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MUSCLE TISSUE
OJORA K. A
Introduction 2
Muscles are multicellular contractile units of contractile cells.
Cells containing contractile proteins.
Structural biology of these proteins generates the forces
necessary for cellular contraction, which drives movement
within certain organs and the body as a whole
On the basis of morphological and functional characteristics
1. Skeletal muscles.
2. Cardiac muscles.
3. Smooth muscle.
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Skeletal muscle
Bundles of very long, cylindrical, multinucleated
cells that show cross-striations.
Contraction is quick, forceful, and usually under
voluntary control.
Itis caused by the interaction of thin actin filaments and
thick myosin filaments whose molecular configuration
allows them to slide upon one another.
Cardiac muscle
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has cross-striations and is composed of elongated,
branched individual cells that lie parallel to each other.
At sites of end-to-end contact are the intercalated
disks, structures found only in cardiac muscle.
Contraction of cardiac muscle is involuntary, vigorous,
and rhythmic.
Smooth muscle
consists of collections of fusiform cells that do not show
cross-striations.
Their contraction process is slow and not subject to
voluntary control.
PLEASE NOTE 5
Some muscle cell organelles have names that
differ from their counterparts in other cells.
Thecytoplasm of muscle cells (excluding the
myofibrils) is called sarcoplasm (Gr. sarkos, flesh,
+ plasma, thing formed), and the smooth
endoplasmic reticulum is called sarcoplasmic
reticulum.
The sarcolemma (sarkos + Gr. lemma, husk) is
the cell membrane, or plasmalemma.
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Skeletal Muscle
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Consists of muscle fibers, bundles of very long
(up to 30 cm) cylindrical multinucleated cells.
Multinucleation results from the fusion of
embryonic mononucleated myoblasts (muscle
cell precursors).
The oval nuclei are usually found at the
periphery of the cell under the cell
membrane.
MEDICAL APPLICATION 8
The variation in diameter of skeletal muscle fibers depends on factors
such as the specific muscle and the age and sex, state of
nutrition, and physical training of the individual.
It is a common observation that exercise enlarges the musculature and
decreases fat depots.
The increase in muscle thus obtained is caused by formation of new
myofibrils and a pronounced growth in the diameter of individual
muscle fibers (hypertrophy).
Hyperplasia does not occur in either skeletal or cardiac muscle but
does take place in smooth muscle.
Hyperplasia is rather frequent in organs such as the uterus, where both
hyperplasia and hypertrophy occur during pregnancy.
Organization of Skeletal Muscle 9
Epimysium- dense connective
perimysium- dense connective
Endomysium- basal lamina
and reticular fibers.
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Blood vessels penetrate the muscle within the connective tissue septa and form a rich capillary
network that runs between and parallel to the muscle fibers.
The capillaries are of the continuous type, and lymphatic vessels are found in the connective
tissue.
Myotendinous junction 12
The electron microscope shows that in this transitional region, collagen fibers of the
tendon insert themselves into complex infoldings of the plasmalemma of the muscle
fibers.
Organization of Skeletal Muscle 13
Fibers
With LM , longitudinally sectioned muscle fibers show cross-striations
of alternating light and dark bands.
The darker bands →A bands (anisotropic, ie, are birefringent in polarized
light).
The lighter bands → I bands (isotropic, ie, do not alter polarized light).
In the EM, each I band is bisected by a dark transverse line, the Z line.
The smallest repetitive subunit of the contractile apparatus, the
sarcomere, extends from Z line to Z line.
The sarcoplasm is filled with long cylindrical filamentous bundles called
myofibrils.
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Longitudinal section of skeletal muscle fibers. Note the dark-stained A bands and the light-stained I
bands, which are crossed by Z lines. Giemsa stain. High magnification.
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The Sarcomere 16
The smallest repetitive subunit of the contractile apparatus,
the sarcomere, extends from Z line to Z line.
EMreveal that this sarcomere pattern is due mainly to the
presence of two types of filaments thick and thin.
The thick filaments occupy the A band.
The thin filaments run between and parallel to the thick
filaments and have one end attached to the Z line.
TheI bands consist of the portions of the thin filaments that
do not overlap the thick filaments.
The
A bands are composed mainly of thick filaments in
addition to portions of overlapping thin filaments.
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The Sarcomere 18
A band shows the presence of a lighter zone in its
center, the H band, that corresponds to a region
consisting only of the rodlike portions of the myosin
molecule.
Bisectingthe H band is the M line, a region at which
lateral connections are made between adjacent thick
filaments.
