Contraction of skeletal muscle

Learning objectives
What evidence supports the sliding
filament mechanism of muscle
contraction?
How does the sliding filament mechanism
cause a muscle to contract and relax?
Where does the energy for muscle
contraction come from?

Muscle Contraction
Knowing the structure of the sarcomere enables
us to understand what happens when a muscle
contracts. The mechanism of muscle contraction
can be deduced by comparing electron
micrographs of relaxed and contracted muscle:

The Sliding Filament Theory

These show that each sarcomere gets shorter
when the muscle contracts, so the whole muscle
gets shorter. But the dark band, which
represents the thick filament, does not change in
length. This shows that the filaments dont
contract themselves, but instead they slide past
each other.

The Sliding Filament Theory

What makes the filaments slide past each other?
Energy is provided by the splitting of ATP, and
the ATPase that does this splitting is located in
the myosin cross bridge head. These cross
bridges can also attach to actin, so they are able
to cause the filament sliding by "walking" along
the thin filament. This cross bridge walking is
called the cross bridge cycle, and it has 4 steps.
One step actually causes the sliding, while the
other 3 simply reset the cross bridge back to its
starting state.

The Sliding Filament Theory

1. The cross bridge swings out from the thick filament and
attaches to the thin filament. [Put oars in water.]
2. The cross bridge changes shape and rotates through 45,
causing the filaments to slide. The energy from ATP splitting is
used for this "power stroke" step, and the products (ADP + Pi)
are released. [Pull oars to drive boat through water.]
3. A new ATP molecule binds to myosin and the cross bridge
detaches from the thin filament. [push oars out of water.]
4. The cross bridge changes back to its original shape, while
detached (so as not to push the filaments back again). It is now
ready to start a new cycle, but further along the thin filament.
[push oars into starting position.]
One ATP molecule is split by each cross bridge in each cycle,
which takes a few milliseconds. During a contraction, thousands
of cross bridges in each sarcomere go through this cycle
thousands of times. Fortunately the cross bridges are all out of
synch, so there are always many cross bridges attached at any
time to maintain the force.

How is the cross bridge cycle switched off in a

relaxed muscle?
This is where the regulatory proteins on the thin filament, troponin
and tropomyosin, are involved. Tropomyosin is a long thin molecule,
and it can change its position on the thin filament. In a relaxed muscle
it is on the outside of the filament, covering the actin molecules so
that myosin cross bridges cant attach. This is why relaxed muscle is
compliant: there are no connections between the thick and thin
filaments. In a contracting muscle the tropomyosin has moved into the
groove of the double helix, revealing the actin molecules and allowing
the cross bridges to attach.

Cross-section of actin filament

Here calcium has

bound to the
troponin to move
tropomysin and
allow the myosin
heads to bind.

Troponin
Tropomysin
Actin
Myosin head binding site

T-System
Transverse system (T-tubules) an infolding of
the sarcolemma (membrane surrounding muscle
fibre) that spread across the muscle fibre.
Sarcoplasmic Reticulum accumulates calcium
ions in a resting muscle.
The depolarisation of the muscle spread across
the sarcolemma and into the T-tubules and the
sarcoplasmic reticulum
Calcium ions are released into the sarcoplasm
(cytoplasm of muscle fibre)

The Neuromuscular Junction

1. A muscle is supplied with only one nerve
fibre
2. There is no threshold for stimulation if
an impulse arrives at the motor end plate
it will always stimulate the muscle fibre.

Controlling Contraction
1. An action potential arrives at the end of a motor neurone,
at the neuromuscular junction.
2. This causes the release of the neurotransmitter
acetylcholine.
3 This initiates an action potential in the muscle cell
membrane.
4. This action potential is carried quickly throughout the large
muscle cell by invaginations in the cell membrane called Ttubules.
5. The action potential causes the sarcoplasmic reticulum
(large membrane vesicles) to release its store of calcium into
the myofibrils.
6. The calcium binds to troponin on the thin filament, which
changes shape, moving tropomyosin into the groove in the
process.
7. Myosin cross bridges can now attach and the cross bridge
cycle can take place.

Neurotransmitter
= acetylcholine
Axon
terminal motor end
plate