CONTRACTION AND RELAXATION OF SKELETAL MUSCLE FIBERS

Sliding Filament Mechanism

- By A.F. Huxley and H.E. Huxley
- Active shortening is caused by the movement of the cross bridges
- In a muscle contraction actin filaments move while myosin filaments remain
static
- Muscle contraction occurs because myosin heads attach to and walk along
the thin filaments at both ends of the sarcomere pulling the thin filaments
towards the M line
o Result:
Thin filaments slide inward and meet at the center of a
sarcomere
I band and H zone narrow and disappear when muscle is
maximally contracted (as thin filaments move inward)
A band and length of thick and thin filaments does not change
Z discs comes closer together (since thin filaments are attached
to them)
SHORTENING of sarcomere then whole muscle

The Contraction Cycle

- The repeating sequence of events that causes the filaments to slide

Onset of contraction sarcoplasm releases calcium ions into the sarcoplasm

- Ca ions bind to troponin
- Troponin moves Tropomyosin away from the myosin-binding sites on actin
- Once binding sites are free CONTRACTION CYCLE begins

4 Steps of the Contraction Cycle

1. ATP HYDROLYSIS
- Myosin head contains an ATP-binding site and an ATPase (enzyme that
hydrolyzes ATP into ADP and a phosphate group)
- Hydrolysis reaction reorients and energizes the myosin head
- ADP and Pi still attached to the myosin head

2. Attachment of myosin to actin to form cross-bridges

- Energized myosin attaches to myosin-binding site on actin
- Release of hydrolyzed phosphate group
- Cross-bridges myosin heads attached to actin

3. Power stroke
- Occurs after cross-bridges forms
- The site on the cross-bridge where ADP is opens, rotates, and releases ADP
- Cross-bridge generates force as it rotates towards the center of the
sarcomere. Sliding the thin filament past the thick filament towards the M line
o Force draws the Z disc toward each other shortening of the
sarcomere Z disc pulls another sarcomere whole muscle fiber
shortens
 o maximal muscle contraction, the distance between two Z discs can
decrease to half the resting length

4. Detachment of myosin from actin

- Cross-bridge remains firmly attached to actin until it binds another molecule
of ATP
- As ATP binds at ATP-binding site on Myosin head, myosin detaches from actin

Contraction repeats and continues as long as ATP is available and the Ca2+ level
near thin filament is high.

Each of the 600 cross-bridges in one thick filament attaches and detaches about
five times per second

Some of the components of a muscle are elastic: They stretch slightly before they
transfer the tension generated by the sliding filaments
- Elastic components include:
o titin molecules
o connective tissue around the muscle fibers (endomysium, perimysium,
and epimysium)
o tendons

As cells of a skeletal muscle starts to shorten

- first full on their connective tissue coverings and then tendons
- coverings and tendons stretch and then become taut
- tension passed through the tendons pulls on the bones to which they are
attached
- movement of a part of the body

Contraction Cycle does not always result in shortening of muscle fibers

- in some, cross-bridges rotate and generate tension, but thin filaments cannot
slide inward because tension not enough
- ex: trying to lift a whole box of books with one hand

Excitation-contraction Coupling
- Events in this coupling that connect excitation (a muscle action potential
propagating along the sarcolemma and into the T tubules) to contraction
(sliding of the filaments):
o Muscle action potential propagates along sarcolemma then into T
tubules
o Ca2+ release channels in the SR membrane to open
o Ca2+ flows out of the SR into the sarcoplasm around the thick and thin
filaments
o Ca2+ concentration in the sarcoplasm rises tenfold or more
o calcium ions combine with troponin, causing it to change shape
o conformational change moves tropomyosin away from the myosin-
binding sites on actin
o myosin heads bind to them to form cross-bridges
o contraction cycle begins
- Ca2+ concentration in sarcoplasm
o Increase contraction
o Decrease relaxation
0.1 micromole per liter
o Normally, large amount of Ca2+ is stored in the Terminal Cistern of the
Sarcoplasmic Reticulum

Ca2+ active transport pumps

o Contained in SR that use ATP to move Ca2+ constantly from the
sarcoplasm into the SR
o Calcium ions flow into the sarcoplasm more rapidly than they are
transported back by the pumps
Calsequestrin
o calcium-binding protein inside the SR
o enabling more Ca3+ to be stored in the SR