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Striated muscle proteins; 21
Thefour main proteins are actin, tropomyosin,
troponin, and myosin.
Thin
filaments are composed of actin, tropomyosin,
troponin.
Thick filaments consist primarily of myosin.
The conducting system for contractile stimuli 22
The depolarization of the sarcoplasmic reticulum membrane,
which results in the release of Ca2+ ions, is initiated at a
specialized myoneural junction on the surface of the
muscle cell.
Surface-initiated depolarization signals would have to diffuse
throughout the cell to effect the release of Ca2+ from internal
sarcoplasmic reticulum cisternae.
To provide for a uniform contraction, skeletal muscle
possesses a system of transverse (T) tubules.
The conducting system for contractile stimuli 23
These fingerlike invaginations of the sarcolemma form a
complex anastomosing network of tubules that encircles
the boundaries of the A-I bands of each sarcomere in
every myofibril.
Adjacent to opposite sides of each T tubule are expanded
terminal cisternae of the sarcoplasmic reticulum.
This specialized complex, consisting of a T tubule with two
lateral portions of sarcoplasmic reticulum, is known as the
triad.
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The conducting system for contractile stimuli 25
At the triad, depolarization of the sarcolemma-derived T tubules is transmitted
to the sarcoplasmic reticulum membrane.
Muscle contraction depends on the availability of Ca2+ ions, and muscle relaxation is
related to an absence of Ca2+.
The sarcoplasmic reticulum specifically regulates calcium flow.
The sarcoplasmic reticulum system consists of a branching network of smooth
endoplasmic reticulum cisternae surrounding each myofibril.
After a neurally mediated depolarization of the sarcoplasmic reticulum membrane, Ca2+
ions concentrated within the sarcoplasmic reticulum cisternae are passively released
into the vicinity of the overlapping thick and thin filaments, whereupon they bind to
troponin and allow bridging between actin and myosin.
When the membrane depolarization ends, the sarcoplasmic reticulum acts as a calcium
sink and actively transports the Ca2+ back into the cisternae, resulting in the cessation of
contractile activity.
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The conducting system for contractile
stimuli
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Electron micrograph of skeletal muscle of a tadpole.
The sliding filament theory 29
Innervation
At the site of innervation, the nerve forms a dilated termination 30
that sits within a trough on the muscle cell surface.
This structure is called the motor end plate, or myoneural
junction.
At this site of the axon terminal are numerous mitochondria
and synaptic vesicles, the latter containing the
neurotransmitter acetylcholine.
The synaptic cleft.
At the junction, the sarcolemma is thrown into numerous deep
junctional folds.
In the sarcoplasm below the folds lie several nuclei and
numerous mitochondria, ribosomes, and glycogen
granules.
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Motor End Plate
Innervation 32
When an action potential invades the motor end plate,
acetylcholine is liberated from the axon terminal, diffuses
through the cleft, and binds to acetylcholine receptors in
the sarcolemma of the junctional folds.
Binding of the transmitter makes the sarcolemma more
permeable to sodium, which results in membrane
depolarization.
Excess acetylcholine is hydrolyzed by the enzyme
cholinesterase bound to the synaptic cleft basal lamina.
Acetylcholine breakdown is necessary to avoid prolonged
contact of the transmitter with receptors present in the
sarcolemma.
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Motor End Plate
MEDICAL APPLICATION 34
Myasthenia gravis is an autoimmune disorder
characterized by progressive muscular weakness
caused by a reduction in the number of
functionally active acetylcholine receptors in
the sarcolemma of the myoneural junction.
Thisreduction is caused by circulating antibodies
that bind to the acetylcholine receptors in the
junctional folds and inhibit normal nerve muscle
communication.
Muscle Spindles 35
Muscle spindles are encapsulated
proprioceptors of striated muscles .
These structures consist of a connective tissue
capsule housig the intrafusal fibers.
Several sensory nerve fibers penetrate the
muscle spindles, where they detect changes
in the length (distention) of extrafusal
muscle fibers and relay this information to the
spinal cord.
Here, reflexes of varying complexity are
activated to maintain posture and to regulate
the activity of opposing muscle groups involved
in motor activities such as walking.
Golgi Tendon Organs 36
In tendons, near the insertion sites of muscle
fibers, a connective tissue sheath
encapsulates several large bundles of
collagen fibers that are continuous with the
collagen fibers that make up the
myotendinous junction.
Sensory nerves penetrate the connective
tissue capsule.
