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MUSCLE PHYSIOLOGY

INTRODUCTION
• Muscle cells, like neurons, can be excited
chemically, electrically, and mechanically to
produce an action potential that is transmitted
along their cell membrane.
• They have a contractile mechanism that is
activated by the action potential.
• The contractile proteins actin and myosin are
abundant in muscle, where they bring about
contraction.
• The actin-binding protein myosin and actin make
up one of the molecular motors that converts the
energy of ATP hydrolysis into movement of one
cellular component along another.
TYPES OF MUSCLE
• Muscle is generally divided into three types, skeletal,
cardiac, and smooth.

• Skeletal muscle makes up the great mass of the


somatic musculature. It has
• - well-developed cross-striations,
• -does not normally contract in the absence of nervous
stimulation,
• - lacks anatomic and functional connections between
individual muscle fibers, and
• -is generally under voluntary control.

TYPES OF MUSCLE
• Cardiac muscle
• -also has cross-striations, but
• -it is functionally syncytial and contracts rhythmically
in the absence of external innervation owing to the
presence in the myocardium of pacemaker cells that
discharge spontaneously.
• Smooth muscle
• -lacks cross-striations.
• -The type found in most hollow viscera is functionally
syncytial and contains pacemakers that discharge
irregularly.
• -The type found in the eye and in some other locations
is not spontaneously active and resembles skeletal
muscle.
Skeletal muscle
STRUCTURE OF SKELETAL MUSCLE.
Made up of bundles of muscle cells known as muscle
fibers or myofibers – long, cylindrical, multinucleated,
extending the entire length of the muscle and
attaching to tendons at both ends.
Striated when viewed under light microscope.
Striations (stripes) are produced by alternating dark
and light bands that span the width of the fiber.
A (anisotropic) band – dark bands
I (isotropic) band – light bands
Under electron microscope, thin dark lines can be
seen in the middle of the I band – called Z line or disk.
Cytoplasm of myofiber is known as
sarcoplasm.
Rich in mitochondria.
Contains 2 types of myofilaments
(myofibrils)
-(a) thick filaments made up of myosin II
--(b) thin filaments made up of actin,
troponin and tropomyosin.
-Other proteins that maintain the
architecture and provide elasticity to
There are 3 functional types of skeletal
muscle proteins
(a)Contractile proteins: Myosin and Actin
(b) Regulatory proteins: Tropomyosin and
Troponin
(c)Attachment proteins: Titin, nubulin and
dystrophin.
-Dystrophin provides support for the muscle
fiber by bridging the cytoskeleton and
myofibrils in the fiber with the extracellular
matrix.
- When dystrophin is defective muscle
ORGANIZATION OF MEMBRANES OF MUSCLE CELL
Cell membrane of muscle fiber (sarcolemma) has
inward extensions (Transverse or T tubules) which
conduct action potentials to the contractile filaments.
In skeletal muscles, the T tubules contain voltage-
sensitive dihydropyridine (DHP) receptors. Located at
junction of A and I bands.
Receptors open the Ca++ channels in the sarcoplasmic
reticulum when the T tubule is depolarized.
In cardiac muscles, the T system is located at the
region of the Z lines and the DHP receptor is a voltage-
gated Ca++ which lets in Ca++ from ECF when the tube
is depolarized.
The sarcoplasmic reticulum
(endoplasmic reticulum) stores large
amounts of Ca++ which are released by
the arrival of action potentials.
The transverse cistern is formed at area
of contact between sarcoplasmic
reticulum and T tubule. Contains the
ryanodine receptor (Calcium channels
specific to the transverse cistern).
MECHANISM OF CONTRACTION
The Sliding Filament Theory: Proposed by Huxley
in 1957.
Theory states that muscle contraction results from
sliding of thin filaments inwards between the thick
filaments.
