Muscle Contraction:

Sliding Filament Theory

This mechanism is formulated by
Hauxley and Hauxley
Sliding Filament Theory

Muscle contraction occurs by a sliding filament

mechanism whereby the sarcomeres shorten (the Z-
lines come closer together) by the action of the actin
filaments sliding over the myosin filaments. Myosin
filaments may look somewhat like a golf club but
they are not inflexible. In fact, muscle contraction
would be impossible if the myosin molecules did not
have a "hinge" along the shaft that allows for a
ratchet movement of the head.
• The force behind muscle contraction is the ratchet
movement of  these tiny myosin
• heads toward the center of their sarcomere. This ratchet
movement occurs many times during a muscle
contraction.
• Electron microscopy combined with chemical
experiments show that muscle is composed of 2
contractile proteins:
• a) Thin filaments: actin, attached to Z line, found in both
A and I bands
• b) Thick filaments: myosin, found in A band
•
 When muscle contracts the actin filaments slide into the A
band, overlapping with myosin.
When muscle contracts:
a) the Z lines move closer together
b) the I band becomes shorter
  c) the A band stays at the same length
This is called the "sliding filament" model of muscle contraction.
The filaments slide together
because myosin attaches to actin and
pulls on it. Myosin head (H) attaches to actin
filament (A), forming a cross bridge.
After the cross bridge is formed the
myosin head bends, pulling on the actin
filaments and causing them to slide:
Muscle contraction is a little like
climbing a rope. The cross bridge cycle
is: grab -> pull -> release, repeated over
and over
• During muscle contraction the myofilaments myosin and
actin slide toward each other and overlap. This shortens
the sacromere and the entire muscle. Muscle cells are
"shocked" by nerve impulses from motor neurons.
• The point of attachment of the nerve to the muscle is
called a neuromuscular junction. A motor neuron and its
muscle cells are referred to as a motor unit. The nerve
impulse is carried from the neuron across the gap to the
membrane (sarcolemma) of the muscle cell by a chemical
called acetycholine.
•After the impulse is passed an enzyme called acetyl
cholinesterase "de-activates" acetylcholine, readying the
muscle for the next nerve impulse. Stimulation of the
muscle cell causes Ca++ ions to be released into the cell.
This binds with the actin filaments causing them to
expose active sites to the myosin cross bridges. The
cross bridges bind to the active sites, forming a new
molecular structure which causes the cross bridge to
bend toward the center, pulling the actin filament with it.
 
 
 
