Skeletal muscle tissue

Subject : histology
Teacher : Aygun Alyarbayova

Name : Arfina Aljaseer

Group : 121 i2b
Muscle Tissue:

Muscle tissue is composed of cells that have the special ability to shorten or contract in
order to produce movement of the body parts. The tissue is highly cellular and is well
supplied with blood vessels. The cells are long and slender so they are sometimes called
muscle fibers, and these are usually arranged in bundles or layers that are surrounded
by connective tissue. Actin and myosin are contractile proteins in muscle tissue.

The 3 types of muscle tissue are cardiac, smooth, and skeletal. Cardiac muscle cells are
located in the walls of the heart, appear striped (striated), and are under involuntary
control. Smooth muscle fibers are located in walls of hollow visceral organs (such as the
liver, pancreas, and intestines), except the heart, appear spindle-shaped, and are also
under involuntary control. Skeletal muscle fibers occur in muscles which are attached to
the skeleton. They are striated in appearance and are under voluntary control.
Skeletal Muscle Tissue

The musculoskeletal system comprises one of the

major tissue/organ systems in the body. The three
main types of muscle tissue are skeletal, cardiac,
and smooth muscle groups. Skeletal muscle attaches
to the bone by tendons, and together they produce
all the movements of the body. The skeletal muscle
fibers are crossed with a regular pattern of fine red
and white lines, giving the muscle a distinctive
striated appearance. Hence they are also known as
striated muscles.

Skeletal muscles are voluntary muscles under the

control of the somatic nervous system. The other
types of muscle are cardiac muscle which is also
striated and smooth muscle which is non-striated;
both of these types of muscle tissue are classified as
involuntary, or, under the control of the autonomic
nervous system.
Skeletal muscle fibers are cylindrical,
multinucleated, striated, and under
voluntary control. A skeletal muscle
contains multiple fascicles – bundles of
muscle fibers. Each individual fiber,
and each muscle is surrounded by a
type of connective tissue layer of
fascia. Muscle fibers are formed from
the fusion of developmental myoblasts
in a process known as myogenesis
resulting in long multinucleated cells.
In these cells the nuclei termed
myonuclei are located along the inside
of the cell membrane. Muscle fibers
also have multiple mitochondria to
meet energy needs. Skeletal muscle
tissue is also extensible and elastic.
Extensible tissue can be stretched and
elastic tissue is able to return to its
original shape following distortion.
Location
Skeletal muscle, as the name implies, is any muscles that
connects to and controls the motions of the skeleton. In all
there are somewhere between 600 and 900 muscles in the
human body, but an exact number is hard. Many muscles are
obscurely small or are sometimes grouped together with
similar muscles. Skeletal muscle is found between bones, and
uses tendons to connect the epimysium to the periosteum, or
outer covering, of bone.

Skeletal muscle is adapted and shaped in many different

ways, which give rise to complex movements. Skeletons are
not always internal as they are in humans. Even animals with
exoskeletons, like crabs and mussels, have skeletal muscle.
While the muscle might be adapted differently depending on
the animal, skeletal muscle is defined by its striations and
connections to skeleton. Everything from the flapping of a
bird’s wings to the crawling of a beetle are carried out by
skeletal muscle.
 Embryonic Development Of Skeletal Muscles
Skeletal muscle is derived from the mesoderm. Cells in the
ventromedial part of the somite form the sclerotome. Cells
in the dorsal part form the dermatome and two edges, the
ventrolateral lip and the dorsomedial lip. Cells from these
two edges migrate ventral to the dermatome and proliferate
to form muscle cell precursors. Collectively, these structures
form the dermomyotome. In turn, the dermomyotome will
differentiate into dermatome cells forming the dermis of the
back and the neck, and myotome cells forming the skeletal
muscles.

