Skeletal Muscle
Learning Objectives
Describe the structure and function of skeletal muscle fibers
By the end of this section, you will be able to:
• Describe the connective tissue layers surrounding skeletal muscle
• Define a muscle fiber, myofibril, and sarcomere
• List the major sarcomeric proteins involved with contraction
• Identify the regions of the sarcomere and whether they change during
contraction
• Explain the sliding filament process of muscle contraction
Skeletal Muscle Definition
Skeletal muscle is one of the three types of muscles in the human body- the others being
visceral and cardiac muscles. In this lesson, skeletal muscles, its definition, structure,
properties, functions, and types are explained in an easy and detailed manner.
Skeletal muscle is a muscle tissue that is attached to the bones and is involved in the
functioning of different parts of the body. These muscles are also called voluntary muscles
as they come under the control of the nervous system in the body.
Also Read: Difference between Voluntary and Involuntary Muscles
Table of Contents
• Structure
• Properties
• Types
• Functions
• Smooth Muscle
• Cardiac Muscle
Structure Of Skeletal Muscle
The structure of skeletal muscles is shown below-
This muscle is attached to the bones by an elastic tissue or collagen fibres called tendons.
These tendons are comprised of connective tissues. The skeletal muscles consist of a
bundle of muscle fibres namely fascicule. These fascicules are cylindrical in shape as
shown in the figure. These muscle fibres are surrounded by blood vessels and a number
of layers of other tissues enclosing it.
Each muscle fibre is lined by plasma membrane namely sarcolemma reticulum. It
encloses a cytoplasm called sarcoplasm which has the endoplasmic reticulum. The
muscle fibres consist of myofibrils, which have two important proteins, namely actin and
myosin in it. The fascicule is enclosed by perimysium and the endomysium is the
connective tissue that encloses the muscle fibres.
Also Refer: Sliding Filament Theory
Properties Of Skeletal Muscle
The skeletal muscles have the following properties:
• Extensibility: It is the ability of the muscles to extend when it is stretched.
• Elasticity: It is the ability of the muscles to return to its original structure when
released.
• Excitability: It is the ability of the muscle to respond to a stimulus.
• Contractility: It is the ability of a muscle to contract when in contact with a
stimulus.
Types Of Skeletal Muscle
There are two types of skeletal muscles named as red and white muscles-
• Red Muscles
Red muscles are due to the red pigment called myoglobin, which is in high amounts in
the human body. These muscles are smaller in diameter and have a large number of
mitochondria in it. The myoglobin stores the oxygen, which is used by the mitochondria
for the synthesis of ATP. Red muscles have a large number of blood capillaries in it.
• White Muscles
Unlike the red muscles, the white muscles are bigger in diameter and have a small
amount of myoglobin in it. They also have less number of mitochondria in it.
Also Refer: Muscular System
Functions Of Skeletal Muscle
Following are the important skeletal muscle function:
1. The skeletal muscles are responsible for body movements such as typing,
breathing, extending the arm, writing, etc. The muscles contract which pulls the
tendons on the bones and causes movement.
2. The body posture is maintained by the skeletal muscles. The gluteal muscle is
responsible for the erect posture of the body. The Sartorius muscles in thighs are
responsible for body movement.
3. The skeletal muscles protect the internal organs and tissues from any injury and
also provide support to these delicate organs and tissues.
4. These also support the entry and exit points of the body. The sphincter muscles
are present around the anus, mouth and the urinary tract. These muscles
contract which reduces the size of the openings and facilitates the swallowing of
food, defecation, and urination.
5. The skeletal muscles also regulate body temperature. After a strenuous exercise,
the body feels hot. This is due to the contraction of skeletal muscles which
converts energy into heat.
Smooth Muscle
Smooth muscle is a type of muscle tissue that is utilized by different systems for the
application of pressure to organs and vessels. It is an involuntary muscle, which shows
no cross stripes even when examined under a microscope.
A smooth muscle is composed of cells that are narrow and spindle-shaped with a single
nucleus that is located centrally.
The cells of smooth muscles are made up of fibres of myosin and actin that run through
the cells and are backed by a framework of various proteins, wherein the filaments are
arranged in a stacked pattern across the cell.
