Muscle Tissue

Yosef Tadesse

1
Outline
• Introduction
• Functional Characteristics of muscle tissue
• Skeletal muscle
– Function
– Development
– Organization
– Muscle Fibers

2
 – Innervation
– Muscle Spindles & Tendon Organs
– Types of Skeletal Muscle Fiber
• Cardiac Muscle
• Smooth Muscle
• Regeneration of Muscle Tissue

3
Introduction
• Muscle tissue is composed of cells
differentiated for optimal use of the universal
cell property termed contractility
• Microfilaments and associated proteins
together generate the forces necessary for
cellular contraction, which drives movement
within certain organs and the body as a whole
• Nearly all muscle cells are of mesodermal
origin and they differentiate mainly by a
gradual process of cell lengthening with
simultaneous synthesis of myofibrillar proteins

4
Functional Characteristics of muscle tissue

• Excitability or irritability
– the ability to respond to a stimulus
• Contractility
– the ability to shorten
• Extensibility
– the ability to stretch
• Elasticity
– the ability to resume normal length after
being stretched or shortened
5
• Muscle cells are elongated and are referred to
as muscle fibers
• Prefixes of myo or mys and sarco refer to
muscle
• Some muscle cell organelles have names that
differ from their counterparts in other cells
– The cytoplasm of muscle cells is called
sarcoplasm
– The smooth ER is called sarcoplasmic
reticulum
– The sarcolemma is the cell membrane, or
plasmalemma
6
• There are three types of muscle tissue
based on
– morphologic and functional characteristics
• The structure of each type is adapted to
its physiologic role
• These muscle tissues differ in
– the structure of their cells
– their body location
– their function
– the means by which they are activated to
contract
7
• Skeletal muscle
– Composed of bundles
of very long,
cylindrical,
multinucleated cells
that show cross-
striations
– Their contraction is
quick, forceful, and
under voluntary
control

8
• Cardiac muscle
– Has cross-striations
– 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

9
• Smooth muscle
– Consists of
collections of
fusiform cells that
do not show
striations
– Their contraction
process is slow and
not subject to
voluntary control 

10
Skeletal muscle
• Wrapped by connective tissue into organs called
muscles
• Attach to bones and some to skin
• When contracts it causes gross body movements
• Skeletal muscle consists of muscle fibers, which are
long, cylindrical multinucleated cells with diameters of
10–100 µm
– Multinucleation results from the fusion of embryonic
mesenchymal cells called myoblasts
• The long oval nuclei are usually found at the periphery
of the cell under the cell membrane
• Striated (fibers contain alternating light and dark
bands)
• Voluntary controlled 11
Skeletal Muscle: Function
• Produces movement
– Movement results from skeletal muscle contraction
– Responsible for locomotion and manipulation
– Allows interaction with external environment
• Maintains posture
– Skeletal muscles are utilized constantly to maintain
sitting, standing, and moving postures
– Postural muscle develop to compensate for the
never ending pull of gravity

12
• Stabilizing Joints
– Skeletal muscle provide the dynamic stability of
joints
– Many joints are poorly reinforced by ligaments and
connective tissue
• Generating Heat
– Muscles generate heat as they contract
– The heat generated is important to maintain normal
body temperature
– Skeletal muscle generates most of the heat because
it represents 40% of body mass

13
 Development of skeletal muscle
• Skeletal muscle begins to differentiate when mesenchymal
cells called myoblasts align and fuse together to make longer,
multinucleated tubes called myotubes
• Myotubes synthesize the proteins to make up myofilaments
and gradually begin to show cross striations
• Myotubes continue differentiating to form functional
myofilaments and the nuclei are displaced against the
sarcolemma
• Part of the myoblast population does not fuse and
differentiate, but remains as a group of mesenchymal cells
called muscle satellite cells located on the external surface
of muscle fibers inside the developing external lamina
– Satellite cells proliferate and produce new muscle fibers
following muscle injury

