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3 kinds of muscle tissue
skeletal
cardiac
smooth
Major purpose of muscle
converting the chemical energy in ATP into mechanical energy of motion
functions of muscles
movement of whole body, body parts, organ contents
stability - maintain posture and prevent movement
communication - speech, facial expressions and writing
controls openings and passageways
sphincters
body heat production
Connective tissues of a muscle - fascia
separates neighboring muscles or muscle groups from each other and the subcutaneous tissue
connective tissues of a muscle - epimysium
fibrous sheath surrounding the entire muscle
outer surface grades into the fascia
connective tissues of a muscle - perimysium
surrounds fascicles
carry larger nerves and blood vessels, and stretch receptors
connective tissues of a muscle - endomysium
thin sleeve of loose connective tissue surrounding each muscle fiber
allows room for capillaries and nerve fibers to reach each muscle fiber
connective tissue elements - tendons
attachments between muscle and bone
epimysium surrounding the entire muscle is continuous with collagen fibers of tendons
dense-regular connective tissue composed of collagen fibers
collagen is somewhat extensible and elastic
stretches slightly under tension and recoils when released
resists excessive stretching and protects muscle from injury
returns muscle to its resting length
contribute to power output and muscle efficiency
skeletal muscle shapes
fusiform - thick in middle and tapered at ends
ex. biceps brachii and gastrocnemius
triangular (convergent) - broad at origin and tapering to a narrow insertion
ex. pectoralis major, temporalis
parallel muscles - parallel fascicles
ex. rectus abdominus, zygomaticus major
circular muscles
act as sphincters
ex. orbicularis oris, urethral and anal sphincters
pennate muscles
fascicles insert obliquely on a tendon (feather shaped)
ex. palmar interosseus, rectus femoris, and deltoid
indirect attachment to bone
tendons
collagen fibers of endo-, peri-, and epimysium continue into the tendon
tendon merges into periosteum of bone
very strong structural continuity from muscle to bone
aponeurosis - tendon is a broad, flat sheet
direct attachment to bone
little separation between muscle and bone
muscle seems to immerge directly from bone
margins of brachialis, lateral head of triceps brachii
origin
bony attachment at stationary end of muscle
insertion
bony attachment to mobile end of muscle
belly
thicker, middle region of muscle between origin and insertion
prime mover (agonist)
muscle that produces most of the force during a joint action
ex. brachialis - prime mover for elbow flexion
synergist
muscle that aids the prime mover
stabilizes the nearby joint
modifies the direction of movement
ex. biceps brachii - synergist for elbow flexion
antagonist
opposes the prime mover
relaxes to give prime mover control over an action
preventing excessive movement and injury
antagonistic pairs - muscles that act on opposing sides of a joint
ex. triceps brachii - antagonist for elbow flexion
fixator
muscle that prevents movement of bone
ex. rhomboideus holds scapula firmly in place
intrinsic muscles
contained within a region
both its origin and insertion occur there
extrinsic muscles
act on a designated region, but has origins elsewhere
ex. fingers
characteristics of muscles
responsiveness (excitability) - to chemical signals, stretch and electrical changes across the plasma membrane
conductivity - local electrical change triggers a wave of excitation that travels along the muscle fiber
contractility - shortens when stimulated
extensibility - capable of being stretched between contractions
elasticity - returns to its original resting length after being stretched
skeletal muscle
voluntary, striated muscle attached to one or more bones
made up of muscle cells called muscle fibers or myofibers
myofiber
composed of myofibrils
also known as muscle cell or muscle fiber
myofibril
long protein bundles that occupies the main portion of the interior of a muscle fiber
myofilaments
make up myofibrils
a protein microfilament responsible for muscle cell contraction
composed of myosin or actin proteins
sacrolemma
plasma membrane of a muscle fiber
sarcoplasm
cytoplasm of a muscle fiber
sarcoplasmic reticulum (SR)
smooth ER that forms a network around each myofibril - calcium reservoir
calcium activates the muscle contraction process
terminal cisternae
dilated end-sacs of SR which cross muscle fiber from one side to the other
T tubules
tubular infoldings of the sarcolemma which penetrate through the cell and emerge on the other side
Triad
a T tubule and two terminal cisterns
glycogen function in myofibers
stored in abundance to provide energy with heightened exercist
myoglobin functions in myofibers
red pigment that stores oxygen needed for muscle activity
Muscle growth and repair
myoblasts - stem cells fuse to form each muscle fiber