It exhibits a ‘staircase’ arrangement of the fibres-actin and myosin, which is typically
different from the structure observed in cardiac and skeletal muscles.
Unlike the striated muscles, the smooth muscle tissues contract at a slower pace
automatically. Most of the musculature of the digestive system and the internal organs
are composed of the smooth muscles.
Upon the release of ATP to be used by myosin, the smooth muscles contract under the
influence of stimuli. This ATP that is released relies on the magnitude of the stimuli
which enables the smooth muscles to possess a graded contraction in contrast to the
‘on-or-off’ contraction observed in the skeletal muscles. The actin filaments run from
one side of the cell to the other which is connected at the cell membrane and the dense
bodies.
Cardiac Muscle
One of the three types of muscles, the cardiac muscles is a muscle tissue found in the
heart, wherein it is performing and bringing about coordinated contractions, which
enable the heart to pump blood throughout the body through the circulatory system.
The cardiac muscle tissues function to cause continuous pumping of the heart through
involuntary movements. This is one of the striking features of cardiac muscles that sets
it apart from the muscle tissues which is under one’s control. It is able to do so through
specialized cells known as the pacemaker cells which govern the heart’s contractions.
The nervous system transmits signals to the specialized cells, which indicates them to
increase or decrease the heart rate. The pacemaker cells are linked to the cardiac
muscle cells, which enables them to transmit signals which cause a wave of
contractions in the cardiac muscles that in turn generates a heartbeat.
The cardiac muscles are composed of the following:
• Nucleus
• Gap junctions
• Intercalated discs
• Desmosomes
Each skeletal muscle is an organ that consists of various integrated tissues. These tissues include
the skeletal muscle fibers, blood vessels, nerve fibers, and connective tissue. Each skeletal
muscle has three layers of connective tissue (called mysia) that enclose it, provide structure to
the muscle, and compartmentalize the muscle fibers within the muscle (Figure 10.2.1). Each
muscle is wrapped in a sheath of dense, irregular connective tissue called the epimysium, which
allows a muscle to contract and move powerfully while maintaining its structural integrity. The
epimysium also separates muscle from other tissues and organs in the area, allowing the muscle
to move independently.
Figure 10.2.1 – The Three Connective Tissue Layers: Bundles of muscle fibers, called
fascicles, are covered by the perimysium. Muscle fibers are covered by the endomysium.
Inside each skeletal muscle, muscle fibers are organized into bundles, called fascicles,
surrounded by a middle layer of connective tissue called the perimysium. This fascicular
organization is common in muscles of the limbs; it allows the nervous system to trigger a
specific movement of a muscle by activating a subset of muscle fibers within a fascicle of the
muscle. Inside each fascicle, each muscle fiber is encased in a thin connective tissue layer of
collagen and reticular fibers called the endomysium. The endomysium surrounds
the extracellular matrix of the cells and plays a role in transferring force produced by the muscle
fibers to the tendons.
In skeletal muscles that work with tendons to pull on bones, the collagen in the three connective
tissue layers intertwines with the collagen of a tendon. At the other end of the tendon, it fuses
with the periosteum coating the bone. The tension created by contraction of the muscle fibers is
then transferred though the connective tissue layers, to the tendon, and then to the periosteum to
pull on the bone for movement of the skeleton. In other places, the mysia may fuse with a broad,
tendon-like sheet called an aponeurosis, or to fascia, the connective tissue between skin and
bones. The broad sheet of connective tissue in the lower back that the latissimus dorsi muscles
(the “lats”) fuse into is an example of an aponeurosis.
Every skeletal muscle is also richly supplied by blood vessels for nourishment, oxygen delivery,
and waste removal. In addition, every muscle fiber in a skeletal muscle is supplied by the axon
branch of a somatic motor neuron, which signals the fiber to contract. Unlike cardiac and smooth
muscle, the only way to functionally contract a skeletal muscle is through signaling from the
nervous system.
Skeletal Muscle Fibers
Because skeletal muscle cells are long and cylindrical, they are commonly referred to as muscle
fibers (or myofibers). Skeletal muscle fibers can be quite large compared to other cells, with
diameters up to 100 μm and lengths up to 30 cm (11.8 in) in the Sartorius of the upper leg.