During the course of embryonic development, mesenchymal

progenitor cells originating from the somites, undergo a
multistep differentiation process to form muscle fibers and
muscle mass. Each somitic area (occipital, cervical, thoracic,
lumbar and sacral) contributes to the formation of muscles.
Muscle satellite cells are formed during embryonic
development as well, and persist in a quiescent state in the
adult muscles, ensuring restoration of muscle cells following
any type of muscle injury.
The head mesoderm constitutes an
additional source of progenitor cells,
which contribute to the formation of
head muscles, or craniofacial muscles.
These progenitor cells are primarily
contributed from the unsegmented
cranial paraxial and splanchnic
mesoderm, which bear a somewhat
different genetic program, when
compared to their counterparts in the
somites. The prechordal mesoderm
cells, which reside in the most anterior
part of the forming head, represent
another population of progenitor cells.
These cells eventually give rise to the
extraoccular muscles (EOM, eye
muscles). The formation of vertebrate
head muscles allow for vision,
mastication and food uptake, facial
expression and breathing in humans.
 Structure
Skeletal muscle tissue is made up of a collection of muscle
fibers wrapped in connective tissue sheaths. There are
three types of connective tissue sheaths named for their
location. Endomysium surrounds individual muscle fibers.
It is made up of a delicate layer of reticular fibers and
permits only small-diameter nerve fibers and capillaries,
thus acting as a site of metabolic exchange.

Perimysium is a slightly thicker layer of connective tissue

consisting mainly of type I and III collagen and surrounds a
group of fibers. This fiber group is referred to as a fascicle
or bundle. Fascicles are the functional units of skeletal
muscle tissue.

The perimysium contains slightly larger blood vessels and

nerve fibers than those traveling through endomysium.

Epimysium surrounds the entire collection of fascicles

making up an individual muscle. This dense connective
tissue made up of mainly type I collagen contains the
neurovascular supply to the muscle.
 Types Of Skeletal Muscles
There are three types of skeletal muscle fibers: Type I, Type IIa, and Type IIb.

Type I muscle fibers, also called slow oxidative fibers, are specialized for aerobic
activity. They are small, contain a high amount of myoglobin, and appear red in fresh
tissue. A muscle twitch is a single contraction of a muscle. Type I fibers make up slow-
twitch, fatigue-resistant motor units. Muscles of the deep back responsible for
maintaining posture are mostly made up of Type I slow oxidative fibers.

Type IIa muscle fibers are also known as fast oxidative glycolytic fibers. These fibers
appear slightly lighter than Type I in fresh tissues. They contain many mitochondria and
have a higher myoglobin content than type IIb fibers. Unlike Type I fibers, Type IIa fibers
have high amounts of glycogen. Because of this they are capable of anaerobic glycolysis
and make up fast-twitch, fatigue resistant motor units. Type IIa fibers are more fatigue
resistant than Type IIb fibers and are used in movements that require high sustained
power. Many athletes have high amounts of these fibers, especially competitive
swimmers.
Type IIb muscle fibers are also referred to as fast glycolytic fibers. They are large fibers and
appear light pink in fresh tissues. Type IIb fibers contain fewer mitochondria and a lower amount
of myoglobin. Although they contain a low level of oxidative enzymes, they show high anaerobic
enzyme activity and contain a high amount of glycogen. Type IIb fibers are more prone to fatigue
than Type I and Type IIa fibers and make fast-twitch, fatigue prone motor units. Type IIb fibers
have the fastest rate of ATPase activity and are found in muscles used for short, rapid bursts of
contraction such as the gastrocnemius, a muscle in the leg that is used in jumping.
 Functional Characteristics
Skeletal muscle, with its associated connective tissue, constitutes approximately 40% of body weight. Skeletal
muscle is so named because most of the muscles are attached to the skeletal system. It is also called striated
muscle because transverse bands, or striations, can be seen in the muscle under the microscope.

Skeletal muscle has four major functional characteristics: contractility, excitability, extensibility, and
elasticity.

1.Contractility is the ability of skeletal muscle to shorten with force. When skeletal muscles contract, they
cause the structures to which they are attached to move. Skeletal muscles shorten forcefully during
contraction, but they lengthen passively. Either gravity or the contraction of an opposing muscle produces a
force that pulls on the shortened muscle, causing it to lengthen.

2.Excitability is the capacity of skeletal muscle to respond to a stimulus. Normally, the stimulus is from nerves
that we consciously control.

