The Physiology of Skeletal Muscle Contraction
What is the Sliding Filament Theory of muscular contraction?
The sliding filament theory is the explanation for how muscles contract to produce force. As we have mentioned on previous pages, the actin and myosin filaments within the sarcomeres of muscle fibres bind to create cross-bridges and slide past one another, creating a contraction. The sliding filament theory explains how these cross-bridges are formed and the subsequent contraction of muscle.
The Sliding Filament Theory
For a contraction to occur there must first be a stimulation of the muscle in the form of an impulse (action potential) from a motor neuron (nerve that connects to muscle).
Note that one motor neuron does not stimulate the entire muscle but only a number of muscle fibres within a muscle.
The individual motor neuron plus the muscle fibres it stimulates, is called a motor unit. The motor end plate (also known as the neuromuscular junction) is the junction of the motor neurons axon and the muscle fibres it stimulates.
When an impulse reaches the muscle fibres of a motor unit, it stimulates a reaction in each sarcomere between the actin and myosin filaments. This reaction results in the start of a contraction and the sliding filament theory.
The reaction, created from the arrival of an impulse stimulates the 'heads' on the myosin filament to reach forward, attach to the actin filament and pull actin towards the centre of the sarcomere. This process occurs simultaneously in all sarcomeres, the end process of which is the shortening of all sarcomeres.
Troponin is a complex of three proteins that are integral to muscle contraction. Troponin is attached to the protein tropomyosin within the actin filaments, as seen in the image below. When the muscle is relaxed tropomyosin blocks the attachment sites for the myosin cross bridges (heads), thus preventing contraction.
When the muscle is stimulated to contract by the nerve impulse, calcium channels open in the sarcoplasmic reticulum (which is effectively a storage house for calcium within the muscle) and release calcium into the sarcoplasm (fluid within the muscle cell). Some of this calcium attaches to troponin which causes a change in the muscle cell that moves tropomyosin out of the way so the cross bridges can attach and produce muscle contraction.
In summary the sliding filament theory of muscle contraction can be broken down into four distinct stages, these are;
1. Muscle activation: The motor nerve stimulates an action potential (impulse) to pass down a neuron to the neuromuscular junction. This stimulates the sarcoplasmic reticulum to release calcium into the muscle cell.
2. Muscle contraction: Calcium floods into the muscle cell binding with troponin allowing actin and myosin to bind. The actin and myosin cross bridges bind and contract using ATP as energy (ATP is an energy compound that all cells use to fuel their activity – this is discussed in greater detail in the energy system folder here at ptdirect).
3. Recharging: ATP is re-synthesised (re-manufactured) allowing actin and myosin to maintain their strong binding state
4. Relaxation: Relaxation occurs when stimulation of the nerve stops. Calcium is then pumped back into the sarcoplasmic reticulum breaking the link between actin and myosin. Actin and myosin return to their unbound state causing the muscle to relax. Alternatively relaxation (failure) will also occur when ATP is no longer available.
In order for a skeletal muscle contraction to occur;
1. There must be a neural stimulus
2. There must be calcium in the muscle cells
3. ATP must be available for energy
So, a few things can stop a contraction;
1. Energy system fatigue: There is no more ATP left in the muscle cell so it can’t keep contracting.
2. Nervous system fatigue: The nervous system is not able to create impulses sufficiently or quickly enough to maintain the stimulus and cause calcium to release.
3. Voluntary nervous system control: The nerve that tells the muscle to contract stops sending that signal because the brain tells it to, so no more calcium ions will enter the muscle cell and the contraction stops.
4. Sensory nervous system information: For example, a sensory neuron (nerves that detect stimuli like pain or how heavy something is) provides feedback to the brain indicating that a muscle is injured while you are trying to lift a heavy weight and consequently the impulse to that muscle telling it to contract is stopped.
In the gym or during exercise virtually all muscular fatigue occurring is energy system fatigue. That is, the rate of work within the muscle can not be maintained because ATP (energy) can no longer be provided. Strength and hypertrophy (training to make muscles stronger or bigger) training are prime examples of the types of training that can cause muscle failure due to energy system fatigue.