Having many nuclei allows for production of the large amounts of proteins and enzymes needed
for maintaining normal function of these large protein dense cells. In addition to nuclei, skeletal
muscle fibers also contain cellular organelles found in other cells, such as mitochondria and
endoplasmic reticulum. However, some of these structures are specialized in muscle fibers. The
specialized smooth endoplasmic reticulum, called the sarcoplasmic reticulum (SR), stores,
releases, and retrieves calcium ions (Ca++).
The plasma membrane of muscle fibers is called the sarcolemma (from the Greek sarco, which
means “flesh”) and the cytoplasm is referred to as sarcoplasm (Figure 10.2.2). Within a muscle
fiber, proteins are organized into organelles called myofibrils that run the length of the cell and
contain sarcomeres connected in series. Because myofibrils are only approximately 1.2 μm in
diameter, hundreds to thousands (each with thousands of sarcomeres) can be found inside one
muscle fiber. The sarcomere is the smallest functional unit of a skeletal muscle fiber and is a
highly organized arrangement of contractile, regulatory, and structural proteins. It is the
shortening of these individual sarcomeres that lead to the contraction of individual skeletal
muscle fibers (and ultimately the whole muscle).
Figure 10.2.2 – Muscle Fiber: A skeletal muscle fiber is surrounded by a plasma membrane
called the sarcolemma, which contains sarcoplasm, the cytoplasm of muscle cells. A muscle fiber
is composed of many myofibrils, which contain sarcomeres with light and dark regions that give
the cell its striated appearance.
The Sarcomere
A sarcomere is defined as the region of a myofibril contained between two cytoskeletal
structures called Z-discs (also called Z-lines or Z-bands), and the striated appearance of skeletal
muscle fibers is due to the arrangement of the thick and thin myofilaments within each
sarcomere (Figure 10.2.2). The dark striated A band is composed of the thick filaments
containing myosin, which span the center of the sarcomere extending toward the Z-dics. The
thick filaments are anchored at the middle of the sarcomere (the M-line) by a protein called
myomesin. The lighter I band regions contain thin actin filaments anchored at the Z-discs by a
protein called α-actinin. The thin filaments extend into the A band toward the M-line and
overlap with regions of the thick filament. The A band is dark because of the thicker myosin
filaments as well as overlap with the actin filaments. The H zone in the middle of the A band is a
little lighter in color because it only contain the portion of the thick filaments that does not
overlap with the thin filaments (i.e. the thin filaments do not extend into the H zone).
Because a sarcomere is defined by Z-discs, a single sarcomere contains one dark A band with
half of the lighter I band on each end (Figure 10.2.2). During contraction the myofilaments
themselves do not change length, but actually slide across each other so the distance between the
Z-discs shortens resulting in the shortening of the sarcomere. The length of the A band does not
change (the thick myosin filament remains a constant length), but the H zone and I band regions
shrink. These regions represent areas where the filaments do not overlap, and as filament
overlap increases during contraction these regions of no overlap decrease.
Myofilament Components
The thin filaments are composed of two filamentous actin chains (F-actin) comprised of
individual actin proteins (Figure 10.2.3). These thin filaments are anchored at the Z-disc and
extend toward the center of the sarcomere. Within the filament, each globular actin monomer
(G-actin) contains a myosin binding site and is also associated with the regulatory proteins,
troponin and tropomyosin. The troponin protein complex consists of three
polypeptides. Troponin I (TnI) binds to actin, troponin T (TnT) binds to tropomyosin, and
troponin C (TnC) binds to calcium ions. Troponin and tropomyosin run along the actin filaments
and control when the actin binding sites will be exposed for binding to myosin.
Thick myofilaments are composed of myosin protein complexes, which are composed of six
proteins: two myosin heavy chains and four light chain molecules. The heavy chains consist of a
tail region, flexible hinge region, and globular head which contains an Actin-binding site and a
binding site for the high energy molecule ATP. The light chains play a regulatory role at the
hinge region, but the heavy chain head region interacts with actin and is the most important
factor for generating force. Hundreds of myosin proteins are arranged into each thick filament
with tails toward the M-line and heads extending toward the Z-discs.