3.Extensibility means that skeletal muscles stretch. After a contraction, skeletal muscles can be stretched to
their normal resting length and beyond to a limited degree.

4.Elasticity is the ability of skeletal muscles to recoil to their original resting length after they have been
stretched.
Skeletal Muscle Growth And Regeneration
Most muscle tissue of the body arises from embryonic mesoderm. Paraxial mesodermal cells adjacent to the
neural tube form blocks of cells called somites. Skeletal muscles, excluding those of the head and limbs,
develop from mesodermal somites, whereas skeletal muscle in the head and limbs develop from general
mesoderm. Somites give rise to myoblasts. A myoblast is a muscle-forming stem cell that migrates to
different regions in the body and then fuses to form a syncytium, or myotube. As a myotube is formed from
many different myoblast cells, it contains many nuclei, but has a continuous cytoplasm. This is why skeletal
muscle cells are multinucleate, as the nucleus of each contributing myoblast remains intact in the mature
skeletal muscle cell. However, cardiac and smooth muscle cells are not multinucleate because the
myoblasts that form their cells do not fuse.
Regeneration
Skeletal muscle contains numerous 'satellite cells' underneath the basal lamina, as shown in the
photograph opposite. These are mono nucleated quiescent cells. When the muscle is damaged, these
cells are stimulated to divide. After dividing, the cells fuse with existing muscle fibres, to regenerate
and repair the damaged fibres. Although the number of muscle cells is set during development,
satellite cells help to repair skeletal muscle cells. A satellite cell is similar to a myoblast because it is a
type of stem cell however, satellite cells are incorporated into muscle cells and facilitate the protein
synthesis required for repair and growth. These cells are located outside the sarcolemma and are
stimulated to grow and fuse with muscle cells by growth factors that are released by muscle fibers
under certain forms of stress. Satellite cells can regenerate muscle fibers to a very limited extent, but
they primarily help to repair damage in living cells. If a cell is damaged to a greater extent than can be
repaired by satellite cells, the muscle fibers are replaced by scar tissue in a process called fibrosis.
Because scar tissue cannot contract, muscle that has sustained significant damage loses strength and
cannot produce the same amount of power or endurance as it could before being damaged.

The skeletal muscle fibres themselves, cannot divide. However, muscle fibres can lay down new
protein and enlarge (hypertrophy).
Innervation Of Skeletal Muscles
Each skeletal muscle fiber is innervated by a single motor axon. The same axon may also innervate
other muscle fibers. All the fibers innervated by the same axon are called a motor unit. Motor
units are small, with one or few fibers, in muscles with delicate action like those moving the fingers
or eyes, they are much larger in muscles with cruder action, like those in the back or thigh.

Muscle action begins at the motor end plate (or neuromuscular junction), which is analogous to a
synapse. The neuromuscular junction is a site located midway along the length of a muscle fiber
where a motor axon terminal releases transmitter (acetylcholine) into the junctional cleft between
axon and muscle membranes. Acetylcholine binds to receptors localized in the muscle membrane
at the motor end plate. Resulting local depolarization at the end plate (the "end plate potential")
normally exceeds threshold and triggers a depolarizing action potential, just like that which travels
along axon membranes. This muscle action potential zips toward both ends of the muscle fiber,
accomplishing coordinated activation of the entire length of the fiber by a single axon terminal.
Vascularization Of Skeletal Muscles
Vascularization of skeletal muscle occurs by four distinct processes: vasculogenesis, angiogenesis,
arteriogenesis, and lymphogenesis.

Vasculogenesis
Vasculogenesis describes the formation of the primitive network of blood vessel in the embryo from
undifferentiated precursor cells (angioblasts), and the differentiation of angioblasts to endothelial cell. This
is the initial step to blood vessel formation. To form a new vessel, angioblasts proliferate and join up with
primary capillary plexus. The endothelial cell grid manufactured by vasculogenesis then serves as an
angiogenesis framework. After primary capillary plexus formation, it is altered by the sprouting and
branching of new vessels from pre-existing ones. The majority of work on skeletal muscle capillaries has
focused on angiogenesis.

