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(Chapter 9; pp.

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

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Pacemaker

Can contract rapidly; Tires easily & must rest; Strong but Adaptable

Neural input can Increase rate

Slow, sustained contraction

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..

2. Maintain posture: constantly working against gravity


3. Joint stabilization: eg: shoulders, knees when moving parts of skeleton 4. Generation of heat: maintenance of body temperature esp: skeletal muscle (at least 40% of body mass) Functional Characteristics of Muscle 1. Excitability (Irritability): ability to receive and respond to a stimulus stimulus usually a chemical (NT, hormone, pH) response = AP along sarcolemma + muscle contraction 2. Contractility: ability to shorten forcibly when adequately stimulated 3. Extensibility: ability to be stretched or extended 4. Elasticity: ability to resume resting length after being stretched

Skeletal Muscle

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

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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

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Structure and Organization of Skeletal Muscle

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Table 9.1a

Skeletal Muscle: 3 connective tissue sheaths

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

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Structure and Organization of Skeletal Muscle

2 m long

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Table 9.1b

Microscopic Anatomy of a Skeletal Muscle Fiber

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

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Microscopic Anatomy of a Skeletal Muscle Fiber

Sarcoplasm has numerous glycosomes and a unique oxygen-binding protein called myoglobin Fibers contain the usual organelles, and myofibrils, sarcoplasmic reticulum, and T tubules

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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

Thick filaments extend the entire length of an A band


Thin filaments extend across the I band and partway into the A band

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

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Longitudinal section of filaments within one sarcomere of a myofibril

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

Flexible hinge region Myosin molecule

Active sites for myosin attachment

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Ultrastructure of Myofilaments: Thick Filaments

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

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Ultrastructure of Myofilaments: Thin Filaments

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

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Sliding Filament Model of Contraction


muscle fiber shortens because sarcomeres shorten; filaments remain same length thin filaments slide over thick filaments 1 - Relaxed: only slight overlap of thick & thin filaments

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

A. Endomysium; E. Myofilament; I. Sarcomere;

B. Epimysium; F. Myofibril; J. Sarcoplasm;

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.

Plasma membrane of the muscle cell.


Cordlike extension of CT beyond muscle - attaches it to bone. A discrete bundle of muscle cells.

J. Carnegie, UofO

Sarcoplasmic Reticulum (SR)


Figure 9.5

SR is an elaborate, smooth endoplasmic reticulum that mostly runs longitudinally and surrounds each myofibril

Paired terminal cisternae form perpendicular cross channels


Functions in the regulation of intracellular calcium levels

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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

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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

Each axon forms several branches as it enters a muscle


Each axon ending forms a neuromuscular junction with a single muscle fiber

Action potential (AP) Nucleus

Myelinated axon of motor neuron Axon terminal of neuromuscular junction Sarcolemma of the muscle fiber

1 Action potential arrives at axon terminal of motor neuron.


channels open and Ca2+ enters the axon terminal.

2 Voltage-gated

Ca2+

Ca2+
Axon terminal of motor neuron Fusing synaptic vesicles

Ca2+

Synaptic vesicle containing ACh Mitochondrion Synaptic cleft

3 Ca2+ entry causes some


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. 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+

Junctional folds of sarcolemma

Sarcoplasm of muscle fiber


Postsynaptic membrane ion channel opens; ions pass.

6 ACh effects are terminated


by its enzymatic breakdown in the synaptic cleft by acetylcholinesterase.

Ach

Degraded ACh Na+

Postsynaptic membrane ion channel closed; ions cannot pass.

