276-311)
2.3.1 Describe the microscopic structure of skeletal muscle fibers and explain the cellular mechanisms of excitation-contraction 2.3.2 Describe the neuromuscular junction 2.3.3 Describe the contractile properties of skeletal muscle (motor unit, isotonic & isometric contractions, spatial & temporal summation, etc) 2.3.4 Associate various muscle types with their metabolism and their speed of contraction and rate of fatigue 2.3.5 Compare the properties of smooth muscle with those of skeletal muscle
Muscle Similarities
Skeletal and smooth muscle cells are elongated (not cardiac muscles) and are called muscle fibers Muscle contraction depends on two kinds of myofilaments actin and myosin Muscle terminology is similar
Sarcolemma muscle plasma membrane Sarcoplasm cytoplasm of a muscle cell Prefixes myo, mys, and sarco all refer to muscle
Pacemaker
Can contract rapidly; Tires easily & must rest; Strong but Adaptable
2.3.1.2 list & briefly describe 4 muscle functions as well as 4 functional characteristics of muscle Muscle Functions 1. Generate movement: locomotion, manipulation, blood flow & pressure, respiration, propelling of food, urine, etc..
Skeletal Muscle
Each muscle is a discrete organ composed of muscle tissue, blood vessels, nerve fibers, and connective tissue
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
Table 9.1a
Endomysium fine sheath of connective tissue composed of reticular fibers surrounding each muscle fiber Perimysium fibrous connective tissue that surrounds groups of muscle fibers called fascicles Epimysium an overcoat of dense irregular connective tissue that surrounds the entire muscle
2 m long
Table 9.1b
Each fiber is a long, cylindrical cell with multiple nuclei just beneath the sarcolemma Fibers are 10 to 100 m in diameter, and up to 30 cm long Each cell is a syncytium produced by fusion of embryonic cells
Sarcoplasm has numerous glycosomes and a unique oxygen-binding protein called myoglobin Fibers contain the usual organelles, and myofibrils, sarcoplasmic reticulum, and T tubules
Myofibrils
Myofibrils are densely packed, rodlike contractile elements They make up most of the muscle cell volume (80%) 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 (striated appearance)
Sarcomeres
Z-disc coin-shaped sheet of proteins (alpha-actinin; desmin IF) that (i) anchors the thin filaments and (ii) connects myofibrils to one another Thin filaments do not overlap with thick filaments in the lighter H zone (and no myosin head) M lines appear darker due to the presence of the protein myomesin (holding thick filaments together)
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Sarcomeres
Thick filament Thin filament In the center of the sarcomere, the thick filaments lack myosin heads. Myosin heads are present only in areas of myosin-actin overlap. Thick filament Thin filament Each thick filament consists of many A thin filament consists of two strands myosin molecules whose heads protrude of actin subunits twisted into a helix at opposite ends of the filament. plus two types of regulatory proteins (troponin and tropomyosin). Portion of a thick filament Portion of a thin filament Myosin head Tropomyosin Troponin Actin
Actin-binding sites
ATPbinding site Heads Tail Actin subunits Actin subunits
Figure 9.3
Thick filaments are composed of the protein myosin Each myosin molecule has a rod-like tail and two globular heads
Tails two interwoven, heavy polypeptide chains Heads two smaller, light polypeptide chains called cross bridges Heads bear actin-binding site and ATPase activity
Figure 9.4a,b
Figure 9.4c
Thin filaments are chiefly composed of the protein actin (microfilaments) Each actin molecule is a helical polymer of globular subunits called G actin (F actin is the fibrous or filamentous form) The subunits contain the active sites to which myosin heads attach during contraction
Tropomyosin and troponin (TnI, TnT and TnC) are regulatory subunits bound to actin
2 - Contracted: thin filaments penetrate more deeply into A band - Z discs pulled toward thick filaments
Overall: distance between Z discs reduced I bands shorten H zones disappear A bands move closer together, but stay same length
C. Fascicle;
D. Fiber; H. Sarcolemma;
G. Perimysium; K. Tendon
1. 2. 3. 4.
Connective tissue surrounding a fascicle. Contractile unit of muscle. A muscle cell. Thin connective tissue investing each muscle cell.
5.
6. 7.
J. Carnegie, UofO
SR is an elaborate, smooth endoplasmic reticulum that mostly runs longitudinally and surrounds each myofibril
T Tubules
Figure 9.5
T tubules are continuous with the sarcolemma They conduct impulses to the deepest regions of the muscle (at each A bandI band junction) These impulses signal for the release of Ca2+ from adjacent terminal cisternae
Triad Relationships
T tubules and SR provide tightly linked signals for muscle contraction A double zipper of integral membrane proteins protrudes into the intermembrane space T tubule proteins act as voltage sensors
SR foot proteins are receptors that regulate Ca2+ release from the SR cisternae Copyright 2006 Pearson Education, Inc., publishing as Benjamin Cummings
Skeletal muscles are stimulated by somatic motor neurons Axons of motor neurons travel from the central nervous system via nerves to skeletal muscles
Myelinated axon of motor neuron Axon terminal of neuromuscular junction Sarcolemma of the muscle fiber
2 Voltage-gated
Ca2+
Ca2+
Axon terminal of motor neuron Fusing synaptic vesicles
Ca2+
ACh
4 Acetylcholine, a neurotransmitter, diffuses across the synaptic cleft and binds to receptors in the sarcolemma. 5 ACh binding opens ion
channels that allow simultaneous passage of Na+ into the muscle fiber and K+ out of the muscle fiber.
