Vous êtes sur la page 1sur 6

MUSCLE TISSUES

I.

Introduction
A.

Muscle tissue is characterized by cells with extensive cytoskeletal components. As a result


of the extensive cytoskeletal components, muscle cells have greatly amplified the basic
cellular process of contractility.

B.

The function of muscle tissue is contraction. Depending on the type of muscle tissue,
muscle cell contraction in muscle tissue can lead to movement of body parts or movement of
fluids through body parts.

C. Special terminology is used in reference to muscle cell and tissue ultrastructure

II.

1.

A Sarcolemma is a muscle cell plasma membrane.

2.

Sarcoplasm is the cytoplasm of a muscle cell.

3.

Sarcoplasmic reticulum is the endoplasmic reticulum in a muscle cell.

4.

Individual muscle cells (or strands of cells in cardiac muscle) are often referred to as
fibers.

Characteristic Features of Muscle Cells and Tissues


A.

Cells are usually closely packed with a small amount of extracellular material and scattered
fibroblasts between muscle cells. Capillaries are numerous in the extracellular material (see
Fig. 11.1, Plate 18).

B.

Elongated muscle cells (or chains of cells) are arranged in parallel arrays which shorten
along the long axis of the cells when contraction occurs. This arrangement allows the tissue
to work effectively to produce movement.

C. The cytoplasm of muscle cells contains extensive longitudinal arrays of modified actin-rich
microfilaments (called thin myofilaments)in combination with thicker myosin-rich filaments
(called thick myofilaments). Extensive arrays of filaments cause most of the cytoplasm of
muscle cells to be very acidophilic in routine LM preparations.
III. Classification of Muscle Cells/Tissue (Page 306)
Muscle tissue is classified according to the organization of cytoskeletal components and the
cellular organization of the tissue (Table 10.1).
A.

If the cytoskeletal components in the cells are arranged in a manner that produces striations
(transverse stripes) visible at the LM level, then the muscle tissue is classified as striated
muscle (Fig. 11.1, Plate 18).
Two types of cellular organization occur in striated muscle tissues.
a.

Cardiac muscle (Fig. 11.14, Plate 20) is uninucleated (contains 1 nucleus per cell).

b.

Skeletal muscle (Fig. 11.1, Plate 18) is highly multinucleated.

B.

If the cytoskeletal components are not arranged in a manner that produces striations
(transverse stripes) visible at the LM level, then the muscle tissue is classified as nonstriated or smooth muscle. Smooth muscle in humans is uninucleated (Fig 11.18, Plate
22).

IV. Structure of Cells in Muscle Tissue


A.

Some features are common to both striated muscle types (Fig. 11.5 11.7)
1.

Striations visible by LM are named as follows:


a.

A bands = darkly stained bands


Within each A band is a central light area, the H zone or H band, containing a
central dark line, the M line.

b.

I bands = lightly stained bands


Within each I band is a central dark line, the Z line or Z disc.

2.

The region between adjacent Z lines is referred to as a sarcomere.

3.

Thick and thin myofilaments are located consistently within regions within sarcomeres.
a.

Thick myofilaments contain primarily myosin and are located in the A band. Thick
myofilaments are connected to adjacent Z lines by very thin, elastic strands of titin.

b.

Thin myofilaments contain actin complexed with troponin and tropomyosin and
extend from the I band into the A band (ending at the beginning of the H zone).

c.

The H zone contains only thick myofilaments (no thin myofilaments).

d.

The I band contains only thin myofilaments (no thick myofilaments).

e.

The M line contains fine filamentous myomesin which connects adjacent thick
filaments.

f.

The Z line contains short intermediate filaments and alpha actinin which
interconnects thin filaments of adjacent sarcomeres.

4.

Muscle contraction occurs when the thick and thin filaments slide past each other. In a
fully contracted sarcomere the thick filaments come to lie next to the Z line and the thin
filaments come to lie next to the M line. The sliding of the filaments past one another
shortens the sarcomere.

5.