Acetylcholinesterase

K+

PLAY

A&P Flix: Events at the Neuromuscular Junction


Figure 9.8

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Axon terminal Open Na+ Channel Na+ Closed K+ Channel

Synaptic cleft

ACh ACh Na+ K+ Na+ K+


++ ++ + +

K+
Action potential

the action potential (AP)

2 Generation and propagation of


Closed Na+ Open K+ Channel Channel Na+

+ + +++ +

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

Events of Excitation-Contraction (E-C) Coupling


Axon terminal of motor neuron Action potential Synaptic cleft is generated ACh Sarcolemma Terminal cisterna of SR
Muscle fiber Ca2+ Triad

One sarcomere

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Figure 9.11, step 1

Events of Excitation-Contraction (E-C) Coupling


Steps in E-C Coupling:
Sarcolemma Voltage-sensitive tubule protein T tubule 1 - Action potential is propagated along the sarcolemma and down the T tubules.

Ca2+ release channel 2 - Calcium ions are released. Terminal cisterna of SR

Ca2+

Actin Troponin Tropomyosin blocking active sites Myosin 3 - Calcium binds to troponin and removes the blocking action of tropomyosin.

Ca2+

Active sites exposed and ready for myosin binding

4 - Contraction begins Myosin cross bridge

The aftermath

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Figure 9.11, step 2

Events of Excitation-Contraction (E-C) Coupling


propagated along the sarcolemma and down the T tubules. Sarcolemma T tubule

1 Action potential is

Steps in E-C Coupling:

Voltage-sensitive tubule protein

Ca2+ release channel Terminal cisterna of SR

Ca2+
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Figure 9.11, step 3

Events of Excitation-Contraction (E-C) Coupling


Actin

Ca2+

Troponin

Tropomyosin blocking active sites Myosin

The aftermath
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Figure 9.11, step 5

Events of Excitation-Contraction (E-C) Coupling


Actin Ca2+ Troponin Tropomyosin blocking active sites Myosin troponin and removes the blocking action of tropomyosin. Active sites exposed and ready for myosin binding

3 Calcium binds to

The aftermath
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Figure 9.11, step 6

Figure 9.11

Events of Excitation-Contraction (E-C) Coupling


Actin
Ca2+ Troponin Tropomyosin blocking active sites Myosin

troponin and removes the blocking action of tropomyosin.


Active sites exposed and ready for myosin binding

3 Calcium binds to

4 Contraction begins
Myosin cross bridge The aftermath
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Figure 9.11, step 7

Each power stroke = 1% shortening Routine contraction = 30-35% shortening


(High Energy) Pulls on actin filament

(Bent Low Energy)

Many such Cross Bridge Cycles are required!


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Figure 9.12

Contraction of Skeletal Muscle Fibers

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

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PUT IN THE CORRECT ORDER:

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

4. Calcium binds to troponin


5. Cycling continues until calcium ions are sequestered by the SR 6. Myosin cross brides detach from actin

7. Troponin changes shape


8. ADP and Pi (inorganic phosphate) are released from the thick filament 9. Myosin heads pull on the thin filaments (power stroke) and slide them toward the centre of the sarcomere 10. ATP binds to the thick filament 11. Tropomyosin is moved exposing active sites on F-actin
J. Carnegie, UofO

IV.

CONTRACTION OF A WHOLE SKELETAL MUSCLE

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

The Motor Unit

motor nerve (at least 1/muscle)


hundreds of motor neuron axons each axon to many axonal terminals each axonal terminal to NMJ of a single muscle fiber (cell) avg. no muscle fibers/neuron = 150 (range = 4 to hundreds) when neuron fires, all fibers contract
MOTOR UNIT = 1 motor neuron + all muscle fibers it supplies

Fig. 9.12

Innervated fibers not clustered 1 motor unit = weak contraction of the entire muscle!

Depends on specific muscle function

Phases of a Muscle Twitch

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

Muscle Twitch Comparisons

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Figure 9.14b

Graded Muscle Responses

Graded muscle responses are:


Variations in the degree of muscle contraction Required for proper control of skeletal movement

Responses are graded by:


Changing the frequency of stimulation Changing the strength of the stimulus

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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

Affects force, but primary goal is to generate smooth, continuous contractions

Treppe Staircase Effect

(Rare)

Fig. 9.15

Muscle Response: Stimulation Strength


In the lab

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

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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

controlled by motor neurons that are least excitable (highest threshold)

Also explains wide range of possible force


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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)

Spinal reflexes account for muscle tone by:


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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

Also depends on Fiber type and Recruitment

The more the merrier!