Other structural proteins are associated with the sarcomere but do not play a direct role in active
force production. Titin, which is the largest known protein, helps align the thick filament and
adds an elastic element to the sarcomere. Titin is anchored at the M-Line, runs the length of
myosin, and extends to the Z disc. The thin filaments also have a stabilizing protein, called
nebulin, which spans the length of the thick filaments.
Figure 10.2.3 – The Sarcomere: The sarcomere, the region from one Z-disc to the next Z-disc,
is the functional unit of a skeletal muscle fiber.
The Sliding Filament Model of Contraction
The arrangement and interactions between thin and thick filaments allows for the sarcomeres to
generates force. When signaled by a motor neuron, a skeletal muscle fiber is activated. Cross
bridges form between the thick and thin filaments and the thin filaments are pulled which slide
past the thick filaments within the fiber’s sarcomeres. It is important to note that while the
sarcomere shortens, the individual proteins and filaments do not change length but simply slide
next to each other. This process is known as the sliding filament model of muscle contraction
(Figure 10.2.4).
Figure 10.2.4 – The Sliding Filament Model of Muscle Contraction: When a sarcomere
shortens, the Z-discs move closer together, and the I band becomes smaller. The A band stays the
same width. At full contraction, the thin and thick filaments have the most amount of overlap.
The filament sliding process of contraction can only occur when myosin-binding sites on the
actin filaments are exposed by a series of steps that begins with Ca++ entry into the
sarcoplasm. Tropomyosin winds around the chains of the actin filament and covers the myosin-
binding sites to prevent actin from binding to myosin. The troponin-tropomyosin complex uses
calcium ion binding to TnC to regulate when the myosin heads form cross-bridges to the actin
filaments. Cross-bridge formation and filament sliding will occur when calcium is present, and
the signaling process leading to calcium release and muscle contraction is known as Excitation-
Contraction Coupling.
Sarcoplasmic reticulum
A cartoon section of skeletal muscle, showing T-tubules running deep into the centre of
the cell between two terminal cisternae/junctional SR. The thinner projections, running
horizontally between two terminal cisternae are the longitudinal sections of the SR.
The sarcoplasmic reticulum (SR) is a membrane-bound structure found within muscle
cells that is similar to the smooth endoplasmic reticulum in other cells. The main
function of the SR is to store calcium ions (Ca2+). Calcium ion levels are kept relatively
constant, with the concentration of calcium ions within a cell being 10,000 times smaller
than the concentration of calcium ions outside the cell.This means that small increases
in calcium ions within the cell are easily detected and can bring about important cellular
changes (the calcium is said to be a second messenger. Calcium is used to
make calcium carbonate (found in chalk) and calcium phosphate, two compounds that
the body uses to make teeth and bones. This means that too much calcium within the
cells can lead to hardening (calcification) of certain intracellular structures, including
the mitochondria leading to cell death. Therefore, it is vital that calcium ion levels are
controlled tightly, and can be released into the cell when necessary and then removed
from the cell.
Structure
The sarcoplasmic reticulum is a network of tubules that extend throughout muscle cells,
wrapping around (but not in direct contact with) the myofibrils (contractile units of the
cell). Cardiac and skeletal muscle cells contain structures called transverse tubules (T-
tubules), which are extensions of the cell membrane that travel into the centre of the
cell. T-tubules are closely associated with a specific region of the SR, known as
the terminal cisternae in skeletal muscle, with a distance of roughly 12 nanometers,
separating them. This is the primary site of calcium release.The longitudinal SR are
thinner projects, that run between the terminal cisternae/junctional SR, and are the
location where ion channels necessary for calcium ion absorption are most
abundant.[4] These processes are explained in more detail below and are fundamental
for the process of excitation-contraction coupling in skeletal, cardiac and smooth
muscle.
Calcium absorption.
The SR contains ion channel pumps, within its membrane that are responsible for
pumping Ca2+ into the SR. As the calcium ion concentration within the SR is higher than
in the rest of the cell, the calcium ions won't freely flow into the SR, and therefore
pumps are required, that use energy, which they gain from a molecule called adenosine
triphosphate (ATP). These calcium pumps are called Sarco(endo)plasmic reticulum
Ca2+ ATPases (SERCA). There are a variety of different forms of SERCA, with SERCA
2a being found primarily in cardiac and skeletal muscle.