Angiogenesis
Angiogenesis and inflammation are critical to the process of muscle regeneration Angiogenesis involves the
growth of new capillaries from existing blood vessels within the skeletal muscle and occurs in the adult
during the ovarian cycle and in physiological repair processes such as wound healing. However, very little
turnover of endothelial cells occurs in the adult vasculature. Angiogenesis in the skeletal muscle can occur
by two primary mechanisms: sprouting and intussusception.
Arteriogenesis
Arteriogenesis, formerly regarded as a variant of angiogenesis, is a relatively new term that was introduced to
distinguish it from other mechanisms of vascular growth, such as angiogenesis and vasculogenesis.
arteriogenesis describes the formation of mature arteries from pre-existent interconnecting arterioles after an
arterial occlusion. Arteriogenesis pointed to the enlargement of existing arterial vessels. This enlargement
indicates an increase in the caliber (diameter) and wall dimensions, resulting in a larger vessel. Transformation
of pericytes into smooth muscle cells and/or apposition of mesenchymal cells, most likely fibroblasts, to the
abluminal surface of capillaries, followed by their gradual change into pericytes or smooth muscle cells lead to
the growth of arteriolar known as “arteriolarization”.

Lymphogenesis
Lymphatic vessels act in close relation with blood vessels. The formed lymph fluid is transported via initial
lymphatic capillaries to collecting vessels, to lymph nodes, and finally back to the blood. Hyperemia-induced
increased filtration of skeletal muscle (during activity), promotes hydrostatic and colloid osmotic pressure and
addresses the need for increased lymph flow to maintain optimal conditions in the muscle. It is clear that
exercise increases skeletal muscle lymph flow significantly in both animals and humans, especially at the
beginning of exercise.
skeletal muscles contain small capillary-sized lymphatic vessels, which are located next to blood capillaries
between muscle fibers but are much fewer in number.
Myofibrils

A myofibril is a basic rod-like organelle of a

muscle cell.Muscles are composed of tubular
cells called myocytes, known as muscle fibres in
striated muscle, and these cells in turn contain
many chains of myofibrils. They are created
during embryonic development in a process
known as myogenesis.

Myofibrils are composed of long proteins

including actin, myosin, and titin, and other
proteins that hold them together. These proteins
are organized into thick and thin filaments
called myofilaments, which repeat along the
length of the myofibril in sections called
sarcomeres. Muscles contract by sliding the
thick (myosin) and thin (actin) filaments along
each other.
Structure

The filaments of myofibrils, myofilaments, consist of two types, thick and thin:

Thin filaments consist primarily of the protein actin, coiled with nebulin filaments. Actin, when
polymerized into filaments, forms the "ladder" along which the myosin filaments "climb" to generate
motion
Thick filaments consist primarily of the protein myosin, held in place by titin filaments. Myosin is
responsible for force generation. It is composed of a globular head with both ATP and actin binding sites,
and a long tail involved in its polymerization into myosin filaments.
The protein complex composed of actin and myosin is sometimes referred to as "actinomyosin".
The Sarcomere: The Basic Contractile Unit of Muscle
A sarcomere is the basic contractile unit of muscle fiber. Each sarcomere is composed of two main protein
filaments, actin and myosin which are the active structures responsible for muscular contraction. The most
popular model that describes muscular contraction is called the sliding filament theory. In this theory, active
force is generated as actin filaments slide past the myosin filaments, resulting in contraction of an individual
sarcomere.

