Muscle Tissue

• Composed of cells that optimize the universal cell property of

contractility.
• Actin microfilaments and associated proteins generate the forces
necessary for the muscle contraction, which drives movement within
the organ systems, of blood, and of the body as a whole.
• Muscle cells are of mesodermal origin and differentiate by a gradual
process of cell lengthening with abundant synthesis of the myofibrillar
proteins actin and myosin.
3 types of muscle tissue
• Skeletal muscle contains bundles of very long, multinucleated cells with
cross-striations. Their contraction is quick, forceful, and usually under
voluntary control.

• Cardiac muscle also has cross-striations and is composed of elongated,

often branched cells bound to another at structures call intercalated discs
that are unique to cardia muscle. Contraction is involuntary, vigorous, and
rhythmic.

• Smooth muscle consists of collections of fusiform cells that lack striations

and have slow, involuntary contractions.
Key notes…
• In all types of muscle, contraction is caused by the sliding interaction
of thick myosin filaments along thin actin filaments.

• The cytoplasm of muscle cells is often called sarcoplasm, the smooth

ER is the sarcoplasmic reticulum, and the muscle cell membrane and
its external lamina are the sarcolemma.
Skeletal muscle
•Long, cylindrical multinucleated cells with
diameters of 10 to 100 µm.
•Elongated nuclei are found in the periphery just
under the sarcolemma, a characteristic nuclear
location unique to skeletal muscle fibers/cells.
Organization of a skeletal muscle
• Thin layers of connective tissue surround and organize the contractile
fibers in all 3 types of muscle (seen specially in skeletal muscle). These
supportive layers are:
• The epimysium, an external sheath of dense connective tissue, surrounds
the entire muscle, carrying the larger nerves, blood vessels, and lymphatics of
the muscle.
• The perimysium is a thin connective tissue layer that immediately surrounds
each bundle of muscle fibers termed a fascicle.
• Within fascicles a very thin, delicate layer of reticular fibers and scattered
fibroblasts, the endomysium, surrounds the external lamina of individual
muscle fibers. In addition to nerve fibers, capillaries form a rich network in
the endomysium bringing oxygen to the muscle fibers
Key notes…
• Some skeletal muscles taper at their ends, where the epimysium is
continuous with the dense regular connective tissue of a tendon at
myotendinous junctions.
Organization within muscle fibers
• Longitudinal sectioned skeletal muscle fibers show cross-striations of
alternating light and dark bands.
• The dark bands are called A bands (anisotropic or birefringent in
polarized light microscopy); the light bands are called I bands
(isotropic, do not alter polarized light).
• In TEM, each I band is seen to be bisected by a dark transverse line,
the Z disc.
• The repetitive functional subunit of the contractile apparatus, the
sarcomere, extends from Z disc to Z disc.
Key notes…
• The A and I banding pattern in sarcomeres is due mainly to the regular arrangement
of thick and thin myofilaments, composed of myosin and F-actin, respectively.
• The thick myosin filaments occupy the A band at the middle region of the sarcomere.
• Myosin has two identical heavy chains and two pairs of light chains. Globular
projections containing the four myosin light chains form a head at one end of each
heavy chain.
• The myosin heads bind both actin, forming transient crossbridges between the thick
and thin filaments, and ATP (catalyzing energy release).
• The thin, helical actin filaments run between the thick filaments. Each G-actin
monomer contains a binding site for myosin.
• Actin filaments are anchored perpendicularly on the Z disc by the actin-binding
protein α-actinin.
Continuation…
• Thin filaments are also tightly associated with two regulatory proteins.
• Tropomyosin
• Troponin, a complex of three subunits: TnT, which attaches to tropomyosin; TnC, which
binds Ca²+; and TnI, which regulates the actin-myosin interaction.
• An important accessory protein in I bands is titin, the largest protein in the body, with scaffolding
and elastic properties, which supports the thick myofilaments and connects the Z disc.
• Another very large accessory protein, nebulin, binds each thin myofilament laterally, helps anchor
them to α-actinin, and specifies the length of the actin polymers during myogenesis.
• The H zone is the lighter zone in the center of the A band with only the rodlike portion of the
myosin molecule and no thin filaments.
• Bisecting the H zone is the M line, containing a myosin-binding protein called myomesin that
holds the thick filaments in place, and creatine kinase. This enzyme catalyzes transfer of
phosphates groups from phosphocreatine, a storage form of high-energy phosphate groups, to
ADP, helping to supply ATP for muscle contraction.
Sarcoplasmic Reticulum and
Transverse Tubule System
• In skeletal muscle, the sarcoplasmic reticulum is specialized for Ca+²
sequestration.
• Transverse of T tubules are long fingerlike invaginations of the cell
membrane that penetrate deeply into the sarcoplasm and encircle
every myofibril near the aligned A- and I-band boundaries of
sarcomeres.
• Adjacent to each side of every tubule are expanded terminal cisterns
of the sarcoplasmic reticulum. This complex of a T tubule with two
closely associated small cisterns of sarcoplasmic reticulum on each
side is known as a triad.
Mechanism of contraction
• During contraction neither thick or thin filaments change their
length.
• Contraction results as the overlapping thin and thick filaments of each
sarcomere slide past one another.
• Contraction is induced when an action potential arrive at a synapse,
the neuromuscular junction, and is transmitted along the T tubules to
the sarcoplasmic reticulum to trigger Ca²+ release.
Key notes…
• In a resting muscle, the myosin heads cannot bind G-actin because the
binding sites are blocked by the troponin-tropomyosin complex on the F-
actin filaments.
• Calcium ions released upon neural stimulation bind troponin, changing
