MUSCULAR SYSTEM: HISTOLOGY AND PHYSIOLOGY

Ref: Seeley’s Anatomy and Physiology, 11th Ed.

Prepared by: Pauline Necor

CONTENTS a. TYPES OF MUSCLE TISSUE

1. Functions of the Muscular System 1 ● Skeletal muscle
a. Types of Muscle Tissue ○ With its associated connective tissue,
2. General Properties of Muscle Tissue 1 constitutes about 40% of the body’s weight
3. Skeletal Muscle Anatomy 1 ○ Responsible for locomotion, facial
a. Whole Skeletal Muscle Anatomy expressions, posture, respiration, and many
b. Skeletal Muscle Fiber Anatomy other body movements
4. Skeletal Muscle Fiber Physiology 6 ○ The nervous system voluntarily controls the
a. Excitability of Muscle Fibers skeletal muscles
b. Action Potentials ● Smooth muscles
c. Excitation-Contraction Coupling ○ Most widely distributed type of muscle in the
d. Cross-Bridge Movement body
e. Muscle Relaxation ○ Found in walls of hollow organs (stomach,
5. Whole Skeletal Muscle Physiology 9 uterus) and tubes (blood vessels and certain
a. Muscle Twitch glands)
b. Muscle Contraction Force ○ Find in the iris of the eye for pupil dilation
6. Muscle Fiber Types 10 ● Cardiac muscle
7. Energy Sources for Muscle Contraction 10 ○ Found only in the heart
a. Oxygen Deficit ○ Contraction moves blood throughout the body
b. Fatigue ○ Contracts spontaneously and rhythmically
8. Muscle Hypertrophy 11 ○ Has autorhythmicity
9. Smooth Muscle 11 ○ Controlled by the autonomic nervous system
10. Cardiac Muscle 12 (ANS) and endocrine system
11. Effects of Aging on Skeletal Muscle 12
2. GENERAL PROPERTIES OF MUSCLE TISSUE
● Contractility
1. FUNCTIONS OF THE MUSCULAR SYSTEM
○ Ability of the muscle to shorten forcefully, or
● Major functions of all three types of muscles: contract
1. Movement of the body ○ Muscle shortening (concentric) is forceful
a. Most skeletal muscles are attached to bone ○ Muscle lengthening (eccentric) is passive
and are responsible for majority of body ■ Muscle lengthens to oppose
movements contraction
2. Maintenance of posture ● Excitability
a. Skeletal muscles constantly maintain tone ○ Capacity of the muscle to respond to an
3. Respiration electrical stimulus
a. Skeletal muscles of the thorax carry out ● Extensibility
breathing movements ○ A muscle can be stretched beyond its normal
4. Production of body heat resting length and still be able to contract
a. Heat is given off as a by-product when skeletal ● Elasticity
muscles contract ○ Ability of the muscle to spring back to its
5. Communication original resting length after being stretched
a. Skeletal muscles are involved in all aspects of
3. SKELETAL MUSCLE ANATOMY
speaking
6. Constriction of organs and vessels a. WHOLE SKELETAL MUSCLE ANATOMY
a. Smooth muscles contract within the walls of ● Each skeletal muscle is a complete organ consisting of
internal organs and vessels, which causes cells, called skeletal muscle fibers
those structures to constrict ● Associated with smaller amounts of connective tissue,
7. Contraction of the heart blood vessels, and nerves
a. The contraction of the cardiac muscle causes ● The connective tissue fibers that surround a muscle
the heart to beat and its internal components extend beyond the center
of the muscle to become tendons, which connect
muscles to bones or to the dermis of the skin