J. Carnegie, UofO

(a)

Direct phosphorylation

Coupled reaction of creatine phosphate (CP) and ADP Energy source: CP

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:

CP

ADP Creatine kinase

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

When muscle contractile activity reaches 70% of maximum:

Bulging muscles compress blood vessels

Oxygen delivery is impaired


Pyruvic acid is converted into lactic acid

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

Muscle Metabolism: Energy for Contraction


(Slow Oxydative Fibers)

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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 theres an end to every good thing

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 (cramps! = myosin head can not be released)
Lactic acid accumulates in the muscle

Ionic imbalances are present (e.g. dehydration; sweat)

Na+-K+ pumps cannot restore ionic balances quickly enough (intense exercise)

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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 (liver)


Glycogen stores must be replaced ATP and CP reserves must be resynthesized

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

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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 What is shivering then?

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FO SO

FG

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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

Aerobic exercise (endurance) results in an increase of:

Muscle capillaries
Number of mitochondria Myoglobin synthesis

Resistance exercise (typically anaerobic) results in:

Muscle hypertrophy

Increased mitochondria, myofilaments, and glycogen stores

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Check the appropriate boxes to characterize each type of skeletal muscle fiber:
Characteristics Slow Oxidative Fast Glycolytic Fast Oxidative

Rapid twitch rate


Fast myosin ATPases Use mostly aerobic metabolism

Large myoglobin stores


Large glycogen stores Fatigue slowly Fibers are white Fibers are small Fibers contain many capillaries & mitochondria
J. Carnegie, UofO

http://www.youtube.com/watch?v=DeTen4GVb-Q

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Table 9.3

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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

Found in walls of hollow organs (except the heart)

Have essentially the same contractile mechanisms as skeletal muscle

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Innervation of Smooth Muscle


Smooth muscle lack neuromuscular junctions

Innervating nerves have bulbous swellings called varicosities


Varicosities release neurotransmitters into wide synaptic clefts called diffuse junctions

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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 caveolae

1 nucleus per cell!

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

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Table 9.3

Proportion and Organization of Myofilaments in Smooth Muscle

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

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

Some smooth muscle cells (stomach and small intestine):

Act as pacemakers and set the contractile pace for whole sheets of muscle Are self-excitatory and depolarize without external stimuli

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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

e.g. ACh (+) vs NE (-) in the bronchioles


vs NE (+) in most blood vessels

Active transport of Ca2+ into SR and ECF


Dephosphorylation of myosin to reduce myosin ATPase activity

Non-neuronal signals: hormones (gastrin), histamine, hypoxia, excess CO2, low pH

Figure 9.29

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Table 9.3

Special Features of Smooth Muscle Contraction

Unique characteristics of smooth muscle include:

Smooth muscle tone (24h maintenance in blood vessel; visceral organs)


Slow, prolonged contractile activity 30x longer than Sk. M. Low energy requirements 1% the energy cost Few mitos; most ATP generated aerobically Sluggish ATPases (Myosin heads)

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Special Features of Smooth Muscle Contraction

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 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 to temporarily store contents

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Hyperplasia

Certain smooth muscles can divide and increase their numbers by undergoing hyperplasia (vs Hypertrophy)

This is shown by estrogens effect on the uterus

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

Secretory functions: collagen, elastin - make their own CT

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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 no recruitment Often exhibit spontaneous action potentials (presence of some Pacemaker cells)

Few nerve terminals usually where pacermaker cells are


Are arranged in opposing sheets and exhibit stressrelaxation response and pretty much what we saw so far!

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Types of Smooth Muscle: Multiunit

Multiunit smooth muscles are found:


In large airways to the lungs, large arteries


In arrector pili muscles, attached to hair follicles In the internal eye muscles (focus)

Their characteristics include:


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

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Copyright 2006 Pearson Education, Inc., publishing as Benjamin Cummings

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.

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