satellite cells - unspecialized myoblasts remaining between the muscle fiber and endomysium
may multiply and produce new muscle fibers to some degree
repair by fibrosis
3 kinds of myofilaments found in a myofibril
thick filaments - myosin proteins
thin filaments - primarily actin proteins
elastic filaments - titin (connectin) proteins
thick myofilaments
made of several hundred myosin molecules
shaped like a gold club
heads directed outward in a helical array around the bundle
bare zone has no heads in the middle
thin myofilaments
fibrous (F) actin - 2 intertwined strands
string of globular (G) actin subunits each with an active site that can bind to head of myosin molecule
tropomyosin - each blocking 6 or 7 active sites on G actin subunits
troponin complex - small, calcium-binding protein on each tropomyosin molecule
elastic myofilaments
titin (connectin) - huge springy protein
flank each thick filament and anchor it to the Z disc
help the cell recoil to its resting length (elasticity)
keeps thick and thin filaments aligned
prevents over stretching
regulatory proteins
tropomyosin and troponin
like a switch that starts and stops contraction
contraction activated by release of calcium into sarcoplasm and its binding to troponin
troponin changes shape and moves tropomyosin off the active sites on actin
contractile proteins
myosin and actin
do the work of contracting the muscle
accessory proteins
at least 7 other accessory proteins in or associated with thick or thin filaments
anchor the myofilaments
regulate length of myofilaments
alignment of myofilaments for optimal effectiveness
dystrophin
links actin in outermost myofilaments to transmembrane proteins and eventually to fibrous endomysium surrounding the entire muscle cell
transfers forces of muscle contraction to connective tissue around myofiber
genetic defects in dystrophin produce muscular dystrophy
normal allele makes dystrophin
absence of it leads to torn cell membranes
Explain the effects of a mutation in Myostatin
removes the body's primary "brake" on muscle growth. This disruption causes significant skeletal muscle hyperplasia (increased number of muscle fibers) and hypertrophy (increased size of fibers), resulting in exceptional muscularity
Describe the arrangement of myofilaments in sarcomeres and how that results in
striations
A band - dark, anisotropic
part of A band where thick and thin filaments overlap is especially dark
H band - in middle of A band - just thick filaments
M line - middle of H band
I band - alternating lighter band - I stands for isotropic
Z disc - provides anchorage for thin filaments and elastic filaments
bisects I band at edge of sarcomere
sarcomere
the segment of the myofibril from one z disc to the next
functional contractile unit of the muscle fiber
muscle shortens because individual sarcomeres shorten
denervation atrophy
shrinkage of paralyzed muscle when connection not restored
somatic motor neuron
stimulate skeletal muscle
cell bodies are located in the brainstem and spinal cord
somatic motor fibers
axons of somatic motor neurons
lead to the skeletal muscle
each nerve fiber branches out to a number of muscle fibers
200 myofibers on avg are controlled by a single somatic motor neuron
each myofiber is supplied by only one motor neuron
motor unit
one nerve fiber and all the muscle fibers innervated by it
myofibers of one motor unit
dispersed throughout the muscle
contract in unison
produce weak contraction over wide area
provides ability to sustain long-term contraction as motor units take turns contracting
effective contraction requires the contraction of several motor units at once
number of muscle fibers per neuron in an average, small and large motor units
average motor unit - 200 muscle fibers per neuron
small motor unit - 3-6 muscle fibers per neuron
fine degree of control
eye and hand muscles
large motor units - 1000 myofibers per neuron
ex. gastrocnemius
powerful contractions supplied by large motor units
neuromuscular junction
when target cell is a muscle fiber
each terminal branch of the nerve fiber within the NMJ forms separate synapse with the muscle fiber
one nerve fiber stimulates the muscle fiber at several points within the NMJ
synapse
point where a nerve fiber meets its target cell
Components of neuromuscular junction
Synaptic knob - swollen end of nerve fiber
contains synaptic vesicles filled with acetylcholine
synaptic vesicles undergo exocytosis to release ACh into synaptic cleft
synaptic cleft - tiny gap between synaptic knob and muscle sarcolemma
Schwann cell - envelopes and isolates all of the NMJ from surrounding tissue fluid
50 million ACh receptors - proteins incorporated into muscle cell plasma membrane
Basal lamina - think layer of collagen and glycoprotein separates Schwann cell and entire muscle cell from surrounding tissue
contains acetylcholinesterase (AChe) that breaks down ACh after contraction
How can neuromuscular toxins disrupt muscle contraction?