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 capillary beds surrounding the muscle fibers
SKELETAL MUSCLE FIBER ANATOMY
● Each muscle cell is called a muscle fiber
○ Long, cylindrical cells with multiple nuclei
located near the plasma membrane
○ A single fiber can extend the entire length of a
muscle
○ 1 millimeter (mm) to about 4 centimeters (cm)
in length
○ 10 micrometers (um) to 100 um in diameter
○ alternating
○ Light and dark bands give the muscle fiber
striations
Muscle Fiber Development
● Muscle fibers develop from less mature, multinucleated
cells called myoblasts
● The multiple nuclei found in skeletal muscle fibers result
from the fusion of myoblast precursor cells, not from the
Connective Tissue Coverings division of nuclei within myoblasts.
● Fascicles ● Myoblasts differentiate into functional muscle fibers as
○ Numerous visible bundles that composes a contractile proteins accumulate within their cytoplasm
muscle ● Nerves begin to innervate the developing muscle fibers.
○ Surrounded by a connective tissue layers ● Hypertrophy
called the perimysium ○ Increase in the size of each muscle fiber, not
● Epimysium from a substantial increase in the number of
○ A layer of dense irregular collagenous muscle fibers
connective tissue that surrounds the entire Histology of Muscle Fibers
muscle
● Two main aspects of muscle contraction
● Fascia
○ Electrical component
○ Sheets of dense irregular collagenous
■ Sarcolemma
connective tissue within the body
■ Transverse tubules
● Muscular fascia
■ Sarcoplasmic reticulum
○ Located superficial to the epimysium
○ Mechanical component
○ Separates individual muscles or groups of
■ Myofibrils
muscles
■ Myofilaments
Nerves and Blood Vessels ● Sarcolemma
● Motor neurons ○ Plasma membrane of a muscle fiber
○ Nerve cells responsible for stimulating skeletal ○ Has two delicate connective tissue layers
muscle contraction located outside
○ Originates from the brain and spinal cord and ■ External lamina and endomysium
extends to the skeletal muscle fibers through ● External Lamina
nerves ○ Deep and thinner to the endomysium
○ At the fascicles, the axons of motor neurons ○ Consists of reticular (collagen) fibers
branch repeatedly, each branch projecting ● Endomysium
toward the center of one muscle fiber. ○ Second layer
○ Every skeletal muscle fiber in the body is in ○ Thicker layer that consists of reticular fibers
contact with a branch of a motor neuron. ● Transverse tubules or T tubules
■ Contact points are called synapses ○ Many tubelike invaginations located along the
or neuromuscular junctions surface of the sarcolemma
○ Each motor neuron innervates more than one ○ Occur at regular intervals along the muscle
muscle fiber fiber and extend inward, connecting the
○ More than one motor neuron innervates most extracellular environment with the interior of
whole muscles the muscle fiber
● Numerous branches of the arteries supply the extensive ○ Lie adjacent to the highly organized smooth

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 endoplasmic reticulum Sarcomeres
● Sarcoplasmic reticulum
○ Highly organized smooth endoplasmic
reticulum
● Sarcoplasm
○ Muscle cell cytoplasm
○ contains numerous bundles of protein
filaments called myofibrils
● Myofibrils
○ threadlike structure, approximately 1–3 um in ● Basic structural and functional unit of skeletal muscle
diameter, that extends the length of the muscle ● The smallest portion of skeletal muscle capable of
fiber contracting
○ contains two kinds of long, thin and thick ● Z disks
protein filaments, myofilaments ○ Separate one sarcomere from the next
● Actin myofilaments ○ Filamentous network of protein that forms a
○ Thin myofilaments stationary anchor for the attachment of actin
● Myosin myofilaments myofilaments
○ Thick myofilaments ○ Each sarcomere extends from one Z disk to
● Actin and myosin myofilaments are arranged into highly the next Z disk
ordered units called sarcomeres ○ Holds actin myofilaments in place
○ Join end to end to form the myofibrils ● The arrangement of the actin and myosin myofilaments
gives the myofibril a banded, or striated, appearance
when viewed longitudinally
○ Two I-bands separated by an A-band
● I bands
○ Light-staining bands
○ Isotropic
○ Each includes a Z disk and extends to the
ends of the myosin myofilaments
○ consists only of actin myofilaments
● A bands
○ Dark-staining band in the center of each
sarcomere
○ Anisotropic
○ Each extends the length of the myosin
myofilament within a sarcomere
○ Actin and myosin myofilaments overlap for
some distance at both ends of the A band
○ In the center of each A band is a smaller band,
called the H zone
■ Actin and myosin myofilaments do
not overlap
■ Only myosin myofilaments are
present
● M line
○ Dark line in the middle of the H zone
○ Consists of delicate filaments that attach to the
center of the myosin myofilaments
○ Helps hold the myosin myofilaments in place
● Titin
○ Largest known proteins
○ Attaches to Z disks and extends along myosin
myofilaments to the M line
○ Part of the titin molecule in the I band
functions as a spring, allowing the sarcomere
to stretch and recoil

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 heads that extend laterally
○ Four light myosin chains are attached to the
heads of each myosin molecule
● The myosin heads have three important properties:
1. The heads can bind to active sites on the actin
molecules to form cross-bridges
2. The heads are attached to the rod portion by a
hinge region that can bend and straighten
during contraction
3. The heads are ATPase enzymes, which break
down adenosine triphosphate (ATP), releasing
energy