Numerous myofilaments are bundled into a myofibril which is approximately 1 micron in


diameter and which extends for the length of the cell. Sarcomeres of adjacent myofibrils
are in register with one another. The alignment of all the sarcomeres of all the myofibrils
is what produces the striations visible at the LM level. Alignment is very precise in
skeletal muscles, less precise in cardiac muscle.

B.

Skeletal muscle (Fig. 11.1, 11.3, 11.4, 11.5, 11.7, Plate 18, 19)
1.

Individual cells are large, elongated, multinucleate, and contain a large number of
myofibrils. The cells are usually strongly striated.

2.

The cytoplasm between myofibrils contains numerous mitochondria and highly


developed smooth ER.

3.

Invaginations of the cell surface called transverse or T tubules extend down into the
cytoplasm between myofibrils. Vesicles of sER are closely associated with the T tubules
where they pass each myofibril. In mammals transverse T tubules pass along the A-I
band junctions. This arrangement of T tubules allows depolarization of the plasma
membrane to affect centrally located myofibrils.

4.

Several different types of skeletal muscle cells can be identified on the basis of
ultrastructural and cytochemical features (Fig. 11.3).
a.

White fibers (fast twitch fibers) have the following features.


(1) Larger cell diameter with extensive myofilaments
(2) More glycogen and little stored lipid
(3) Lower numbers of mitochondria (correlates with lower succinic dehydrogenase
(SDH) activity)
(4) Little myoglobin

b.

Red fibers (slow twitch fibers) have the following features.


(1)
(2)
(3)
(4)

Smaller cell diameter with fewer myofilaments


Less glycogen and more stored lipid
Higher numbers of mitochondria (correlates with higher SDH activity)
Larger amounts of myoglobin (causes the red color in fresh muscle tissue)

c. Intermediate fibers have features intermediate between white and red fibers.
5.

Skeletal muscle normally contracts only in response to synaptic stimuli from motor
neurons. Each skeletal muscle cell contracts independently, but a single motor neuron
may innervate many muscle cells which then respond simultaneously as a motor unit.

6.

The sequence of events which leads to contraction is complex but occurs very quickly.
a.

Neurotransmitter release from a motor neuron synapse with a skeletal muscle cell
triggers depolarization of the sarcolemma by opening sodium channels across the
membrane (Figs. 11.9 - 11.11).

b.

The depolarization triggers an action potential which spreads along the surface of
the entire muscle cell and at least partially down each T tubule (Figs. 11.11 and
11.8).

c.

The depolarization and/or the sodium ions which enter the cell during the
depolarization trigger calcium ion release from the sarcoplasmic reticulum (Fig.
11.11).

d.

The calcium ions remove the troponin blockage of the active sites on the actin in the
thin myofilaments, activating the thin filaments.

7.

e.

The thin and thick myofilaments interact, using ATP energy to power sliding of the
filaments past each other to shorten each sarcomere (pages 288 299).

f.

Active transport moves the calcium ions back into the sarcoplasmic reticulum,
inactivating the thin myofilaments and causing the muscle to relax (Fig. 11.11).

Skeletal muscle cells are attached at both ends to the connective tissue framework of
the muscle. Shortening of the cell therefore creates force on the connective tissue. The
attachment sites resemble large hemidesmosomes when viewed by TEM, but the
filaments involved on the cytoplasmic side of the junction are the thin myofilaments of
the terminal sarcomeres which are modified microfilaments rather than intermediate
filaments. The attachment junctions are therefore related to zonula adherens and are
called fasciae adherens (see Plate 19).

C. Cardiac muscle (Fig. 11.14, 11.16, Plate 20)


1.

Individual cells are short and uninucleate and are weakly striated. Individual cells are
joined end to end to form long fibers. Cells and fibers are often branched.

2.

Cells are joined end to end (to form fibers) by specialized junctions called intercalated
discs. Intercalated disks have transverse and longitudinal components. The transverse
components are located at Z lines of sarcomeres on the myofibrils. Each intercalated
disk contains 3 types of junctions.
a.

Macula communicans junctions are primarily located on longitudinal regions of


intercalated disks. These allow ions and electrical stimuli to pass from cell to cell.

b.

Macula adherens junctions occur at intervals on the transverse regions of


intercalated disks and contribute to mechanical attachment.

c.