• Energy from ATP is used to break the
bond, straighten the cross bridge, and
allow the cross bridge to form a new bond
with another active site further down the
actin filament. This cycle continues until
the muscle contraction is complete. Then
ATP is used to cause active transport to
move the calcium ions out of the muscle
fiber causing relaxation of the muscle.
Role of Calcium ion and
Energy Source (ATP)
Role of Calcium ions
•A sudden inflow of Ca is the trigger for muscle
contraction. In the resting state the protein tropomyosin
winds around actin and covers the myosin binding sites.
The Ca binds to a second protein, troponin, and this
action causes the tropomyosin to be pulled to the side,
exposing the myosin binding sites. With the sites
exposed muscle will contract if ATP is present.
•Impulses conducted along the transverse tubules
stimulate the release of Ca++ from the sarcoplasmic
reticulum into the cytoplasm. Ca++ difuses toward the
myofibrils and causes contraction.
• Troponin and Tropoyosin are two regulatory proteins
associated with the thin filaments. Tropomyosin lies against
the thin filament and troponin is bound to tropomyosin. In a
resting muscle fiber, the concentration of Ca++ in the
cytoplasm is very low, and tropomyosin is located close to
the myosin-binding sites on the thin filament. In this position,
tropomyosin physically blocks the myosin heads from
binding actin, thus preventing contraction. In a stimulated
muscle fiber, the Ca++ released by the sarcoplasmic
reticulum binds to troponin. The Ca++ troponin complex
pulls the tropomyosin away from the myosin-binding sites on
actin, allowing cross-bridges to form.
• Cross-bridges cycles continue
as long as Ca++ remains
attached to troponin. When
nerve activity ceases,
impulses in the muscle fiber
also cease, and Ca++ is
actively transported from the
cytoplasm back into the
sarcoplasmic reticulum. As
Ca++ is released from
troponin, tropomyosin returns
to its inhibitory position on
the thin filament, again
preventing the myosin heads
from binding to actin. The
muscle fiber relaxes.
• Muscle contraction stops when Ca++ is
removed from the immediate environment
of the myofilaments. The sarcoplasmic
reticulum actively pumps Ca++ back into
itself and this requires utilization of ATP.
Troponin-tropomyosin reassume their
inhibitory position between the actin and
myosin molecules once Ca++ is removed.
Role of the energy source (ATP)
•ATP is the chief energy currency of all cells.The
bulk of the energy that plants harvest during
photosynthesis is channeled into production of
ATP, and so is most of the energy stored in fat
and starch. Cells use their supply of ATP to power
almost very energy-requiring process they carry
out, from supplying activation energy for
chemical reactions and actively transporting
substances across membranes, to moving through
their environment and growing. ATP as a small
unstable energy carrier that shuttles back and
forth within the cell, picking up energy in one
place and releasing it in another. ATP is so
important in all organisms.
» When a skeletal muscle is at rest, the
myosin heads function as enzymes,
cleaving ATP into ADP and P. The
hydrolysis of ATP activates the
myosin heads, putting them in an
orientation that allows them to bind
to specific sites on the actin of the
thin filaments when the muscle is
stimulated to contract.
» ATP is active transport, the movement of
substances across a membrane against their
concentration gradients. In this case, the
splitting of ATP activates a carrier protein
in the membrane, perhaps by changing its
shape so that it can transport a particular
molecule or ion across the membrane.
• Once the substance
has been released on
the other side, the
carrier protein returns
to its nonactived
shape, ready to
become energized by
another. ATP
molecule and shuttle
another molecule or
ion across the
membrane.
• ATP is the energy supply for contraction.
It is required for the sliding of the
filaments which is accomplished by a
bending movement of the myosin heads. It
is also required for the separation of actin
and myosin which relaxes the muscle.
When ATP runs down after death muscle
goes into a state of rigor mortis.
Role of Motor Units
Role of motor units

•The motor nerve and all the fibers it

innervates is called the motor unit. The
number of fibers is dependent on the
necessity for fine control. In general, small
muscles that react rapidly with fine control
have one nerve and only a  few muscle
fibers. Those muscles that do not require
fine control, such as the gastrocnemius (calf
muscle), may have several hundred muscle
fibers per motor unit.
• The contraction of individual muscle fibers is all-or-none.
Therefore, any graded response must come from the number
of motor units stimulated at any one time. Summation is the
adding together of individual muscle twitches to make a
whole muscle contraction. This can be accomplished by
increasing the number of motor units contracting at one time
(spatial summation) or by increasing the frequency of
contraction of individual muscle contractions (temporal
summation). These processes almost always occur
simultaneously within normal muscle contraction. Usually,
individual motor units fire asynchronously.
• All motor units are not created equal. Therefore one motor
unit within a particular muscle may be as much as 50 times as
strong as another.
Smaller motor units are much more easily excited than
larger ones because they are innervated by smaller
nerve fibers that have a naturally lower threshold for
excitation. In spatial summation motor units are
recruited by increasing the strength of the stimulus
thereby increasing the strength of the contraction.
•Motor unit is the smallest functional element of a
skeletal muscle. The division of the muscle into motor
units allows the muscle’s strength of contraction to be
finely graded, a requirement for coordinated
movements of the skeleton. Muscles that require a
finer degree of control have smaller motor units than
muscles that require less precise control but must exert
more force.
• The weakest contractions of a muscle are accomplished
by the activation of a few small motor units. If a slightly
stronger contraction is necessary. Additional small motor
units are also activated. The initial increments to the total
force generated by the muscle are therefore relatively
small. If ever greater forces are required, more and larger
motor units are brought into action, and the force
increments become larger. The use of increased numbers
and sizes of motor units in a contraction is termed
recruitment, and it is another way in which the strength of
muscle contraction is governed by the nervous system.