Neuromuscular Junction Structure

● Each muscle fiber is in contact with a branch of a motor
neuron axon from the brain or spinal cord that carry
Actin and Myosin Myofilament Structure electrical signals called action potentials
● Each actin myofilament is composed of two strands of ● Action potentials
fibrous actin (F actin) ○ Stimulate muscle fiber action potentials
○ a series of tropomyosin molecules, and a followed by muscle contraction
series of troponin molecules ● Neuromuscular junction
● The two strands of F actin are coiled to form a double ○ AKA Synapse
helix, which extends the length of the actin myofilament ○ Point of contact of motor neuron axon
● Each F actin strand is a polymer of approximately 200 branches with the muscle fiber
small, globular units called globular actin (G actin) ○ Consists of a group of enlarged axon terminals
monomers that rests in an invagination of the sarcolemma
● Tropomyosin is an elongated protein that winds along ○ Consists of the axon terminals and the area of
the groove of the F actin double helix the muscle fiber sarcolemma they innervate
● Troponin is composed of three subunits: ○ Each axon terminal is the presynaptic
1. binds to actin terminal
2. binds to tropomyosin ■ Each presynaptic terminal contains
3. binding site for Ca2+ numerous mitochondria and many
● Myosin myofilaments are composed of many elongated small, spherical sacs
myosin molecules shaped like golf clubs ○ Synaptic cleft
○ Consists of two myosin heavy chains wound ■ Space between the presynaptic
together to form a rod portion lying parallel to terminal and the muscle fiber
the myosin myofilament and two myosin

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○ Postsynaptic Membrane Sliding Filament Model
■ AKA motor end-plate ● When a muscle contracts, the actin and myosin
■ Muscle plasma membrane in the area myofilaments in the sarcomere slide past one another
of the junction but remain the same length as when the muscle is at
○ Synaptic vesicles rest
■ contain acetylcholine (ACh) ● When the myofilaments slide past each other and the
● a neurotransmitter sarcomeres shorten, the myofibrils also shorten
● a substance released from a because the myofibrils consist of sarcomeres joined
presynaptic membrane that end to end
diffuses across the synaptic ● The myofibrils extend the length of the muscle fibers,
cleft and alters the activity of and when they shorten the muscle fibers shorten
the postsynaptic cell ● Groups of muscle fibers make up a muscle fasciculus,
● Neurotransmitters can and several muscle fascicles make up a whole muscle
stimulate (or inhibit) the ● When sarcomeres shorten, myofibrils, muscle fibers,
production of an action muscle fascicles, and muscles shorten to produce
potential in the postsynaptic muscle contraction
membrane (the ● During muscle relaxation, the sarcomeres lengthen
sarcolemma) by binding to
ligand-gated ion channels

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 4. SKELETAL MUSCLE FIBER PHYSIOLOGY
a. EXCITABILITY OF MUSCLE FIBERS
● Action potentials travel from the brain or spinal cord
along the axons to muscle fibers and cause them to
contract
● Electrically excitable cells are polarized
○ The inside of the plasma membrane is
negatively charged
● Resting membrane potential
○ Charge difference across the plasma
membrane of an unstimulated cell
Ion Channel
● Ligand-gated ion channels
○ Opens when a ligand (a chemical signal such
as neurotransmitter) binds to a receptor that is
part of the ion channel
○ The axons of neurons supplying skeletal
muscle fibers release a neurotransmitter,
which binds to ligand-gated Na+ channels in
the membranes of the muscle fibers.
○ As a result, the Na+ channels open, allowing
Na+ to enter the cell
● Voltage-gated ion channels
○ Open and close in response to a particular b. ACTION POTENTIALS
membrane potential
● Reverse of the resting membrane potential
○ When a neuron or muscle fiber is stimulated,
● The inside of the plasma membrane becomes
the charge difference changes, and a
positively charged compared with the outside
particular charge causes certain voltage-gated
● Ion channels open, changing the characteristics of the
ion channels to open or close
plasma membrane permeability
○ Voltage-gated Na+, K+, and Ca2+ channels
● The diffusion of ions changes the charge across the
The Resting Membrane Potential plasma membrane and produces an action potential
● Action potentials cannot be produced without a resting ● Action potential
membrane potential ○ Lasts 1 millisecs to a few millisecs
● The RMP is the result of three factors: ○ Has two phases:
1. The concentration of K+ inside the plasma ■ Depolarization
membrane is higher than outside the plasma ■ Repolarization
membrane
2. The concentration of Na+ outside the plasma
membrane is higher than inside the plasma
membrane
3. The plasma membrane is more permeable to
K+ than to Na+
● Since excitable cells have many K+ leak ion channels,
at rest, K+ moves out of the cell faster than Na+ moves
into the cell
● It is the active transport of Na+ and K+ by the
sodium-potassium pump that maintains the uneven
distribution of Na+ and K+ across the plasma
membrane
● In a resting cell, the sodium-potassium pump transports K+
from outside the cell to the inside and transports Na+ from
inside the cell to the outside
● The potential differences across the plasma membranes of
neurons and muscle fibers are between −70 and −90 mV