SERCA consists of 13 subunits (labelled M1-M10, N, P and A). Calcium ions bind to the
M1-M10 subunits (which are located within the membrane), whereas ATP binds to the
N, P and A subunits (which are located outside the SR). When 2 calcium ions, along
with a molecule of ATP, bind to the cytosolic side of the pump (i.e. the region of the
pump outside the SR), the pump opens. This occurs because ATP (which contains
three phosphate groups) releases a single phosphate group (becoming adenosine
diphosphate). The released phosphate group then binds to the pump, causing the pump
to change shape. This shape change causes the cytosolic side of the pump to open,
allowing the two Ca2+ to enter. The cytosolic side of the pump then closes and the
sarcoplasmic reticulum side opens, releasing the Ca 2+ into the SR.[6]
A protein found in cardiac muscle, called phospholamban (PLB) has been shown to
prevent SERCA from working. It does this by binding to the SERCA and decreasing its
attraction (affinity) to calcium, therefore preventing calcium uptake into the SR. Failure
to remove Ca2+ from the cytosol, prevents muscle relaxation and therefore means that
there is a decrease in muscle contraction too. However, molecules such
as adrenaline and noradrenaline, can prevent PLB from inhibiting SERCA. When these
hormones bind to a receptor, called a beta 1 adrenoceptor, located on the cell
membrane, they produce a series of reactions (known as a cyclic AMP pathway) that
produces an enzyme called protein kinase A (PKA). PKA can add a phosphate to PLB
(this is known as phosphorylation), preventing it from inhibiting SERCA and allowing for
muscle relaxation.[7]
Calcium storage
Located within the SR is a protein called calsequestrin. This protein can bind to around
50 Ca2+, which decreases the amount of free Ca2+ within the SR (as more is bound to
calsequestrin).[8] Therefore, more calcium can be stored (the calsequestrin is said to be
a buffer). It is primarily located within the junctional SR/luminal space, in close
association with the calcium release channel (described below).[9]
Calcium release
Calcium ion release from the SR, occurs in the junctional SR/terminal cisternae through
a ryanodine receptor (RyR) and is known as a calcium spark.
There are three types of ryanodine receptor, RyR1 (in skeletal
muscle), RyR2 (in cardiac muscle) and RyR3 (in the brain).[11] Calcium release through
ryanodine receptors in the SR is triggered differently in different muscles. In cardiac and
smooth muscle an electrical impulse (action potential) triggers calcium ions to enter the
cell through an L-type calcium channel located in the cell membrane (smooth muscle) or
T-tubule membrane (cardiac muscle). These calcium ions bind to and activate the RyR,
producing a larger increase in intracellular calcium. In skeletal muscle, however, the L-
type calcium channel is bound to the RyR. Therefore, activation of the L-type calcium
channel, via an action potential, activates the RyR directly, causing calcium release
(see calcium sparks for more details). Also, caffeine (found in coffee) can bind to and
stimulate RyR. Caffeine makes the RyR more sensitive to either the action potential
(skeletal muscle) or calcium (cardiac or smooth muscle), thereby producing calcium
sparks more often (this is partially responsible for caffeine's effect on heart rate).
Triadin and Junctin are proteins found within the SR membrane, that are bound to the
RyR. The main role of these proteins is to anchor calsequestrin (see above) to the
ryanodine receptor. At ‘normal’ (physiological) SR calcium levels, calsequestrin binds to
the RyR, Triadin and Junctin, which prevents the RyR from opening.[14] If calcium
concentration within the SR falls too low, there will be less calcium bound to the
calsequestrin. This means that there is more room on the calsequestrin, to bind to the
junctin, triadin and ryanodine receptor, therefore it binds tighter. However, if calcium
within the SR rises too high, more calcium binds to the calsequestrin and therefore it
binds to the junctin-triadin-RyR complex less tightly. The RyR can therefore open and
release calcium into the cell.[15]
In addition to the effects that PKA had on phospholamban (see above) that resulted in
increased relaxation of the cardiac muscle, PKA (as well as another enzyme
called calmodulin kinase II) can also phosphorylate ryanodine receptors. When
phosphorylated, RyRs are more sensitive to calcium, therefore they open more often
and for longer periods of time. This increases calcium release from the SR, increasing
the rate of contraction.[16] Therefore, in cardiac muscle, activation of PKA, through
the cyclic AMP pathway, results in increased muscle contraction
(via RyR2 phosphorylation) and increased relaxation
(via phospholamban phosphorylation), which increases heart rate.