its shape and moving tropomyosin on the F-actin to expose the myosin-
binding active sites and allow crossbridges to form.
• Binding actin produces a conformational change or pivot in the myosins,
which pulls the thin filaments farther into the A band, toward the Z disc.
• Energy for the pivot and pulling of the actin is provided by the hydrolysis
of ATP bound to the myosin heads.
Medical application
• Myasthenia gravis is an autoimmune disorder that involves circulating
antibodies against proteins of acetylcholine receptors. Antibody
binding to the antigenic sites interferes with acetylcholine activation
of their receptors, leading to intermittent periods of skeletal muscle
weakness.
Muscle Spindles and Tendon Organs
• Striated muscles and myotendinous junctions contain sensory receptors acting as proprioceptors
(a sensory receptor that receives stimuli from within the body, especially one that responds to
position and movement), providing the CNS data from the musculoskeletal system.
• Proprioceptors are specialized sensory receptors that are located within joints, muscles, and
tendons.
• These receptors are sensitive to both tension and pressure, they play a role in relaying
information concerning muscle dynamics to the conscious and subconscious parts of the central
nervous system.
• They provide the brain with information concerning kinesthetic sense, or conscious appreciation
of the position of body parts with respect to gravity.
• Most of this proprioceptive information is processed at a subconscious level, therefore we do
not have to dedicate any conscious activity toward tasks such as maintaining posture or position
of body parts.
• Golgi tendon organs (GTOs) are an example of proprioceptors. They are located in tendons near
the myotendinous junction and are in series, that is, attached end to end, with extrafusal muscle
fibers.
Continuation…
• Among the muscle fascicle are stretch detectors known as muscle
spindles.
• Muscle spindles are skeletal muscle sensory receptors within the
body of a muscle that primarily detect changes in the length of
muscle contributing to fine motor control and providing axial and
limb position information to the central nervous system.
Muscle fiber types
• Muscles that are often active with slow contractions for long periods
tend to have more mitochondria for oxidative phosphorylation and
ATP production. Such fibers have high levels of myoglobin, which
contains iron and oxygen stores, giving such fibers a red color in fresh
tissue.
• Muscles specialized for short-term work and fast contraction are
typically larger in diameter and depend more heavily on anaerobic
(glycolytic) metabolism of glucose, much of which is derived from
stored glycogen.
Key terms…
• Muscle fiber is actually a muscle cell.
• Myofibrils are longitudinal arrays of contractile filaments in the
cytoplasm of muscle cells.
• Myofibrils are composed of thick (myosin) and thin (actin)
myofilaments.
• A sarcomere is the segment of a myofibril that forms the basic
functional unit of skeletal muscle.
Cardiac Muscle
• During embryonic development, the mesoderm cells of the primitive
heart tube align in chainlike arrays rather than fusing into
multinucleated cells like that of skeletal muscle fibers.
• Cardiac muscle cells form complex junctions between interdigitating
processes.
• Cells within a fiber often branch and bind to cells in adjacent fibers.
• Muscle cells exhibit cross striations comparable to skeletal muscle
fibers.
• Each cardiac muscle cell possesses only one (sometimes two)
centrally located nucleus.
Key points…
• A unique and distinguishing characteristics of cardiac muscle is the presence of dark-
staining transverse lines that cross the chains of cardiac cells at irregular intervals where
the cells join. These intercalated discs represent the interface between adjacent muscle
cells and contain junctional complexes (desmosomes, fascia adherens, and gap junctions).
• Cardiac muscles contain numerous mitochondria, reflecting their need for continuous
aerobic metabolism.
• Fatty acids are the major fuel of the heart and are stored as triglycerides which can be
seen in numerous lipid droplets in many cardiac cells.
• Muscle of the ventricles is much thicker than that of the atria.
• The muscles of the atria have cytoplasmic granules that release the atrial natriuretic
factor (ANF) that acts on the target cells of the kidney to affect sodium and water
balance.
• Cardiac muscle fiber contraction is intrinsic and spontaneous.
Smooth Muscles
• Is specialized for slow, steady contraction and is controlled by a variety of
involuntary mechanisms.
• Fibers of smooth muscle cells (also called visceral muscle) are elongated, tapering,
and nonstriated cells enclosed by a thin basal lamina and fine network of reticular
fibers, the endomysium.
• Each cell has a single long nucleus located in the cell’s central, broadest part.
• All cells are linked by numerous gap junctions.
• Smooth muscle fibers have rudimentary sarcoplasmic reticulum and lack T
tubules.
• Caveolae (short membrane invaginations) of smooth muscles contains several
pumps and ion channels and may serve to organize proteins signaling release
calcium release at myofibrils.
Continuation…
• In smooth muscle cells, bundles of thin and thick myofilaments crisscross obliquely
through the cell.
• Actin filaments lack troponin, using instead calmoudin and Ca²+ -sensitive myosin light
chain kinase (MLCK) in the contraction mechanism.
• Smooth muscle fibers lack MEP’s.
• Because smooth muscle is most often spontaneously active without nervous stimuli, its
nerve supply serves primarily to modify activity rather than to initiate it.
• Smooth muscle receives both adrenergic and cholinergic nerve endings that act
antagonistically, stimulating or depressing its activity.
• Smooth muscle contraction is determined largely by the degree of autonomic
innervation and the density of the gap junctions.; both conditions vary considerably in
different organs.
• Smooth muscles demonstrate the characteristic features of secretory cells. For
example, endomysium, and the basal lamina are produced and deposited by smooth
muscle cells.