ANA205

MUSCULAR TISSUE
Lecture Notes
BY DR. TAIWO-OLA
Introduction: Terminology
• Myofiber or Myocyte: a muscle cell
• Sarcolemma: the plasma membrane of a muscle cell
• Sarcoplasm: the cytoplasm of the muscle cell
• Sarcoplasmic reticulum: the endoplasmic reticulum of a muscle
cell
• Sarcosome: the mitochondria of a muscle cell
• Sarcomere: the contractile or functional unit of muscle
Structure of Muscle
Introduction
The functions of muscles are:
• contraction for locomotion and skeletal movement
• contraction for propulsion
• contraction for pressure regulation

Classification of Muscle:

Muscle tissue may be classified according to a morphological

classification or a functional classification.

1. Morphological classification (based on structure)

There are two types of muscle based on the morphological

classification system
• Striated
• Non striated or smooth.

2. Functional classification

There are two types of muscle based on a functional classification

Types of muscle:
There are generally considered to be three types of
muscle in the human body.
• Skeletal muscle: which is striated and voluntary
• Cardiac muscle: which is striated and involuntary
• Smooth muscle: which is non striated and involuntary
 Skeletal Muscle
Contractions move part of the skeleton. Also called 'voluntary'
because usually its contractions are under your control.

Skeletal muscle has a stripy appearance, because of the repeating

structure of the muscle: there are many myofibrils (fibers), each one of
which is made up of repeating units called muscle sarcomeres.
Each sarcomere is 2.5 mm long.

Skeletal muscle moves joints by strong and rapid contractions.

Each muscle is a bundle of muscle fibres, each of which is a long
multinucleated cell.

Single mononucleated cells called myoblasts fuse together to form

this multinucleated fibre (myotube) during development. This process
also occurs to regenerate muscle fibres.
If you cut a muscle longitudinally and stain it (H&E), then
you can see long muscle fibres, that look stripy. Have a
look at this H&E stained section and notice the stripes.
The dark purple nuclei can be seen on the edges of the
muscle fibres.
This picture shows a transverse section through skeletal
muscle.
If you cut a muscle transversely, like this, then you can
see the roughly circular outline of the individual muscle
fibres.
Cardiac Muscle
Cardiac muscle makes up the muscular walls of
the heart (myocardium). It is 'involuntary' because its
contractions are not under your control. However, it has
a similar ultrastructural organisation to skeletal muscle.
So, it too has a stripy appearance because of the
repeating units called muscle sarcomeres.
Cardiac muscle is striated, like skeletal muscle, as
the actin and myosin are arranged in sarcomeres, just
as in skeletal muscle. However, cardiac muscle
is involuntary.
• Cardiac muscle cells usually have a single (central)
nucleus. The cells are often branched, and are
tightly connected by specialised junctions. The
region where the ends of the cells are connected to
another cell is called an intercalated disc.
• The intercalated disc contains gap
junctions, adhering junctions and desmosomes. Gap
junctions allow the muscle cells to be electrically
coupled, so that they beat in synchrony.
This is a low power section through the wall of the heart,
showing the striated appearance of the muscle, and the
nuclei.
What you can't see very clearly here are the intercalated
discs, which are shown in a higher power image next
slide.
Here, in this higher magnification picture (compare the
size of the scale bar with that in the picture above),
you can now see individual cardiac muscle cells, that
branch. The densely staining regions at the ends of the
muscle fibres are intercalated discs.
Skeletal and Cardiac Muscle
Ultrastructure

This is a high power, light micrograph of a muscle fibre

showing the banding pattern. There are light stripes - which
are called the 'Z' lines, and darker wider stripes called the 'A'
bands. (A - for anisotropic - because in a polarizing light
microscope, the dark bands are birefringent). The Z-lines are
midway inside a light band, called the 'I' band. (I for isotropic
- because in a polarising microscope, these bands are much
less birefringent than the A bands).
The muscle sarcomere is the repeating unit (S) between
two Z-lines.
N - is a nucleus on the outside of the fibre.
This is an electron micrograph (EM) of a skeletal muscle fibre. (The
sarcomeres of cardiac muscle have a very similar organisation).
Notice how the stripes appear less regular than in the light
microscope. This is because the repeating muscle sarcomeres are
arranged in longitudinal structures called 'myofibrils' (from top right
to bottom left of the picture).
This is the same EM with labels to show the organisation of the
muscle sarcomeres. Z-lines mark the boundaries of the sarcomeres.
The dark staining region in the centre of the sarcomere is called the A
(anisotropic) band. The lighter staining band, through which the Z-line
passes is called the I (isotropic) band.
A diagram of a muscle sarcomere:
 Muscle sarcomere
• Thin filaments, which consist mostly of the proteins actin, troponin and
tropomyosin, insert directly into the Z-lines.
• Thick filaments consist mostly of the proteins myosin and titin. Titin
connects the thick filaments to the Z-line.
• There are about 300 molecules of myosin in each thick filament, and at
the end of each molecule are globular heads that stick out and bind to
actin in the thin filament. When the muscle is stimulated, the heads bind
to actin, and pull on actin to pull the thin filaments towards the M-line,
increasing the overlap between thick and thin filaments, and making the
muscle shorten. This uses ATP for energy.
• In relaxed muscle, troponin/tropomyosin stops myosin from binding to
actin. When muscle is stimulated, calcium is released, binds to troponin,
and this then allows myosin to bind to actin.
 Smooth Muscle
Found in the walls of most blood vessels and tubular
organs such as the intestine. It is also 'involuntary'.
However, it does NOT have a stripy appearance, because
it does not have repeating sarcomeres. The contractile
proteins, myosin and actin are much more randomly
arranged than in skeletal or cardiac muscle. Smooth
muscle is made up of cells that contain a single central
nucleus. The cells stick together and are connected by
specialised cell junctions, called gap junctions. The cells
are spindle shaped, and the nucleus is central.
Look at this section of smooth muscle, which shows smooth muscle cells both
in longitudinal section (LS) and in transverse section (TS). Compare this to
the diagram above, and make sure you can recognise smooth muscle cells in
TS and LS.
The cells do not have a striated appearance. This is because the actin and
myosin in these muscles is more randomly arranged.
Look at this scanning EM of smooth muscle
from the urinary bladder of a guinea pig. Scale
bar: 50 mm. It shows bundles of smooth
muscle fibres.
 There are many functions of smooth muscle,
from moving food along the digestive tract, to pulling
hair erect in response to cold or fear.
Single smooth muscle cells are often found
surrounding ducts, or blood vessels, lying within the
basement membrane - i.e. they are part of the
epithelial layer. These are called myo-epithelial cells.
When these cells contract, they squeeze the ducts,
helping to extrude the contents.
Look at this section of ducts, that are surrounded by myo-epithelial
cells.
The myoepithelial cells are very thin, and have dense flattened
nuclei. The position of the myoepithelial cells around the duct is
outlined in the diagram below.
 Cell Junctions
There are several kinds of cell-cell junctions.
Cardiac cells are special, amongst the muscle types,
because they are connected to each other by intercalated discs -
structures that are only found in cardiac muscle cells. These can
be seen in this diagram, as darkly staining irregular lines, at 90
degrees to the striped sarcomeric pattern.