Fasciae adherens resemble zonula adherens junctions but occur as irregular


regions rather than a continuous band. Fasciae adherens occur in transverse
regions of intercalated disks and contribute to mechanical attachment. The thin
myofilaments of the terminal sarcomeres insert into the fasciae adherens.

3.

Register of sarcomeres is less precise (relative to striated skeletal muscle) so striations


are not as obvious as in skeletal muscle.

4.

T tubules are larger in diameter than in skeletal muscle and are located at Z lines of the
sarcomeres on the myofibrils. Vesicles of sER are located around T tubules but sER is
not as closely associated with the T tubules as in skeletal muscle.

5.

Cardiac muscle contracts spontaneously at a frequency determined by the cells with the
fastest intrinsic frequency. The triggering depolarization spreads as an action potential
along each cell and moves from cell to cell through the maculae communicans in the
intercalated disks.

6.

The contraction mechanism triggered by depolarization in cardiac muscle closely


resembles the process in skeletal muscle.

7.

Cardiac muscle cells are mechanically joined end to end by their intercalated discs to
form fibers. Contraction of each cell therefore shortens the fiber. The ends of the fiber
attach to the connective tissue around the valve through which the blood will leave the
heart chamber. Contraction of the cell therefore decreases the volume of the heart
chamber and pulls the wall toward the valve, efficiently forcing out the blood in the
chamber.

D. Smooth muscle (Fig. 11.18 11.23, Plate 22)


1.

Individual cells are long tapered cylinders or spindles and are uninucleate with a
centrally located nucleus. The cells are unstriated.

2.

Invaginations of the cell surface called caveolae extend into the peripheral cytoplasm
and are associated with numerous pinocytic vesicles in the cytoplasm.

3.

The cytoskeleton is organized differently than in striated muscle cells.

4.

a.

Longitudinally oriented thin myofilaments are abundant in the peripheral cytoplasm.


Thick myofilaments are less abundant than in skeletal or cardiac muscle. Thick
myofilaments can be demonstrated but may not be permanent structures in all types
of smooth muscle cells. The myofilaments are not organized into myofibrils with
sarcomeres, since the thick myofilaments are "side-polar" rather than the bipolar
thick filaments found in striated muscle cells (Fig. 11.21).

b.

Thin filaments are attached to the plasma membrane so the membrane shortens
along with the myofilaments when the cell contracts. The attachment occurs
through alpha actinin-rich dense bodies (Fig. 11.22). Dense bodies also occur in
the cytoplasm, apparently interconnecting sets of thin filaments.

c.

Intermediate filaments (containing desmin) are involved in the cytoskeletal


framework for the cell, but intermediate filaments can be removed without
destroying the contractile ability of the cell.

d.

Contraction appears to be initiated by chemical modification of the myosin in


response to chemical signals from autonomic nerve endings (or other sources)
received by cell surface receptors. Calcium ions are involved but do not appear to
be part of the triggering events which activate actin-myosin complexes in the same
way they do in skeletal and cardiac muscle.

Two functional types of smooth muscle tissue have been described.


a.

Visceral smooth muscle (gut wall, etc.) cells are connected by numerous macula
communicans junctions which allow passage of electrical impulses between cells.
Autonomic nerve endings associate very loosely with the cells, and many muscle
cells are not near nerve endings. This type of smooth muscle contracts as a unit,
frequently in a progressive wave fashion. Visceral smooth muscle is capable of
spontaneous contraction, particularly if stretched.

b.

Multiunit smooth muscle (vascular walls, etc.) cells have fewer macula
communicans junctions. Most muscle cells are closely innervated by an autonomic
nerve ending. Cells in this muscle contract relatively individually in response to
nervous stimulation.

5.

Smooth muscle cells are surrounded by a basal lamina and reticular lamina (both
secreted by the smooth muscle cells), outside of which (and conneced to the reticular
lamina) are fine collagen and elastic fibers forming a stromal framework for the muscle
layer. Hemidesmosome (macula adherens) junctions connect the cytoskeleton of the
cell to the basal lamina. Therefore when the cell contracts, it pulls on the connective
tissue framework, producing tension.