- The length of the A band is unchanged, I.e.
there is no change in the length of the thick filaments.
- the distance between the Z line and the
adjacent edge of the H zone also remain unchanged
- The H zone and the I band decrease in width,
i.e. there is an increase in the overlap between the
thick and thin filaments.
MOLECULAR BASIS OF MUSCLE CONTRACTION:
EXCITATION-CONTRACTION COUPLING.
Based on the premise that the sliding filaments
arises from an interaction between the myosin
cross-bridges and the actin filaments.
Requires the presence of Ca++ and utilizes
energy from ATP.
• Spread of AP throughout the T system
triggers the voltage-sensitive DHP receptors to
open Ca++ channels. This triggers release of Ca+
+
thus raising [Ca++ ] in sarcoplasm.
• Ca++ binds to troponin C, causing conformational
change in the troponin which is transmitted to
tropomyosin.
• The shape change in troponin causes its attached
tropomyosin to shift position in the actin filament thus
exposing binding sites for the myosin cross bridges.
• Myosin cross bridges previously activated by the
hydrolysis of ATP, attach to actin.
• After attachment, pulls thin filaments over thick
filaments (power stroke).
• Attachment of fresh ATP allows the cross bridges to
detach from actin and repeat the contraction cycle as
long as Ca++ remains attached to troponin.
•When AP stops, Ca++ becomes actively
transported into sarcoplasmic reticulum
by Ca++ -ATPase pumps and tropomyosin
returns to its inhibitory position
(relaxation).
• (If ATP is not available, the detachment
will not occur leading to sustained
contracture (rigor mortis) as in death).
Sequence of events in
contraction of skeletal muscle
• Steps in contraction1
• -(1) Discharge of motor neuron.
• - (2) Release of transmitter (acetylcholine) at
motor end-plate.
• -(3) Binding of acetylcholine to nicotinic
acetylcholine receptors.
• -(4) Increased Na+ and K+ conductance in end-
plate membrane.
• -(5) Generation of end-plate potential.
• - (6) Generation of action potential in muscle
fibers..
Sequence of events in
contraction of skeletal muscle
• -(7) Inward spread of depolarization along T
tubules.
• -(8) Release of Ca2+ from terminal cisterns of
sarcoplasmic reticulum and diffusion to thick
and thin filaments.
• -(9) Binding of Ca2+ to troponin C, uncovering
myosin-binding sites on actin.
• - (10) Formation of cross-linkages between actin
and myosin and sliding of thin on thick
filaments, producing shortening
Sequence of events in
relaxation of skeletal muscle.
• Steps in relaxation
• -(1) Ca2+ pumped back into
sarcoplasmic reticulum.
• -(2) Release of Ca2+ from troponin.
• -(3) Cessation of interaction between
actin and myosin
SOURCES OF ENERGY
(1)Phosphocreatinine: A store of high energy
phosphate bond. Reaction is catalysed by creatine
kinase.
Phosphocreatinie + ADP ------------- ATP + Creatinine
(2) Action of myokinase in ADP to form ATP
2 ADP ----------------- ATP + AMP
(3) Metabolism of glucose, fats and proteins to
generate ATP. Breakdown of glucose or glycogen
to pyruvic acid (anaerobic glycolysis) is most rapid
source of ATP. Over 95% of all energy used by the
muscle for sustained, long term contraction is
derived from this source.
Types of Contraction

• Muscular contraction involves shortening of the contractile elements, but


because muscles have elastic and viscous elements in series with the
contractile mechanism,
• - it is possible for contraction to occur without an appreciable decrease in
the length of the whole muscle .Such a contraction is called isometric
("same measure" or length).
• -Contraction against a constant load, with approximation of the ends of
the muscle, is isotonic ("same tension").
• -Note that since work is the product of force times distance, isotonic
contractions do work whereas isometric contractions do not. In other
situations, muscle can do negative work while lengthening against a
constant weight.
Fiber Types