These structures, known as Golgi tendon
organs, contribute to proprioception by
detecting tensional differences in
tendons.
Cardiac Muscle 37
Mature cardiac muscle exhibit a cross-striated banding pattern
identical to that of skeletal muscle.
Each cardiac muscle cell possesses only one or two centrally
located pale-staining nuclei.
Surrounding the muscle cells is a delicate sheath of endomysial
connective tissue containing a rich capillary network.
A unique and distinguishing characteristic of cardiac muscle is the
presence of dark-staining transverse lines that cross the
chains of cardiac cells at irregular intervals.
Intercalated disc 38
Junctional complexes found at the interface between adjacent cardiac muscle cells.
Appear as straight lines or may exhibit a steplike pattern.
It has two regions a transverse portion,and a lateral portion.
There are three main junctional specializations within the disk.
1. Zonulae adherentes which serve as anchoring sites for actin filaments of the
terminal sarcomeres.
2. Maculae adherentes (desmosomes) bind the cardiac cells together, preventing them
from pulling apart under constant contractile activity.
3. Gap junctions provide ionic continuity between adjacent cells.
The significance of ionic coupling is that chains of individual cells act as a syncytium,
allowing the signal to contract to pass in a wave from cell to cell.
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Drawing of a section of heart muscle, showing central nuclei, cross-striation, and intercalated
disks.
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D desmosome
E endothelial cell EL
external lamina F fibroblast
FA fascia adherens G
glycogen ID interdigitation L
leucocyte M mitochondria
N nexus or gap junction S
sarcomere SR sarcoplasmic
reticulum T T tubule
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Cardiac / skeletal muscles
CM fibres have an arrangement of contractile proteins similar to that of SM and
are consequently striated .
The T tubules are more numerous and larger in ventricular muscle than in
skeletal muscle.
Cardiac T tubules are found at the level of the Z band rather than at the A I junction
in SM.
The SR is not as well developed and wanders irregularly through the myofilaments.
Triads are not common in cardiac cells, because the T tubules are generally associated
with only one lateral expansion of sarcoplasmic reticulum cisternae.
Thus, heart muscle characteristically possesses diads composed of one T tubule and
one sarcoplasmic reticulum cisterna.
Cardiac muscle cells contain numerous mitochondria, which occupy 40% or more of the
cytoplasmic volume, reflecting the need for continuous aerobic metabolism in heart
muscle.
Smooth Muscle 44
Smooth muscle is composed of elongated, nonstriated cells,
each of which is enclosed by a basal lamina and a network of
reticular fibers .
The basal lamina and a network of reticular fibers serve to
combine the force generated by each smooth muscle fiber into a
concerted action, eg, peristalsis in the intestine.
cells are fusiform.
cell has a single nucleus located in the center of the broadest part
of the cell.
The borders of the cell become scalloped when smooth muscle
contracts, and the nucleus becomes folded or has the appearance
of a corkscrew.
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A rudimentary sarcoplasmic reticulum is present; it consists of a
closed system of membranes, similar to the sarcoplasmic 46
reticulum of striated muscle.
The characteristic contractile activity of smooth muscle is
related to the structure and organization of its actin and myosin
filaments.
These bundles consist of thin filaments containing actin and
tropomyosin and thick filaments consisting of myosin.
Both structural and biochemical studies reveal that smooth
muscle actin and myosin contract by a sliding filament
mechanism similar to what occurs in striated muscles.
Smooth muscle filaments. 47
Desmin (skeletin)- major protein of intermediate
filaments in all smooth muscles, and vimentin is an
additional component in vascular smooth muscle.
Smooth muscle is innervated by both sympathetic
and parasympathetic nerves of the autonomic
system.
Elaborateneuromuscular junctions such as those in
skeletal muscle are not present in smooth muscle.
Frequently, autonomic nerve axons terminate in a
series of dilatations in the endomysial connective
tissue.
Regeneration of Muscle Tissue 48
The three types of adult muscle have different potentials for regeneration
after injury.
Cardiac muscle
no regenerative capacity beyond early childhood.
Defects or damage (eg, infarcts) in heart muscle are generally replaced by the
proliferation of connective tissue, forming myocardial scars.
skeletal muscle
undergo limited regeneration.
The source of regenerating cells is believed to be the satellite cells.
They are considered to be inactive myoblasts that persist after muscle
differentiation.
Smooth muscle
an active regenerative response.
After injury, viable mononucleated smooth muscle cells and pericytes from
blood vessels undergo mitosis and provide for the replacement of the
damaged tissue.
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