14
15
MEDICAL APPLICATION
• 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
• Exercise enlarges the musculature by formation
of new myofibrils and a pronounced growth in
the diameter of individual muscle fibers
• This process, characterized by increase of cell
volume, is called hypertrophy
• Tissue growth by an increase in the number of
cells is termed hyperplasia
16
Organization
• Each skeletal muscle is a discrete organ
with thousands of fibers
• Muscle fibers predominate the tissue but
it also contains blood vessels, nerve
fibers, and connective tissue
• 3 layers of connective tissue surround
the muscle fibers
– Epimysium: an external sheath of dense
connective tissue surrounding the entire
muscle

17
 – Perimysium:
• From the epimysium, thin septa of
connective tissue extend inward,
surrounding the fascicles or bundles of
fibers within a muscle
• The connective tissue around each
fascicle is called the perimysium
– Endomysium:
• Delicate connective tissue surround each
muscle fiber
• External to the epimysium is the deep fascia
that binds muscles into functional groups

18
• All the connective tissue layers are continuous
with one another as well as with the tendons
that join muscles to bone
– It transmit the mechanical forces generated by the
contracting muscle fibers because individual muscle
cells seldom extend from one end of a muscle to the
other
• Blood vessels penetrate the muscle within the
connective tissue septa and form a rich
capillary network in the endomysium
• Lymphatic vessels and larger blood vessels are
found in the other connective tissue layers 

19
20
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22
Muscle Fibers
• As observed with the light microscope,
longitudinally sectioned skeletal muscle fibers
show cross-striations of alternating light and
dark bands
• The darker bands are called A bands; the lighter
bands are called I bands
• In the EM each I band is seen to be bisected by a
dark transverse line, the Z line
• The repetitive functional subunit of the
contractile apparatus, the sarcomere, extends
from Z line to Z line and is about 2.5 µm long in
resting muscle
23
24
25
26
Myofibrils
• The sarcoplasm is filled primarily with long
cylindrical filamentous bundles called
myofibrils running parallel to the long axis of
the fiber
• The myofibrils have a diameter of 1–2 µm and
consist of an end-to-end repetitive
arrangement of sarcomeres
• The A and I banding pattern in sarcomeres is
due mainly to the regular arrangement of two
types of myofilaments—thick and thin—that
lie parallel to the long axis of the myofibrils in
a symmetric pattern 27
• Thick filaments
– 1.6 µm long and 15 nm wide
– occupy the A band, the central portion of the
sarcomere
• Thin filaments
– 1.0 µm long and 8 nm wide
– run between and parallel to the thick filaments
– have one end attached to the Z line
• As a result of this arrangement
– I bands consist of the portions of the thin
filaments that do not overlap the thick filaments
– A bands are composed mainly of thick filaments in
addition to overlapping portions of thin filaments
28
• H zone
– Close observation of the A band shows the
presence of a lighter zone in its center, the H
zone
– Corresponds to a region consisting only of the
rod-like portions of the myosin molecule of
thick filament with no thin filaments present
• M line
– Bisecting the H zone
– A region where lateral connections are made
between adjacent thick filaments

29
Each muscle fiber contains several parallel bundles called
myofibrils
Each myofibril consists of a long series of sarcomeres
which contain thick and thin filaments and are separated
from one another by Z discs 30
Thin filaments are actin filaments with one end bound to actinin, the major
protein of the Z disc. Thick filaments are bundles of myosin, which span the
entire A band and are bound to proteins of the M line and to the Z disc across
31
the I bands by a very large protein called titin, which has spring-like domains.
The molecular organization of the sarcomeres has bands of
greater and lesser protein density, resulting in staining
differences that produce the dark and light-staining bands
seen by light microscopy and TEM. 32
• Thin and thick filaments overlap for some
distance within the A band
• As a consequence, a cross section in the
region of filament overlap shows each
thick filament surrounded by six thin
filaments in the form of a hexagon
• Thin filaments are composed of F-actin,
associated with tropomyosin, and troponin
• Thick filaments consist primarily of
myosin
• Myosin and actin together represent 55%
of the total protein of striated muscle
33
• F-actin
– Consists of long filamentous polymers
containing two strands of globular (G-actin)
monomers, 5.6 nm in diameter, twisted
around each other in a double helical
formation
– Each G-actin monomer contains a binding site
for myosin
– Actin filaments are anchored perpendicularly
on the Z line by actin-binding protein -
actinin 