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 difference across the plasma membrane. An action
potential, which is a reversal of that charge difference,
stimulates cells to respond by contracting.

● Threshold
○ The depolarization changes the membrane
potential to a value
○ Action potential is triggered
● Depolarization phase
○ A brief period during which further
depolarization occurs and the inside of the cell
becomes positively charged
● Repolarization phase
○ The return of the membrane potential to its
resting value
● Action potentials occur according to the all-or-none
principle
○ All action potentials are identical
● The resting membrane potential results from a charge

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 c. EXCITATION-CONTRACTION COUPLING
● Action potentials produced in the sarcolemma of a
skeletal muscle fiber can lead to contraction of the fiber.
● Excitation-contraction coupling
○ Mechanism of an action potential that causes
muscle contraction
○ Occurs due to interactions of T tubules and
sarcoplasmic reticulum
■ Ca2+ in the sarcoplasmic reticulum
binds to the troponin component of
actin myofilaments
■ T tubules project into the muscle fiber
and wrap around sarcomeres in the
region where actin and myosin
myofilaments overlap
■ Sarcoplasmic reticulum is enlarged to
form terminal cisternae near the T
tubules
● Triad
○ T tubules
○ 2 adjacent terminal cisternae

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 d. CROSS-BRIDGE MOVEMENT
● Cross-bridge cycling
1. Exposure of active sites
a. When Ca2+ binds to troponin, it moves the
tropomyosin exposing the active sites
2. Cross-bridge formation
a. When active sites are exposed, the myosin
head attach to it using 1 phosphate
3. Power stroke
a. Myosin head moves using the stored energy
(ADP), which causes the actin myofilament to
slide past the myosin myofilament
4. Cross-bridge release
a. ATP molecule binds to myosin head causing it
to detach from the active site
5. Hydrolysis (breakdown of ATP)
a. ATP is split into ADP and P e. MUSCLE RELAXATION
6. Recovery stroke ● Occurs when ACh is not longer released at the
a. Myosin head returns to its resting position neuromuscular junction
● Three major ATP dependent events are required for
muscle relaxation:
1. After an action potential has occurred in the
muscle fiber, the sodium-potassium pump
must actively transport Na+ and K+ to return to
and maintain resting membrane potential.
2. ATP is required to detach the myosin heads
from the actin and return them to their resting
position.
3. ATP is needed for the active transport of Ca2+
into the sarcoplasmic reticulum.
5. WHOLE SKELETAL MUSCLE PHYSIOLOGY
a. MUSCLE TWITCH
● A muscle twitch is a single contraction of a muscle fiber
in response to a stimulus
● 3 phases:
○ Lag phase
■ Latent phase
■ Beginning of contraction
■ Gap between the time of stimulus
application to the motor neuron
○ Contraction phase
○ Relaxation phase

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 b. MUSCLE CONTRACTION FORCE 7. ENERGY SOURCES FOR MUSCLE CONTRACTION
● The strength of muscle contraction varies from weak to ● Adenylate kinase
strong ○ Or myokinase
● Muscle contraction forces is increased in two ways: ○ Transfer 1 phosphate from 1 ADP to a second
○ Summation ADP, resulting to 1 ATP and 1 AMP
■ An increase in stimulus frequency ● Conversion of a molecule called creatine phosphate to
increases the overall force of ATP
contraction ● During exercise periods of rest, when there is excess
■ Tetanus is a sustained contraction ATP, the excess ATP is used to store creatine
that occurs when the frequency of phosphate
stimulation is so rapid that no ● During exercise, especially at the onset of exercise, the
relaxation occurs small cellular ATP reserve is quickly depleted
○ Recruitment ● Anaerobic production of ATP
■ More motor units are stimulated, ○ Breaks down glucose to produce ATP and
which increase the total number of lactate
muscle fibers contracting ○ Produces less ATP but can produce ATP in a
matter of a few seconds
● Aerobic production of ATP
○ Produces more ATP, but slower to produce
● Conversion of two ADP to one ATP and one AMP
(adenosine monophosphate) during heavy exercise
○ If the use of ATP is greater than the production
of ATP, the ATP:ADP ratio decreases, which
interferes with the functioning of all major
ATP-dependent structures in the muscle fibers
■ An enzyme transfers one phosphate
from one ADP to another ADP,
generating one ATP and one AMP
■ The presence of AMP triggers a
switch from anaerobic respiration to
aerobic respiration of blood glucose
and fatty acids