interfere with synaptic function
some pesticides contain cholinesterase inhibitors
bind to AChe and prevent it from degrading ACh
spastic paralysis - a state of continual contraction of the muscles
tetanus (lockjaw) is a form of spastic paralysis caused by toxin Clostridium tetani
glycine in spinal cord normally stops motor neurons from producing unwanted muscle contractions
tetanus toxin blocks glycine release in the spinal cord and causes overstimulation and spastic paralysis of the muscles
flaccid paralysis
a state in which the muscles are limp and cannot contract
compete with ACh for receptor sites, but do not stimulate the muscles
What makes muscle fibers and neurons electrically excitable cells?
their plasma membrane exhibits voltage changes in response to stimulation
Unstimulated (resting) cell
there are more anions (negative ions) on the inside of the plasma membrane than on the outside
plasma membrane is electrically polarized (charged)
there are excess sodium (Na+) ions in the ECF
there are excess potassium (K+) ions in the ICF
Resting membrane potential of a myofiber is -90 mV
Voltage (electrical) potential
a difference in electrical charge from one point to another
Stimulated (active) muscle fiber or nerve cell
ion gates open in the plasma membrane
Na+ instantly diffuses down its concentration gradient into the cell
these cations override the negative charges in the ICF
depolarized occurs as inside of plasma membrane becomes briefly positive
Na+ gates close and K+ gates open
K+ rushes out of cell
repolarization as loss of positive K+ ions turns membrane negative again
action potential
quick up and down voltage shift from the negative RMP to a positive value, and back to the negative value again
a quickly fluctuating voltage seen in an active stimulated cell
an action potential at one point on a plasma membrane causes another one to happen immediately in front of it, which triggers another one a little farther along and so forth
4 Major Phases of Contraction and Relaxation
excitation
the process in which nerve action potentials lead to muscle action potentials
excitation-contraction coupling
events that link the action potentials on the sarcolemma to activation of the myofilaments, thereby preparing them to contract
contraction
step in which muscle fiber develops tension and may shorten
relaxation
when work is done, a muscle relaxes and returns to its resting length
Order of muscle contraction and relaxation
nerve fiber releases ACh
ACh binds to receptors on the muscle fiber, triggering an action potential
Calcium is released from the SR of the muscle fiber
Calcium binds to troponin
Tropomyosin moves to expose active sites on the thin filament
Myosin binds to actin and pulls creating a power stroke that shortens the sarcomere
muscle contracts
AChe breaks down ACh
Calcium is removed from troponin and returns to the SR
Muscle relaxes
rigor mortis
hardening of muscles and stiffening of body beginning 3-4 hours after death
deteriorating SR releases calcium
deteriorating sarcolemma allows calcium to enter cytosol
calcium activates myosin-actin cross-bridging
muscle contracts, but cannot relax
Muscle relaxation requires ATP, but NO ATP after death
Rigor mortis peaks at 12 hours AD, then diminishes over next 48-60 hours
Length-tension relationship
the amount of tension generated by a muscle and the force of contraction depends on how stretched or contracted it was BEFORE being stimulated
overly contracted at rest - a weak contraction occurs
too stretched before stimulated - a weak contraction occurs
optimum resting length
produces greatest force when muscle contracts
CNS continually monitors and adjusts length of the resting muscle
maintains a state of partial contraction - muscle tone (tonus)
threshold
the minimum voltage necessary to generate an action potential in the muscle fiber and produce a contraction
Twitch
a quick cycle of contraction when stimulus is at threshold or higher
phases of a twitch contraction:
latent period
contraction phase
relaxation phase
latent period of twitch contraction
2 msec delay between the onset of stimulus and onset of twitch response
time required for excitation, excitation-contraction coupling and tensing of elastic components of the muscle
internal tension - force generated during latent period and no shortening of muscle occurs
contraction phase of twitch contraction
phase in which filaments slide and the muscle shortens
once elastic components are taut, muscle begins to produce external tension - in muscle that moves a load
short-lived phase
relaxation phase of twitch contraction
SR quickly reabsorbs calcium, myosin releases the thin filaments and tension declines
muscle returns to resting length
entire twitch lasts from 7-100 msec
factors affecting twitch strength
stimulus frequency
stimuli arriving closer together produce stronger twitches
concentration of calcium in sarcoplasm
how stretched the muscle was before it was stimulated
temperature of the muscles
warmed up muscle contracts more strongly
enzymes work more quickly
lower than normal pH of sarcoplasm weakens the contraction
state of hydration of muscle
affects overlap of thick and thin filaments
What happens when you stimulate the nerve with higher and higher voltages?
it produces stronger contractions
higher voltages excite more and more nerve fibers in the motor nerve which stimulates more and more motor units to contract
recruitment or multiple motor unit (MMU) summation
the process of bringing more motor units into play
What happens when stimulus intensity (voltage) remains constant but stimulus
frequency varies?