34
• Tropomyosin
– Each tropomyosin subunit is a long, thin
molecule about 40 nm in length containing
two polypeptide chains, which assembles to
form a long polymer located in the groove
between the two twisted actin strands
• Troponin
– a complex of three subunits:
• TnT, which attaches to tropomyosin
• TnC, which binds calcium ions
• TnI, which inhibits the actin-myosin interaction
– Troponin complexes are attached at specific
sites at regular intervals along each
tropomyosin molecule
35
Each thin filament is composed of F-actin,
tropomyosin, and troponin complexes
36
• Myosin
– A much larger complex (molecular mass ~500 kDa)
– Can be dissociated into two identical heavy chains
and two pairs of light chains
– Myosin heavy chains are thin, rod-like molecules
(150 nm long and 2–3 nm thick) made up of two
heavy chains twisted together as myosin tails
– Small globular projections at one end of each heavy
chain form the heads, which have ATP binding sites
as well as the enzymatic capacity to hydrolyze ATP
(ATPase activity) and the ability to bind actin
– The four light chains are associated with the head
– Several hundred myosin molecules are arranged
within each thick filament with their rodlike
portions overlapping and their globular heads
directed toward either end
37
Each thick filament consists of many myosin heavy
chain molecules bundled together along their rod-
like tails, with their heads exposed and directed
toward neighboring thin filaments
38
• Analysis of thin sections of striated
muscle shows the presence of cross
bridges between thin and thick filaments
– These bridges are formed by the head of the
myosin molecule plus a short part of its
rodlike portion
– These bridges are involved in the conversion
of chemical energy into mechanical energy 

39
Besides interacting with the neighboring thin
filaments, thick myofilament bundles are held in
place by myosin-binding proteins within the M line
40
Sarcoplasmic Reticulum & Transverse
Tubule System
• In muscle the smooth endoplasmic reticulum
(SER) is specialized for Ca2+ ion sequestration
• The depolarization of this sarcoplasmic
reticulum membrane, which results in the
release of Ca2+ ions, is initiated at the
specialized myoneural junction on the surface
of the muscle cell
• Surface-initiated depolarization signals would
have to diffuse throughout the cell to produce
Ca2+ release from internal sarcoplasmic
reticulum cisternae
• Diffusion of the depolarization signal would
lead to a wave of contraction of myofibrils
41
• The sarcoplasmic reticulum system
consists of a branching network of small
cisternae surrounding each myofibril
• To provide for a uniform contraction, skeletal
muscle fibers have a system of transverse (T)
tubules
– These fingerlike invaginations of the sarcolemma
form a complex network of tubules that encircles
every myofibril near the A-I band boundaries of
each sarcomere
• Adjacent to opposite sides of each T tubule are
expanded terminal cisternae of the
sarcoplasmic reticulum
• This specialized complex, consisting of a T
tubule and usually two small cisternae of
sarcoplasmic reticulum, is known as a triad
42
• 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, which is necessary
for rapid contraction and relaxation
cycles
43
• After a neurally mediated depolarization
of the sarcoplasmic reticulum membrane,
Ca2+ ions concentrated within these
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 molecules
• When the membrane depolarization ends,
the sarcoplasmic reticulum actively
transports the Ca2+ back into the
cisternae, ending contractile activity
44
45
T tubules 46
Arrows: triad
47
Innervation
• Myelinated motor nerves branch out within the
perimysium connective tissue, where each nerve
gives rise to several terminal twigs
• At the site of innervation, the axon loses its
myelin sheath and forms a dilated termination
situated within a trough on the muscle cell
surface
• This structure is called the motor end-plate,
or the neuromuscular junction
• At this site, the axon is covered only by a thin
cytoplasmic extension from a Schwann cell

