Figure 9.21a–c

95 Length Tension Relationships
Figure 9.22

96 Muscle Fiber Type: Functional Characteristics
Speed of contraction – determined by speed in which ATPases split ATP ATP-forming pathways Oxidative fibers – use aerobic pathways Glycolytic fibers – use anaerobic glycolysis These two criteria define three categories – slow oxidative fibers, fast oxidative fibers, and fast glycolytic fibers

97 Fast versus Slow Fibers
Figure 10.21

98 Slow oxidative fibers Half the diameter of fast fibers
Take three times as long to contract after stimulation Abundant mitochondria aerobic metabolism Extensive capillary supply High concentrations of myoglobin Fatigue resistant Red color

99 Fast glycolytic fibers
Large in diameter Contain densely packed myofibrils Large glycogen reserves Relatively few mitochondria Produce rapid, powerful contractions of short duration Fast fatigue White color

100 Fast oxidative or Intermediate fibers
Similar to fast fibers Greater resistance to fatigue Pink color Intermediate glycogen storage many mitochondria Many capillaries Uses mainly aerobic metabolism

101 Load and Contraction Figure 9.23

102 Effects of Aerobic Exercise
Aerobic exercise results in an increase of: Muscle capillaries Number of mitochondria Myoglobin synthesis Mainly in slow oxidative fibers Not significant muscle hypertrophy

103 Effects of Resistance Exercise
Resistance exercise (typically anaerobic) results in: Muscle hypertrophy Increased mitochondria, myofilaments, and glycogen stores Increases muscle fibers

104 The Overload Principle
Forcing a muscle to work promotes increased muscular strength Muscles adapt to increased demands Muscles must be overloaded to produce further gains

105 Smooth Muscle Composed of spindle-shaped fibers with a diameter of 2-10 m and lengths of several hundred m Lack the coarse connective tissue sheaths of skeletal muscle, but have fine endomysium Organized into two layers (longitudinal and circular) of closely apposed fibers

106 Smooth Muscle Found in walls of hollow organs (except the heart)
Have essentially the same contractile mechanisms as skeletal muscle

107 Smooth Muscle Figure 9.24

108 Peristalsis When the longitudinal layer contracts, the organ dilates and shorten When the circular layer contracts, the organ elongates and constrict Peristalsis – alternating contractions and relaxations of smooth muscles that mix and squeeze substances through the lumen of hollow organs

109 Innervation of Smooth Muscle
Smooth muscle lacks neuromuscular junctions Innervating nerves have bulbous swellings called varicosities Varicosities release neurotransmitters into wide synaptic clefts called diffuse junctions

110 Innervation of Smooth Muscle
Figure 9.25

111 Microscopic Anatomy of Smooth Muscle
SR is less developed than in skeletal muscle and lacks a specific pattern T tubules are absent Plasma membranes have pouchlike infoldings called caveoli

112 Microscopic Anatomy of Smooth Muscle
Ca2+ is sequestered in the extracellular space near the caveoli, allowing rapid influx when channels are opened There are no visible striations and no sarcomeres Thin and thick filaments are present

113 Proportion and Organization of Myofilaments in Smooth Muscle
Ratio of thick to thin filaments is much lower than in skeletal muscle Thick filaments have heads along their entire length There is no troponin complex

114 Proportion and Organization of Myofilaments in Smooth Muscle
Thick and thin filaments are arranged diagonally, causing smooth muscle to contract in a corkscrew manner Noncontractile intermediate filament bundles attach to dense bodies (analogous to Z discs) at regular intervals

115 Smooth Muscle

116 Proportion and Organization of Myofilaments in Smooth Muscle
Figure 9.26

117 Contraction of Smooth Muscle
Whole sheets of smooth muscle exhibit slow, synchronized contraction They contract in unison, reflecting their electrical coupling with gap junctions Action potentials are transmitted from cell to cell

118 Contraction of Smooth Muscle
Some smooth muscle cells: Act as pacemakers and set the contractile pace for whole sheets of muscle Are self-excitatory and depolarize without external stimuli