• Although skeletal muscle fibers resemble one another


in a general way, skeletal muscle is a very
heterogeneous tissue made up of fibers that vary in
myosin ATPase activity, contractile speed, and other
properties.
• The fibers fall roughly into two types, type I and type
II, although each of these types is itself a spectrum.
Muscles containing many type I fibers are called red
muscles because they are darker than other muscles.
• The red muscles, which respond slowly and have a long
latency, are adapted for long, slow, posture-
maintaining contractions.
• The long muscles of the back are red muscles
Fiber Types

• . White muscles, which contain


mostly type II fibers, have short
twitch durations and are specialized
for fine, skilled movement.
• The extraocular muscles and some of
the hand muscles contain many type
II fibers and are generally classified
as white muscles.
Disease of Muscle

• Myasthenia gravis is due to blockade or


destruction of Ach receptors by antibodies
secreted by the immune system of the affected
person. Results in muscle weakness.
• Tetanus toxin and botulinum toxin are bacterial
products that cause paralysis by preventing
neurotransmission.
• Tetanus toxin blocks inhibitory synapses
causing spastic paralysis.
• Botulinum toxin causes flaccid paralysis by
preventing release of Ach.
CARDIAC MUSCLE
MORPHOLOGY
• The striations in cardiac muscle are similar to those
in skeletal muscle, and Z lines are present.
• The muscle fibers branch and interdigitate, but each
is a complete unit surrounded by a cell membrane.
• Where the end of one muscle fiber abuts on
another, the membranes of both fibers parallel each
other through an extensive series of folds. These
areas, which always occur at Z lines, are called
intercalated disks .
• They provide a strong union between fibers,
maintaining cell-to-cell cohesion, so that the pull of
one contractile unit can be transmitted along its
axis to the next.
MORPHOLOGY
• Along the sides of the muscle fibers next to the
disks, the cell membranes of adjacent fibers fuse for
considerable distances, forming gap junctions.
• These junctions provide low-resistance bridges for
the spread of excitation from one fiber to another .
• They permit cardiac muscle to function as if it were
a syncytium, even though there are no protoplasmic
bridges between cells.
• The T system in cardiac muscle is located at the Z
lines rather than at the A-I junction, where it is
located in mammalian skeletal muscle.
ELECTRICAL PROPERTIES
• The resting membrane potential of individual
mammalian cardiac muscle cells is about -90 mV
(interior negative to exterior).
• Stimulation produces a propagated action potential
that is responsible for initiating contraction.
• Ca++ enters cytoplasm from sarcoplasmic reticulum
and extracellular fluid.
• Can contract without nerve stimulation; action
potentials originate in pacemaker cells of heart.
• Gap junctions present as intercalated discs.
PACEMAKER TISSUE