48
• Within the axon terminal are numerous
mitochondria and synaptic vesicles, containing
the neurotransmitter acetylcholine
• Between the axon and the muscle is a space, the
synaptic cleft, in which lies an amorphous basal
lamina matrix from the muscle fiber
• At the junction, the sarcolemma is thrown into
numerous deep junctional folds, which provide
for greater surface area
• In the sarcoplasm below the folds lie several
nuclei and numerous mitochondria, ribosomes,
and glycogen granules

49
• When an action potential reaches the motor end
plate, acetylcholine is liberated from the axon
terminal, diffuses across the cleft, and binds to
acetylcholine receptors in the folded sarcolemma
• Binding of the transmitter opens Na+ channels in
the sarcolemma, producing 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 its
receptors
50
• The depolarization initiated at the motor
end-plate is propagated along the surface
of the muscle cell and deep into the
fibers via the transverse tubule system
• At each triad, the depolarization signal is
passed to the sarcoplasmic reticulum and
results in the release of Ca2+, which
initiates the contraction cycle
• When depolarization ceases, the Ca2+ is
actively transported back into the
sarcoplasmic reticulum cisternae, and the
muscle relaxes
51
• A single nerve fiber (axon) can innervate
one muscle fiber, or it may branch and be
responsible for innervating 160 or more
muscle fibers
• In the case of multiple innervation, a single
nerve fiber and all the muscle fibers it
innervates are called a motor unit
• Individual striated muscle fibers do not
show graded contraction—they contract
either all the way or not at all
• To vary the force of contraction, the
fibers within a muscle bundle do not all
contract at the same time
52
• Since muscles are composed of many motor
units, the firing of a single motor axon will
generate tension proportional to the number of
muscle fibers innervated by that axon
• Thus, the number of motor units and the variable
size of each unit can control the intensity of a
muscle contraction
• The ability of a muscle to perform delicate
movements depends on the size of its motor
units
– For example, because of the fine control required by
eye muscles, each of their fibers is innervated by a
different nerve fiber
• In larger muscles exhibiting coarser movements,
such as those of the limb, a single, profusely
branched axon innervates a motor unit that
consists of more than 100 individual muscle
fibers 53
54
Silver staining can reveal the nerve bundle (NB), the
terminal axonal twigs, and the motor end plates (MEP)
on striated muscle fibers (S). 55
A SEM shows the branching ends of a motor axon, each covered by an
extension of the last Schwann cell and expanded terminally as a motor
end plate embedded in a groove in the external lamina of the muscle
fiber. 56
57
MEDICAL APPLICATION
• Myasthenia gravis
– 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
– This reduction is caused by circulating antibodies
that bind to the acetylcholine receptors in the
junctional folds and inhibit normal nerve-muscle
communication
– As the body attempts to correct the condition,
membrane segments with affected receptors are
internalized, digested by lysosomes, and replaced by
newly formed receptors
– These receptors, however, are again made
unresponsive to acetylcholine by similar antibodies,
and the disease follows its progressive course 
58
Muscle Spindles & Tendon Organs
• Striated muscles and myotendinous
junctions contain sensory receptors that
are encapsulated proprioceptors
• Muscle spindles
– Stretch detectors among the muscle
fascicles
– Consist of a connective tissue capsule
surrounding a fluid-filled space that contains
a few thin, nonstriated muscle fibers densely
filled with nuclei and called intrafusal fibers
59
– These thin fibers differ from the ordinary
skeletal muscle fibers in having essentially no
myofibrils
– Their many nuclei can either be closely
aligned (nuclear chain fibers) or piled in a
central dilatation (nuclear bag fibers)
– Several sensory nerve axons penetrate each
muscle spindle and wrap around individual
intrafusal fibers
– Muscle spindles detect contraction of
neighboring (extrafusal) muscle fibers during
body movement and participate in the
nervous control of body posture and the
coordinate action of opposing muscles
60
• Golgi tendon organs
– In tendons, near the insertion sites of muscle fibers, a
connective tissue sheath encapsulates the large
collagen bundles of the myotendinous junction
– Sensory nerves penetrate this capsule and form
another sensory receptor known as (Golgi) tendon
organs
– Tendon organs detect changes in tension within
tendons produced by muscle contraction and act to
inhibit motor nerve activity if tension becomes
excessive
• Because both of these sensory receptors detect
increases in tension, they help to regulate the
amount of effort required to perform movements
that call for variable amounts of muscular force 
61
62
Types of Skeletal Muscle Fiber
• Skeletal muscle fibers are classified
into three types based on their
physiological, biochemical, and
histochemical characteristics
• All three fiber types are normally
found throughout most muscles
– red (slow twitch)
– white (fast twitch)
– intermediate fibers
63
Type I or Red oxidative (slow twitch) fibers
• Relatively thin fibers
• Named for the abundant myoglobin (oxygen binding
pigment) in their sarcoplasm
• Obtain their energy from aerobic (oxygen requiring)
reactions and thus have relatively large numbers of
mitochondria and a rich blood supply
• Adapted for slow, continuous contractions over
prolonged periods
• Resistant to fatigue as long as oxygen is present
• Used in many of the postural muscles of the axial
skeleton
• Because their fibers are thin, they do not generate
much power
64
Type IIb or (fast twitch), white
glycolytic fibers
• About twice the diameter of red fibers
• Have fewer mitochondria and capillaries and less
myoglobin, but abundant glycogen, making them
very pale in color
• Depend largely on anaerobic pathways (glycolysis)
for energy
• Adapted for rapid contractions, but fatigue quickly
• Contain more myofibrils and generate more power
• Common in the muscle of the upper limbs
• Used to lift heavy objects for brief periods