6. MUSCLE FIBER TYPES

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 ○ Local inflammatory reaction
■ The presence of immune system
intermediates increases the
perception of pain, which most likely
serves as a signal to protect those
tissues from further damage
8. MUSCLE HYPERTROPHY
● Increase in size
○ Increase in number of…
■ Myofibrils
■ Sarcomeres
■ Nuclei from satellite cells
■ Mitochondria
■ Blood vessels
■ Connective tissues
● Training the muscles
○ Better neuromuscular coordination
○ Improved metabolism
○ Increased circulation
○ More efficient respiration
○ Greater capacity for heart to pump blood

a. OXYGEN DEFICIT 9. SMOOTH MUSCLE

● After increased level of activity, breathing pattern does ● Smooth muscle cells are spindle-shaped with a single
not readily return to pre-exercise level nucleus.
○ ATP is used to repay the oxygen deficit ● Have actin myofilaments and myosin myofilaments but
○ The recovery oxygen consumption is the are not striated.
amount of O2 needed in chemical reaction that ● The sarcoplasmic reticulum is poorly developed, and
occur to restore pre-exercise conditions caveolae may function as a T tubule system.
■ Conversion of lactate to glucose ● Calcium ions enter the cell to initiate contraction;
■ Replenishment of the depleted ATP calmodulin binds to Ca2+ and activates an enzyme that
and creatine phosphate stores in transfers a phosphate group from ATP to myosin. When
muscle fibers phosphate groups are attached to myosin,
■ Replenishment of O2 stores in the cross-bridges form.
lungs, blood, and muscles ● Relaxation results when myosin phosphatase removes
a phosphate group from the myosin molecule.
b. FATIGUE
○ If phosphate is removed while the
● The decrease capacity to do work and the reduced cross-bridges are attached, relaxation occurs
efficiency of performance that normally follows a period very slowly, and this is referred to as the latch
of activity state.
● Multiple mechanisms underlying fatigue: ○ If phosphate is removed while the
○ Acidosis of ATP depletion due to either cross-bridges are not attached, relaxation
increased ATP consumption or decreased occurs rapidly.
production ● Types:
■ Decreased effectiveness of Ca2+ on ○ Visceral smooth muscle fibers
actin and overall less Ca2+ release ■ contract slowly, have gap junctions
from the sarcoplasmic reticulum (and thus function as a single unit),
○ Oxidative stress ■ can be autorhythmic.
■ Build up of excess reactive oxygen ○ Multiunit smooth muscle fibers
species (ROS) ■ contract rapidly in response to
● Causes the breakdown of stimulation by neurons and function
proteins, lipids, or nucleic independently.
acids ● Functional properties
● Triggers interleukin (IL)-6, a ○ Can contract auto rhythmically in response to
mediator of inflammation stretch or when stimulated by the autonomic

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 nervous system or hormones. Smooth muscle contraction
○ Maintains a steady tension for long periods.
○ The force of smooth muscle contraction
remains nearly constant, despite changes in
muscle length.
○ Does not develop an oxygen deficit.

10. CARDIAC MUSCLE

● Striated
● Have a single nucleus,
● Are connected by intercalated disks (and thus function
as a single unit)
● Capable of autorhythmicity
11. EFFECTS OF AGING ON SKELETAL MUSCLE
● Reduction of muscle mass
● Slower response time for muscle contraction
● Reduction in stamina
● Increased recovery time
● Fast-twitch muscle fibers decrease in number more
rapidly than slow-twitch fibers
● Fewer action potentials are produced in muscle fibers
● The number of motor neurons decrease
● Decrease in the density of capillaries in skeletal
muscles

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