• The heart continues to beat after all nerves to it


are sectioned; indeed, if the heart is cut into
pieces, the pieces continue to beat.
• This is because of the presence in the heart of
specialized pacemaker tissue that can initiate
repetitive action potentials.
• The pacemaker tissue makes up the conduction
system that normally spreads impulses
throughout the heart. It is characterized by an
unstable membrane potential that slowly
decreases after each impulse until the firing level
is reached and another impulse is generated
SMOOTH MUSCLE
morphology
• Smooth muscle is distinguished anatomically
from skeletal and cardiac muscle because it
lacks visible cross-striations.
• Actin and myosin-II are present, and they
slide on each other to produce contraction.
• However, they are not arranged in regular
arrays, as in skeletal and cardiac muscle, and
so the striations are absent.
• Instead of Z lines, there are dense bodies in
the cytoplasm and attached to the cell
membrane, and these are bound by α-actinin
to actin filaments.
morphology
• Smooth muscle also contains tropomyosin, but
troponin appears to be absent.
• The isoforms of actin and myosin differ from
those in skeletal muscle.
• There is a sarcoplasmic reticulum, but it is
poorly developed.
• In general, smooth muscles contain few
mitochondria and depend to a large extent on
glycolysis for their metabolic needs.
types
• There is considerable variation in the
structure and function of smooth muscle in
different parts of the body.
• In general, smooth muscle can be divided
into visceral smooth muscle and multi-
unit smooth muscle.
• Visceral smooth muscle occurs in large
sheets, has low-resistance bridges
between individual muscle cells, and
functions in a syncytial fashion.
types
• The bridges, like those in cardiac muscle, are junctions
where the membranes of the two adjacent cells fuse to
form gap junctions.
• Visceral smooth muscle is found primarily in the walls of
hollow viscera, e.g. the musculature of the intestine, the
uterus, and the ureters.
• Multi-unit smooth muscle is made up of individual units
without interconnecting bridges.
• It is found in structures such as the iris of the eye.
SUMMARY AND
COMPARISON
SUMMARY AND
COMPARISON
• SKELETAL MUSCLE
• Striated; actin and myosin arranged in sarcomeres.
• Well-developed sarcoplasmic reticulum and T tubules.
• Contains troponin in thin filaments.
• Ca++ released into cytoplasm from sarcoplasmic
reticulum.
• Cannot contract without nerve stimulation; denervation
results in muscle atrophy.
• Muscle fibers stimulated independently; no gap junction.
SUMMARY AND
COMPARISON
• CARDIAC MUSCLE
• Striated; actin and myosin arranged in sarcomeres.
• Moderately developed sarcoplasmic reticulum and T
tubules.
• Contains troponin in the thin filaments.
• Ca++ enters cytoplasm from sarcoplasmic reticulum
and extracellular fluid.
• Can contract without nerve stimulation; action
potentials originate in pacemaker cells of heart.
• Gap junctions present as intercalated discs.
SUMMARY AND
COMPARISON
• SMOOTH MUSCLE
• Not striated; more actin than myosin; actin inserts into
dense bodies and membrane.
• Poorly developed sarcoplasmic reticulum, no T tubule.
• Contains calmodulin, a protein that when bound to Ca++ ,
activates the enzyme myosin light-chain kinase (MLCK).
• Ca++ enters cytoplasm from extracellular fluid, sarcoplasmic
reticulum and perhaps mitochondria.
• Maintains tone in absence of nerve stimulation, visceral
smooth muscle produces pacemaker potential; denervation
results in hypersensitivity to stimulation.
• Gap junctions generally present.
• Multiple-Choice Questions

Self Study
In questions 1-5, select the single best answer.

1. The action potential of skeletal muscle

(A) has a prolonged plateau phase

(B) spreads inward to all parts of the muscle via the T tubules

(C) causes the immediate uptake of Ca2+ into the lateral sacs of the sarcoplasmic reticulum

(D) is longer than the action potential of cardiac muscle

(E) is not essential for contraction

2. The functions of tropomyosin in skeletal muscle include

(A) sliding on actin to produce shortening

(B) releasing Ca2+ after initiation of contraction

(C) binding to myosin during contraction

(D) acting as a "relaxing protein" at rest by covering up the sites where myosin binds to actin

(E) generating ATP, which it passes to the contractile mechanism


Self Study
• 3. The cross-bridges of the sarcomere in skeletal muscle are made up of

(A) actin

(B) myosin

(C) troponin

(D) tropomyosin

(E) myelin

4. The contractile response in skeletal muscle

(A) starts after the action potential is over

(B) does not last as long as the action potential

(C) produces more tension when the muscle contracts isometrically than when the muscle contracts
isotonically

(D) produces more work when the muscle contracts isometrically than when the muscle contracts
isotonically

(E) decreases in magnitude with repeated stimulation


Self Study
• 5. Gap junctions
(A) are absent in cardiac muscle
(B) are present but of little functional importance in
cardiac muscle
(C) are present and provide the pathway for rapid
spread of excitation from one cardiac muscle fiber to
another
(D) are absent in smooth muscle
(E) connect the sarcotubular system to individual
skeletal muscle cells
Self Study
• 1. Compare the electrical and mechanical
events in smooth and cardiac muscle with
those in skeletal muscle, and compare
their responses to acetylcholine and
norepinephrine
• 2. "Muscle is a machine for converting
chemical into mechanical energy."
Analyze and discuss this statement.
Prepared By;

Dr. Patrick Maison

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