Up to 10 stimuli per second:
each stimulus produces identical twitches and full recovery between twitches
10-20 stimuli per second:
produces treppe (staircase) phenomenon
muscle still recovers fully between twitches, but each twitch develops more tension than the one before
stimuli arrive so rapidly that SR doesn’t have time between stimuli to completely absorb all calcium released
Calcium conc. in cytosol rises higher and higher with each stimulus causing subsequent twitches to be stronger
heat released by each twitch causes muscle enzymes like myosin ATPase to work more efficiently and produce stronger twitches as muscle warms up
20-40 stimuli per second:
produces incomplete tetanus
each stimulus arrives before the previous twitch is over
new twitch rides piggy back on previous one, generating higher tension
temporal summation - 2 stimuli arrive close together
wave summation - results from one wave of contraction added to another
each twitch reaches a higher level of tension than the one before
muscle relaxes only partially between stimuli
produces a state of sustained fluttering contraction (incomplete tetanus)
40-50 stimuli per second:
muscle has no time to relax between stimuli
twitches fuse to a smooth, prolonged contraction called complete tetanus
muscle in complete tetanus produces 4x the tension as a single twitch
isometric muscle contraction
muscle is producing internal tension which an external resistance causes it to stay the same length or become longer
can be a prelude to movement when tension is absorbed by elastic component of muscle
important in postural muscle function and antagonistic muscle joint stabilization
isotonic muscle contraction
muscle changes in length with no change in tension
concentric contraction
muscle shortens while maintains tension
a type of isotonic contraction
eccentric contraction
muscle lengthens as it maintains tension
a type of eccentric contraction
2 main pathways of ATP synthesis
anaerobic fermentation
enables cells to produce ATP in absence of oxygen
yields little ATP
produces lactic acid, a major factor in muscle fatigue
aerobic respiration
produces far more ATP
less toxic end produces (CO2 and water)
requires a continual supply of oxygen
What 2 enzyme systems control phosphate transfers for immediate energy needs?
myokinase - transfers Pi from one ADP to another converting the later to ATP
creatine kinase - obtains Pi from a phosphate-storage molecule creatine phosphate (CP)
fast-acting system that helps maintain the ATP level while other ATP-generating mechanisms are being activated
phosphagen system
ATP and CP collectively
provides nearly all energy used for short bursts of intense activity
one minute of brisk walking
6 seconds of sprinting or fast swimming
important in activities requiring brief, but maximum effort
football, baseball, and weight lifting
short term energy needs
glycogen-lactic acid system
the pathway from glycogen to lactic acid
produces enough ATP for 30-40 seconds of maximum activity
long-term energy needs
aerobic respiration
theoretically produces 36-38 ATP per glucose
efficient means of meeting the ATP demands of prolonged exercise
one’s rate of oxygen consumption rises from 2-3 min and levels off to a steady state in which aerobic ATP production keeps pace with demand
little lactic acid accumulates under steady state conditions
causes of muscle fatigue
ATP synthesis declines as glycogen is consumed
ATP shortage slows down Na+K+ pumps
compromises ability to maintain the resting membrane potential and excitability of the muscle fibers
lactic acid lowers pH of sarcoplasm
inhibits enzymes involved in contraction, ATP synthesis, and other aspects of muscle function
release of K+ with each action potential causes the accumulation of extracellular K+
hyperpolarizes the cell and makes the muscle fiber less excitable
motor nerve fibers use up their ACh
CNS fatigues by unknown processes
endurance
the ability to maintain high-intensity exercise for more than 4-5 minutes
determined in large part by one’s maximum oxygen uptake (VO2max)
maximum oxygen uptake (VO2max)
the point at which the rate of oxygen consumption reaches a plateau and does not increase further with an added workload
proportional to body size
peaks at around age 20
usually greater in males than females
can be twice as great in trained endurance athletes as in untrained person
results in twice the ATP production
oxygen debt
heavy breathing continues after strenuous exercise
excess post-exercise oxygen consumption (EPOC) - difference between the resting rate of oxygen consumption and the elevated rate following exercise
typically about 11 liters extra is needed after strenuous exercise
repaying the oxygen debt
oxygen debt is needed to:
replace oxygen reserves that were depleted in the 1st minute of exercise
replenish the phosphagen system
oxidize lactic acid
serve the elevated metabolic rate
oral creatine supplement