Na+ K+
Ach
Acetylcholinesterase
K+
PLAY
Synaptic cleft
K+
Action potential
+ + +++ +
generation of the end plate potential on the sarcolemma Sarcoplasm of muscle fiber
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1 Local depolarization:
K+ 3 Repolarization
Figure 9.9
One sarcomere
Ca2+
Actin Troponin Tropomyosin blocking active sites Myosin 3 - Calcium binds to troponin and removes the blocking action of tropomyosin.
Ca2+
The aftermath
1 Action potential is
Ca2+
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Ca2+
Troponin
The aftermath
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3 Calcium binds to
The aftermath
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Figure 9.11
3 Calcium binds to
4 Contraction begins
Myosin cross bridge The aftermath
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Figure 9.12
Contraction refers to the activation of myosins cross bridges (force-generating sites) Shortening occurs when the tension generated by the cross bridge exceeds forces opposing shortening (load)
When nervous stimulation ceases, Ca2+ is pumped back into the SR and contraction ends
Contraction ends when cross bridges become inactive, the tension generated declines, and relaxation is induced (Ca2+ removal => blockage restored)
http://www.youtube.com/watch?v=DeTen4GVb-Q
http://www.youtube.com/watch?v=DeTen4GVb-Q
1. Myosin heads bind to active sites on actin molecules
2. ATP is hydrolyzed 3. Myosin heads return to high-energy shape, ready for the next power stroke
IV.
2.3.3.1 define motor unit; describe the influences of wave summation and motor unit summation on the contractile response of skeletal muscle; define tetanus in terms of muscular contraction
Fig. 9.12
Innervated fibers not clustered 1 motor unit = weak contraction of the entire muscle!
A muscle twitch is the response of a muscle to a single, brief threshold stimulus Latent period first few msec after stimulus; EC coupling taking place
Period of contraction cross bridges form; muscle shortens Period of relaxation Ca2+ reabsorbed; muscle tension goes to zero
The force exerted by a contracting muscle on an object is called muscle tension The opposing force exerted on the muscle is called the load
Myogram
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Figure 9.14a
Figure 9.14b
Variations in the degree of muscle contraction Required for proper control of skeletal movement
Speed of stimulation: wave summation & tetanus: rapid rate of stimuli: each contraction builds on the previous one ([Ca2+]; Heat) but AP refractory period is always honored (repolarization) tetanus: fused contractions - inter-stimulus interval too short to allow inter-twitch muscle relaxation - eventually followed by muscle fatigue
(Rare)
Fig. 9.15
Threshold stimulus the stimulus strength at which the first observable muscle contraction occurs Beyond threshold, muscle contracts more vigorously as stimulus strength is increased Force of contraction is precisely controlled by multiple motor unit summation This phenomenon, called recruitment, brings more and more muscle fibers into play
Not random
In the body
Size Principle
Motor units with the smallest fibers are controlled by highly excitable motor neurons activated first
As contractile strength increases larger and larger muscle fibers are recruited
Figure 9.17
Muscle tone:
It 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 Activating one motor unit and then another Responding to activation of stretch receptors in muscles and tendons (Chapter 13)
1. After ACh attaches to its receptors at the neuromuscular junction, the next step is: a) K+-gated channels open b) Ca++ binds to troponin c) the T tubules depolarize d) cross bridges attach e) ATP is hydrolyzed 2. Which of these bands or lines does NOT narrow when skeletal muscle contracts? a) H zone b) A band c) I band d) M line e) b and d 3. is a continuous contraction that shows no evidence of relaxation. a) tetanus b) all or none c) isometric J. Carnegie, UofO d) treppe
2.3.3.2 differentiate between isotonic and isometric contractions, giving an example of each (1) isotonic: muscle changes in length & moves load - what are concentric vs eccentric isotonic contractions? NB: eccentric contractions are 50% more forceful, use less ATP/O2 & fewer muscle fibers but more prone to delayed-onset soreness than concentric contractions
2.3.3.2 differentiate between isotonic and isometric contractions, giving an example of each (2) isometric: tension increases but muscle remains same length most body movements are a mix of isotonic and isometric contractions realize that in isotonic contractions, the thin filaments are sliding, but in isometric contractions the cross bridges are generating force, but do not move thin filaments
2.3.3.3 Define the optimal length-tension relationship for muscle in terms of muscle anatomy
Factors influencing the Force of Muscle Contraction Degree of muscle stretch length/tension relationship
Fig. 9.21
Fig. 9.22
2.3.3.4 Indicate the influence of load on the velocity and duration of skeletal muscle contraction
Velocity & Duration of Contraction depends on: Load: greater load = longer latent period = slower contraction = shorter contraction duration Fig. 9.23