The mechanism behind the termination of calcium release through the RyR is still not
fully understood. Some researchers believe it is due to the random closing of ryanodine
receptors (known as stochastic attrition), or the ryanodine receptors becoming inactive
after a calcium spark,[17] while others believe that a decrease in SR calcium, triggers the
receptors to close (see calcium sparks for more details).
Role in rigor mortis
The breakdown of the sarcoplasmic reticulum, along with the resultant release of
calcium, is an important contributor to rigor mortis, the stiffening of muscles after death.
If the concentration of calcium increases in the sarcoplasm then it can also cause
muscles stiffness.
• Isotonic and Isometric, Eccentric
• Muscle contractions are defined by the changes in the length of the muscle
during contraction.
• Muscle contractions can be described based on two variables:
• force and length.
• Force itself can be differentiated as either tension or load.
• Muscle tension is the force exerted by the muscle on an object whereas a load is
the force exerted by an object on the muscle.
• Muscle fiber generates tension through the action of actin and myosin cross-
bridge cycling. While under tension, the muscle may lengthen, shorten, or remain
the same. Although the term contraction implies shortening, when referring to the
muscular system, it means muscle fibers generating tension with the help of
motor neurons. Several types of muscle contractions occur and they are defined
by the changes in the length of the muscle during contraction.
• When muscle tension changes without any corresponding changes in muscle
length, the muscle contraction is described as isometric. If the muscle length
changes while muscle tension remains the same, then the muscle contraction is
isotonic. In an isotonic contraction, the muscle length can either shorten to
produce a concentric contraction or lengthen to produce an eccentric
contraction. In natural movements that underlie locomotor activity, muscle
contractions are multifaceted as they are able to produce changes in length and
tension in a time-varying manner.[7] Therefore, neither length nor tension is likely
to remain constant when the muscle is active during locomotor activity.
• Isotonic Contractions
• Isotonic contractions maintain constant tension in the muscle as the muscle
changes length. This can occur only when a muscle’s maximal force of
contraction exceeds the total load on the muscle. Isotonic muscle contractions
can be either concentric (muscle shortens) or eccentric (muscle lengthens).
• Concentric Contractions
• A concentric contraction is a type of muscle contraction in which the muscles
shorten while generating force. This is typical of muscles that contract due to the
sliding filament mechanism, and it occurs throughout the muscle. Such
contractions also alter the angle of the joints to which the muscles are attached,
as they are stimulated to contract according to the sliding filament mechanism.
• This occurs throughout the length of the muscle, generating force at the musculo-
tendinous junction; causing the muscle to shorten and the angle of the joint to
change. For instance, a concentric contraction of the biceps would cause the arm
to bend at the elbow as the hand moves from near to the leg to close to the
shoulder (a biceps curl). A concentric contraction of the triceps would change the
angle of the joint in the opposite direction, straightening the arm and moving the
hand toward the leg.
• Eccentric Contractions
• An eccentric contraction results in the elongation of a muscle. Such contractions
decelerate the muscle joints (acting as “brakes” to concentric contractions) and
can alter the position of the load force. These contractions can be both voluntary
and involuntary. During an eccentric contraction, the muscle elongates while
under tension due to an opposing force which is greater than the force generated
by the muscle. Rather than working to pull a joint in the direction of the muscle
contraction, the muscle acts to decelerate the joint at the end of a movement or
otherwise control the repositioning of a load.
• This can occur involuntarily (when attempting to move a weight too heavy for the
muscle to lift) or voluntarily (when the muscle is “smoothing out” a movement).
Over the short-term, strength training involving both eccentric and concentric
contractions appear to increase muscular strength more than training with
concentric contractions alone.
• Isometric Contractions
• In contrast to isotonic contractions, isometric contractions generate force without
changing the length of the muscle . This is typical of muscles found in the hands
and forearm: the muscles do not change length, and joints are not moved, so
force for grip is sufficient. An example is when the muscles of the hand and
forearm grip an object; the joints of the hand do not move, but muscles generate
sufficient force to prevent the object from being dropped.