• A sarcomere is defined as the segment between two neighbouring Z-lines. In electron micrographs of cross-
striated muscle, the Z-line appears in between the I-bands as a dark line that anchors the actin
myofilaments.
• Surrounding the Z-line is the region of the I-band. I-band is the zone of thin filaments that is not
superimposed by myosin
• Following the I-band is the A-band. Named for their properties under a polarized light microscope. An A-
band contains the entire length of a single thick filament. The anisotropic band contains both thick and
thin filaments.
• Within the A-band is a paler region called the H-zone . Named for their lighter appearance under a
polarization microscope. H-band is the zone of the thick filaments that has no actin.
• Within the H-zone is a thin M-line, appears in the middle of the sarcomere formed of cross-connecting
elements of the cytoskeleton.
Mechanism Of Muscle Contraction

The process of muscular contraction

occurs over a number of key steps,
including:

1. Depolarisation and calcium ion release

2. Actin and myosin cross-bridge
formation
3. Sliding mechanism of actin and myosin
filaments
4. Sarcomere shortening (muscle
contraction)
1. Depolarisation and Calcium Ion Release

• An action potential from a motor neuron triggers the release of acetylcholine into the motor
end plate
• Acetylcholine initiates depolarisation within the sarcolemma, which is spread through the
muscle fibre via T tubules
• Depolarisation causes the sarcoplasmic reticulum to release stores of calcium ions (Ca2+)
• Calcium ions play a pivotal role in initiating muscular contractions.

2. Actin and Myosin Cross-Bridge Formation

• On actin, the binding sites for the myosin heads are covered by a blocking complex (troponin
and tropomyosin)
• Calcium ions bind to troponin and reconfigure the complex, exposing the binding sites for the
myosin heads
• The myosin heads then form a cross-bridge with the actin filaments.
3. Sliding Mechanism of Actin and Myosin

• ATP binds to the myosin head, breaking the cross-bridge between actin and myosin
• ATP hydrolysis causes the myosin heads to change position and swivel, moving them
towards the next actin binding site
• The myosin heads bind to the new actin sites and return to their original conformation
• This reorientation drags the actin along the myosin in a sliding mechanism
• The myosin heads move the actin filaments in a similar fashion to the way in which an oar
propels a row boat
4. Sarcomere Shortening
• The repeated reorientation of the myosin heads drags the actin filaments along the length of
the myosin
• As actin filaments are anchored to Z lines, the dragging of actin pulls the Z lines closer
together, shortening the sarcomere
• As the individual sarcomeres become shorter in length, the muscle fibres as a whole
contracts
Control Of Skeletal Muscle Contraction
Two main proprioceptors influence the strength and duration of muscle contraction: Golgi tendon organs and
muscle spindle fibers. Golgi tendon organs relay information about the force of contraction. Muscle spindles
detect changes in muscle length.

Golgi tendon organs

Golgi tendon organs are receptors found at the junction between myofibrils and tendons. The receptor
endings of a Golgi tendon organ are intertwined with the collagenous fibers of the tendon. When the muscle
in contracted, the tendon is stretched and the receptors are compressed in a web of connective tissue. The
pressure initiates an action potential in the nerve ending and sends a signal through the central nervous
system. The role of the Golgi tendon organ is to prevent muscles from over-contracting.

Muscle spindle fibers

Muscle spindles detect change in muscle length, position, and velocity. Muscle spindle fibers are long, thin
encapsulated fibers aligned in parallel to myofibrils. They are also called intrafusal muscle fibers, while
extrafusal fibers are are present outside the spindle capsule. The ends of the intrafusal muscle fibers are
attached to myofibrils so that when the muscle is stretched, the muscle spindle fibers stretch as well.
Stretching the spindle causes it to depolarize and relay that information through the spinal cord.
References

https://ib.bioninja.com.au/higher-level/topic-11-animal-physiology/112-movement/muscle-
contraction.html
https://histology.siu.edu/ssb/muscle.htm
https://opentextbc.ca/anatomyandphysiologyopenstax/chapter/development-and-regeneration-
of-muscle-tissue/
https://www.brainkart.com/article/Characteristics-of-Skeletal-Muscle_21789/
https://www.kenhub.com/en/library/anatomy/development-of-musculoskeletal-system
https://biologydictionary.net/skeletal-muscle/
http://www.proteinatlas.org/learn/dictionary/normal/skeletal+muscle/detail+1/magnification+1
https://edu.glogster.com/glog/skeletal-muscle-tissue/24qnuwwbacg
https://www.intechopen.com/chapters/66964
https://training.seer.cancer.gov/anatomy/cells_tissues_membranes/tissues/muscle.html