(a)
Direct phosphorylation
CP
Creatine
ATP
Creatine Kinase
Oxygen use: None Products: 1 ATP per CP, creatine Duration of energy provision: 15 seconds
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Figure 9.19a
Anaerobic glycolysis (Fast Glycolytic Fibers): only 2 ATP/glucose but O2 not used and is 2.5x faster than aerobic pathway Together, ATP, CP and Anaerobic glycolysis can support strenuous muscle activity for ~ 1 min. usually pyruvic acid enters aerobic pathway
Diffuses into the bloodstream Is picked up and used as fuel by the liver, kidneys, and heart
Figure 9.19
Energy systems used during sports activities: weight lifting, diving, sprinting: ATP & CP - very short surge of power tennis, soccer, 100 m swim: almost entirely anaerobic - on-and-off; burstlike marathon runs, jogging: mainly aerobic; but anaerobic may function until aerobic reaches full efficiency Mainly endurance rather than power aerobic endurance: length of time a muscle can use aerobic anaerobic threshold: point at which muscle converts to anaerobic
Muscle fatigue the muscle is in a state of physiological inability to contract Muscle fatigue occurs when:
There is a relative deficit of ATP, causing contractures (cramps! = myosin head can not be released)
Lactic acid accumulates in the muscle
Na+-K+ pumps cannot restore ionic balances quickly enough (intense exercise)
Oxygen Debt
Vigorous exercise causes dramatic changes in muscle chemistry For a muscle to return to a resting state:
Oxygen debt the extra amount of O2 needed for the above restorative processes
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 What is shivering then?
FO SO
FG
Figure 9.24
(O2)
a motor unit consists of just one muscle fiber type but one muscle usually contain a mix of fiber types
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Table 9.2
Effects of Exercise
Muscle capillaries
Number of mitochondria Myoglobin synthesis
Muscle hypertrophy
Check the appropriate boxes to characterize each type of skeletal muscle fiber:
Characteristics Slow Oxidative Fast Glycolytic Fast Oxidative
http://www.youtube.com/watch?v=DeTen4GVb-Q
Table 9.3
Table 9.3
Smooth Muscle
Composed of spindle-shaped fibers with a diameter of 5-10 m and lengths of several hundred m Sk. M. are ~10x wider; 1000x longer Lack the coarse connective tissue sheaths of skeletal muscle, but have fine endomysium Organized into two layers (longitudinal and circular) of closely apposed fibers Parallel to organ Organ Dilates & Contracts
Organ Elongates
Peristalsis
Ca2+ is sequestered in the extracellular space near the caveolae, allowing rapid influx when channels are opened
SR still releases some Ca2+ and contributes to its removal
Table 9.3
Ratio of thick to thin filaments is much lower (1:13) than in skeletal muscle (1:2) Thick filaments have heads along their entire length (more strength) There is no troponin complex
Thick and thin filaments are arranged diagonally (no sarcomere; no striations), causing smooth muscle to contract in a corkscrew manner
Noncontractile intermediate filament bundles attach to dense bodies (analogous to Z discs) atasregular intervals (linked to thin filaments) Copyright 2006 Pearson Education, Inc., publishing Benjamin Cummings
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
Act as pacemakers and set the contractile pace for whole sheets of muscle Are self-excitatory and depolarize without external stimuli
Stimuli can vary Different autonomic nerves release diff. NT NT may excite or inhibit a particular group of smooth muscle cells
Relaxation requires:
Ca2+ detachment from calmodulin
Figure 9.29
Table 9.3
Response to Stretch: Sk. M. can still contract when stretch up to 60%; Smooth muscle can still generate tension if stretch up to 150%!
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 to temporarily store contents
Hyperplasia
Certain smooth muscles can divide and increase their numbers by undergoing hyperplasia (vs Hypertrophy)
At puberty, estrogen stimulates the synthesis of more smooth muscle, causing the uterus to grow to adult size During pregnancy, estrogen stimulates uterine growth to accommodate the increasing size of the growing fetus
Contract rhythmically as a unit Are electrically coupled to one another via gap junctions no recruitment Often exhibit spontaneous action potentials (presence of some Pacemaker cells)
Rare gap junctions Infrequent spontaneous depolarizations Structurally independent muscle fibers A rich nerve supply (still Autonomic/Involuntary), which, with a number of muscle fibers, forms motor units Graded contractions in response to neural stimuli recruitment Can also reponsd to Hormonal control
1.
The first energy source used to regenerate ATP when muscles are extremely active is: a) fatty acids b) glucose c) creatine phosphate d) pyruvic acid When paying back the oxygen debt: a) lactic acid is formed b) lactic acid is converted to pyruvic acid c) ATP formation requires creatine phosphate d) muscle cells utilize glycogen reserves Holding up the corner of a heavy couch to vacuum under it involves which type(s) of contraction? a) tetanic b) isotonic c) isometric d) tone e) a and c
J. Carnegie, UofO
2.
3.