119 Contraction Mechanism
Actin and myosin interact according to the sliding filament mechanism The final trigger for contractions is a rise in intracellular Ca2+ Ca2+ is released from the SR and from the extracellular space Ca2+ interacts with calmodulin and myosin light chain kinase to activate myosin

120 Contraction Mechanism
Ca2+ binds to calmodulin and activates it Activated calmodulin activates the kinase enzyme Activated kinase transfers phosphate from ATP to myosin cross bridges Phosphorylated myosin heads form cross bridges with actin

121 Contraction Mechanism
Power stroke is executed Smooth muscle relaxes when intracellular Ca2+ levels drop

122 Figure 9.27

123 Special Features of Smooth Muscle Contraction
Unique characteristics of smooth muscle include: Smooth muscle tone Slow, prolonged contractile activity Low energy requirements Response to stretch

124 Response to Stretch Smooth muscle exhibits a phenomenon called stress-relaxation response in which: Smooth muscle responds to stretch only briefly, and then adapts to its new length The new length, however, retains its ability to contract This enables organs such as the stomach and bladder to temporarily store contents

125 Hyperplasia Certain smooth muscles can divide and increase their numbers by undergoing hyperplasia This is shown by estrogen’s effect on the uterus At puberty, estrogen stimulates the synthesis of more smooth muscle, causing the uterus to grow to adult size

126 Types of Smooth Muscle: Single Unit
The cells of single-unit smooth muscle, commonly called visceral muscle: Contract rhythmically as a unit Are electrically coupled to one another via gap junctions Often exhibit spontaneous action potentials Are arranged in opposing sheets and exhibit stress-relaxation response

127 Types of Smooth Muscle: Multiunit
Multiunit smooth muscles are found: In large airways to the lungs In large arteries In arrector pili muscles In the intrinsic eye muscles

128 Types of Smooth Muscle: Multiunit
Their characteristics include: Rare gap junctions Infrequent spontaneous depolarizations Structurally independent muscle fibers A rich nerve supply, which, with a number of muscle fibers, forms motor units Graded contractions in response to neural stimuli - recruitment

129 Table 9.3.3

130 Table 9.3.4

131 Developmental Aspects
Muscle tissue develops from embryonic mesoderm called myoblasts Multinucleated skeletal muscle cells form by fusion of myoblasts Syncytium The growth factor agrin stimulates the clustering of ACh receptors at newly forming motor end plates

132 Developmental Aspects
As muscles are brought under the control of the somatic nervous system, the numbers of fast and slow fibers are also determined Cardiac and smooth muscle myoblasts do not fuse but develop gap junctions at an early embryonic stage

133 Developmental Aspects: Regeneration
Cardiac and skeletal muscle become amitotic, but can lengthen and thicken Myoblastlike satellite cells show very limited regenerative ability Cardiac cells lack satellite cells Smooth muscle has good regenerative ability

134 Developmental Aspects: After Birth
Muscular development reflects neuromuscular coordination Development occurs head-to-toe, and proximal-to-distal Peak natural neural control of muscles is achieved by midadolescence Athletics and training can improve neuromuscular control

135 Developmental Aspects: Male and Female
There is a biological basis for greater strength in men than in women Women’s skeletal muscle makes up 36% of their body mass Men’s skeletal muscle makes up 42% of their body mass

136 Developmental Aspects: Male and Female
These differences are due primarily to the male sex hormone testosterone With more muscle mass, men are generally stronger than women

137 Developmental Aspects: Age Related
With age, connective tissue increases and muscle fibers decrease Muscles become stringier and more sinewy By age 80, 50% of muscle mass is lost (sarcopenia)

138 Developmental Aspects: Age Related
Regular exercise reverses sarcopenia Aging of the cardiovascular system affects every organ in the body Atherosclerosis may block distal arteries, leading to intermittent claudication and causing severe pain in leg muscles

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