Skeletal muscle does not have any cell-cell junctions.

Smooth muscle contains gap junctions, to allow a

rapid spread of depolarisation, as in cardiac muscle.

 Intercalated discs contain three different types of

cell-cell junctions:

• Fascia adherens junctions (anchoring junctions)

where actin filaments attach thin filaments in the muscle

sarcomeres to the cell membrane.

• Expanded desmosomes, sites of strong adhesion, that help

to keep the muscle cells connected when they contract.

• Gap junctions, large and small, which provide direct contact

between the cardiac cells, facilitating electrical

communication, so that waves of depolarisation spread

rapidly over the entire heart, by passing from cell to cell.

Muscle Regeneration
• Skeletal muscle contains numerous 'satellite cells'
underneath the basal lamina, as shown in the
photograph opposite. These are mononucleated
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.
The skeletal muscle fibres themselves, cannot divide.
However, muscle fibres can lay down new protein and
enlarge (hypertrophy).
• Cardiac muscle can also hypertrophy. However, there are
no equivalent to cells to the satellite cells found in skeletal
muscle. Thus when cardiac muscle cells die, they are not
replaced.

• Smooth cells have the greatest capacity to regenerate of

all the muscle cell types. The smooth muscle cells
themselves retain the ability to divide, and can increase in
number this way. As well as this, new cells can be produced
by the division of cells called pericytes that lie along some
small blood vessels. Smooth muscle can also hypertrophy.
Muscle: Stimulation
Skeletal muscle is stimulated via a nerve impulse, which
depolarises the muscle. However, not all the muscle fibres in
the muscle fibre will necessarily be activated at once.
Sometimes, a subset of muscle fibres is activated, depending on
how much force is needed.
When the muscle is stimulated, calcium ions are released
from its store inside the sarcoplasmic reticulum, into the
sarcoplasm (muscle ). Then the calcium ions bind to a protein
called troponin, on the thin filaments, which in turn allows
myosin to bind to actin. This interaction makes the thick and
thin filaments slide past each other, to make the muscle
• Invaginations of the plasma membrane (sarcolemma) of the muscle

fibres are called T (or transverse) tubules. The T-tubules lie over
the junction between the A- and I-bands (see diagram).
• The two terminal cistemae of the SR together with their associated

T tubule are known as a triad.

• Inside the muscle fibre, the T-tubules lie next to the terminal

cisternae of an internal membrane system derived from the

endoplasmic reticulum, called the sarcoplasmic reticulum
(SR), which is a store of calcium ions. Stimulation of the muscle
fibre, causes a wave of depolarisation to pass down the t-tubule, and
the SR to release calcium ions into the sarcoplasm. Calcium is
pumped back up into the SR to lower calcium ion concentration in
the sarcoplasm, to relax the muscle (turn off contraction).
• Cardiac Muscle also has T-tubules, and SR. However the T-

tubules lie over the Z-line in cardiac muscle, are less

numerous and wider. The SR is smaller and less elaborate,

and stores less calcium ions. Cardiac muscle cells also

depend on extracellular calcium ions, that enter through the

T-tubules and triggers release of calcium ions from the SR.

• Cardiac muscle cells are electrically connected through gap

junctions, so that waves of electrical stimuli pass around the

heart from cell to cell, and all the cells are stimulated to

contract.
• Smooth Muscle. The thick and thin filaments are
attached to alpha-actinin in dense bodies (equivalent
to Z-lines in skeletal muscle), which are attached to
the plasma membrane by intermediate filaments. The
thin filaments do not have troponin. This type of
muscle responds to a increase in calcium, following
nerve stimulation through a protein
called calmodulin. Binding of calcium to calmodulin,
results in the activation of an enzyme (myosin light
chain kinase) that phosphorylates myosin, which
activates it, enabling it to interact with actin.
References
• Histology Guide © Faculty of Biological
Sciences, University of Leeds