65
Type IIa or intermediate oxidative-
glycolytic fibers
• Intermediate between the other fiber types both in color,
size and in energy metabolism
• Like white fibers they contract quickly
• Like red fibers they are oxygen dependent
• Have many mitochondria, a rich supply of capillaries and much
myoglobin, but also have considerable glycogen
• They utilize both oxidative metabolism and anaerobic
glycolysis
• Less fatigue resistant but more powerful than red fibers, but
not as strong as white
• Abundant in the muscles of the lower limbs
• Used to move the body for long periods of time in activities
like walking and jogging

66
Cross-section of skeletal muscle stained histochemically to detect the
density of myofibrillar myosin-ATPase can be used to demonstrate the
distribution of slow (S) type I fibers, intermediate (I) type IIa fibers,
and fast (F) type IIb fibers.  67
Cardiac Muscle
• During embryonic development, the
mesoderm cells of the primitive heart
tube align into chainlike arrays
• Rather than fusing into multinucleated
cells, as in the development of skeletal
muscle fibers, cardiac muscle cells form
complex junctions between extended
processes
• Cells within a fiber often branch and bind
to cells in adjacent fibers
68
69
• Mature cardiac muscle cells are
approximately 15 µm in diameter and from
85 to 100 µm in length
• They exhibit a cross-striated banding
pattern comparable to that of skeletal
muscle
• Cardiac muscle cell possesses only one or
two centrally located pale-staining nuclei
• Surrounding the muscle cells is a delicate
sheath of endomysium containing a rich
capillary network
70
• Intercalated discs
– 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
– These intercalated discs represent the
interface between adjacent muscle cells
where many junctional complexes are present