increases level of creatine phosphate in muscle tissue and increases speed of ATP regeneration
useful in burst type exercises like weight-lifting
risks are not well known
muscle cramping, electrolyte imbalances, dehydration, water retention, gastro issues, weight gain, seizures, strokes
kidney disease from overloading kidney with metabolite creatine
body can stop producing natural creatine
carbohydrate loading
dietary regimen
packs extra glycogen into muscles cells
extra glycogen is hydrophilic and adds 2.7 g water/g glycogen
slow oxidative (SO), slow twitch, red, or type I fibers
dark meat
abundant mitochondria, myoglobin, and capillaries - deep red color
adapted for aerobic respiration and fatigue resistance
relative long twitch lasting about 100 msec
soleus of calf and postural muscles of the back
fast glycolytic (FG), fast-twitch, white, or type II fibers
white meat
fibers are well adapted for quick responses, but not for fatigue resistance
rich in enzymes of phosphagen and glycogen-lactic acid systems
less mitochondria, myoglobin, and blood capillaries which gives paler appearance
SR releases and reabsorbs Ca quickly, so contractions are quicker
extrinsic eye muscles, gastrocnemius, and biceps brachii
factors affecting muscle strength
muscle size
fascicle arrangement
pennate stronger than parallel, and parallel stronger than circular
size of motor units
larger the motor unit, the stronger the contraction
multiple motor unit summation = recruitment
when stronger contraction is required, the nervous system activates more motor units
temporal summation
nerve impulses usually arrive at a muscle in a series of closely spaced action potentials
the greater the stimulation frequency, the more strongly a muscle contracts
length-tension relationship
fatigue
resistance training (weight lifting)
contraction of a muscle against a load that resist movement
a few minutes of resistance exercise a few times a week is enough to stimulate muscle growth
growth is from cellular enlargement
muscle fibers synthesize more myofilaments and myofibrils and grow thicker
endurance training (aerobic exercise)
improves fatigue resistant muscles
slow twitch fibers produce more mitochondria, glycogen, and acquire a greater density of blood capillaries
improves skeletal strength
increases the red blood cell count and oxygen transport capacity of the blood
enhances the function of the cardiovascular, respiratory, and nervous systems
cardiac muscle
limited to the heart where it functions to pump blood
required properties of cardiac muscle
contraction with regular rhythm
muscle cells of each chamber must contract in unison
contractions must last long enough to expel blood
must work in sleep or wakefulness, without fail and without conscious attention
must be highly resistant to fatigue
striated
intercalated discs joins myocytes
electrical gap junctions allow each myocyte to directly stimulate its neighbors
SR less developed, but T tubules are larger
damaged cardiac muscle cells repair by fibrosis
very slow twitches
smooth muscle
composed of myocytes with fusiform shape
one nucleus located near the middle of cell
no visible striations
z discs are absent and replaced by dense bodies
cytoplasm has extensive cytoskeleton of intermediate filaments
No T tubules
capable of mitosis and hyperplasia
some lack nerve supply, others receive autonomic fibers, not somatic motor fibers as in skeletal muscle
2 types of smooth muscle
multiunit smooth muscle
occurs in some of the largest arteries and pulmonary air passages, in piloerector muscle of hair follicle, and in iris of eye
autonomic innervation similar to skeletal muscle
terminal branches of a nerve fiber synapse with individual myocytes and form a motor unit
each motor unit contracts independently of the others
single unit smooth muscle
more widespread
found in most blood vessels, digestive, respiratory, urinary, and reproductive tracts (visceral muscle)
2 layers
gap junction
large number of cells contract together as a single unit
stimuli of smooth muscle
chemical stimuli
hormones, carbon dioxide, low pH, oxygen deficiency
in response to stretch
single unit smooth muscle in stomach and intestines has pacemaker cells that set off waves of contraction throughout entire layer of muscle
varicosities
in a single unit smooth muscle, each autonomic nerve fibers have up to 20,000 beadlike swellings known as varicosities
each contains synaptic vesicles and a few mitochondria
nerve fiber passes amid several myocytes and stimulates all of them at once when it releases its neurotransmitter
contraction and relaxation of smooth muscle
contraction and relaxation are very slow compared to skeletal muscle
latent period is 50-100 msec compared to skeletal muscle’s 2 msec
tension peaks at about 500 msec
declines over a period of 1-2 seconds
slows myosin ATPase enzyme and slow pumps that remove Ca+2
Ca+2 bonds to calmodulin instead of troponin
activates kinases and ATPases that hydrolyze ATP