71
– Transverse regions of these steplike discs have
many desmosomes and fascia adherentes (which
resemble the zonula adherentes between
epithelial cells) and together these serve to
bind cardiac cells firmly together to prevent
their pulling apart under constant contractile
activity
– The more longitudinal portions of each disc
have multiple gap junctions, which provide ionic
continuity between adjacent cells
– These act as "electrical synapses" and allow
cells of cardiac muscle to act as in a
multinucleated syncytium, with contraction
signals passing in a wave from cell to cell
72
• The structure and function of the contractile
proteins in cardiac cells are essentially the
same as in skeletal muscle
• The T tubule system and sarcoplasmic
reticulum, however, are not as regularly
arranged in cardiac fiber
• The T tubules are more numerous and larger in
cardiac muscle than in skeletal muscle and the
sarcoplasmic reticulum is less well developed
• T tubules are less well-organized and are usually
associated with one expanded terminal cisterna
of SR, forming dyads

73
Cardiac muscle: nuclei (N); intercalated discs (I)
74
Intercalated disc (arrows): desmosomes (D) and
adherent junctions called fascia adherentes (F),
mitochondria (M) and myofibrillar structures similar to
those of skeletal muscle but slightly less organized 75
Cardiac muscle: abundant mitochondria (M); sparse
sarcoplasmic reticulum (SR); T tubules are less well-
organized and are usually associated with one expanded
terminal cisterna of SR, forming dyads (D) 76
• 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
– By comparison, only about 2% of skeletal muscle
fiber is occupied by mitochondria
• Fatty acids are the major fuel of the heart and
are stored as triglycerides in numerous lipid
droplets seen in many cardiac muscle cells
• Glycogen particles may also be present
• Lipofuscin pigment granules are often found
near the nuclei of cardiac muscle cells

77
• A few differences in structure exist between
atrial and ventricular muscle
– The arrangement of myofilaments is the same in
both, but atrial muscle has markedly fewer T
tubules, and the cells are somewhat smaller
– Membrane-limited granules are found at the poles of
atrial muscle nuclei and are associated with Golgi
complexes in this region
– These granules release the peptide hormone atrial
natriuretic factor (ANF) which acts on target cells in
the kidney to affect Na+ excretion and water
balance
– The contractile cells of the heart's atria thus also
serve an endocrine function

78
Smooth Muscle
• Smooth muscle fibers are elongated,
tapering, and nonstriated cells, each of
which is enclosed by a thin basal lamina
and a fine network of reticular fibers
• The connective tissues serve to combine
the forces generated by each smooth
muscle fiber into a concerted action, eg,
peristalsis in the intestine

79
• Smooth muscle cells may range in length from
20 µm in small blood vessels to 500 µm in the
pregnant uterus
• Each cell has a single nucleus located in the
center of the cell's broadest part
• To achieve the tightest packing, the narrow
part of one cell lies adjacent to the broad
parts of neighboring cells
• Such an arrangement viewed in cross section
shows a range of diameters, with only the
largest profiles containing a nucleus
• The borders of the cell become scalloped when
smooth muscle contracts and the nucleus
becomes distorted
80
• Concentrated near the nucleus are
mitochondria, polyribosomes, cisternae of
rough ER, and the Golgi apparatus
• Pinocytotic vesicles are frequent near the
cell surface
• A rudimentary sarcoplasmic reticulum is
present in smooth muscle cells, but T
tubules are not
• The characteristic contractile activity of
smooth muscle is related to the structure
and organization of its actin and myosin
filaments, which do not exhibit the
organization present in striated muscles
81
• In smooth muscle cells, bundles of thin and
thick myofilaments crisscross obliquely through
the cell, forming a latticelike network
• Smooth muscle actin and myosin contract by a
sliding filament mechanism similar to that in
striated muscles
• However, myosin proteins are bundled
differently and the cross-bridges interact with
fewer F-actin filaments
• The thin filaments of smooth muscle cells lack
troponin complexes and instead utilize
calmodulin, a calcium-binding protein

82
Smooth muscle in the wall of the small intestine, cells of
the inner circular (IC) layer are cut lengthwise and cells
of the outer longitudinal layer (OL) cross transversely 83
Smooth muscle in bladder: fibers in cross-section (XS)
and longitudinal section (LS); much collagen in the
branching perimysium (P), but very little evidence of
endomysium is apparent; Mallory trichrome 84
• Smooth muscle cells have an elaborate array of
intermediate filaments
– Desmin is the major intermediate filament protein in
all smooth muscles
– Vimentin is an additional component in vascular
smooth muscle
• Both intermediate filaments and F-actin
filaments insert into dense bodies which can be
membrane-associated or cytoplasmic
• Dense bodies contain actinin and are thus
functionally similar to the Z discs of striated
and cardiac muscles
• The attachments of thin and intermediate
filaments to the dense bodies helps transmit
contractile force to adjacent smooth muscle
cells and their surrounding network of reticular
fibers
85
Thin filaments attach to dense bodies located in the cell membrane
and deep in the cytoplasm 86
• Contraction of smooth muscle is not under
voluntary control, but is regulated by autonomic
nerves, certain hormones, and local physiological
conditions such as the degree of stretch
• The cells occur either as multiunit smooth
muscle, in which each cell is innervated and can
contract independently, or more commonly as
unitary smooth muscle, in which only a few cells
are innervated but all cells are interconnected
by gap junctions
• Gap junctions allow the stimulus for contraction
to spread as a synchronized wave among
adjacent cells
87
• Smooth muscle lacks neuromuscular junctions like those
in skeletal muscle
• Instead axonal swellings with synaptic vesicles simply
lie in close contact with the sarcolemma, with little or
no specialized structure to the junctions
• Because smooth muscle is usually spontaneously active
without nervous stimuli, its nerve supply serves
primarily to modify activity rather than initiate it
• Smooth muscle receives both adrenergic and
cholinergic nerve endings that act antagonistically,
stimulating or depressing its activity
• In addition to contractile activity, smooth muscle cells
also synthesize collagen, elastin, and proteoglycans

88
Contraction decreases the length of the cell, deforming
the nucleus and promoting contraction of the whole
muscle 89
Regeneration of Muscle Tissue
• The three types of adult muscle have different
potentials for regeneration after injury
• Skeletal muscle
– Although the nuclei are incapable of
undergoing mitosis, the tissue can undergo
limited regeneration
– The source of regenerating cells is the
sparse population of mesenchymal satellite
cells that lies within the external lamina of
each mature muscle fiber

90
– Satellite cells are inactive, reserve
myoblasts that persist after muscle
differentiation
– After injury or certain other stimuli, the
normally quiescent satellite cells become
activated, proliferating and fusing to form
new skeletal muscle fibers
– A similar activity of satellite cells has been
implicated in muscle growth after extensive
exercise, a process in which they fuse with
their parent fibers to increase muscle mass
beyond that occurring by cell hypertrophy

91
• Cardiac muscle
– Lacks satellite cells and has virtually no
regenerative capacity
– Defects or damage (eg, infarcts) in heart muscle
are generally replaced by fibroblast
proliferation and growth of connective tissue,
forming myocardial scars
• Smooth muscle
– Composed of simpler, mononucleated cells, is
capable of a more active regenerative response
– After injury, viable smooth muscle cells undergo
mitosis and replace the damaged tissue
– Contractile pericytes from the walls of small
blood vessels participate in the repair of
vascular smooth muscle
92
 Summary of muscle tissues
Type of Shape of Nuclei Control Striatio Regeneration
muscle cells (location) ns

Skeletal Long and Many Voluntary Present  hypertrophy

Cylindrical Peripheral and hyperplasia
 Slight
regeneration

Cardiac Short, One or two Involunta Present  Hypertrophy

cylindrical central ry  No
and regeneration
branching

Smooth Fusiform One Involunta Absent  hypertrophy

Central ry and hyperplasia
 Good
regeneration
93