muscle

muscle is the body's contractile tissue. ‘Contraction’, in the physiological sense, may involve shortening and change of shape, or it may generate force without any change in length. All contraction depends on physicochemical alterations in the molecules of protein filaments within the cells, resulting in the generation of force at linkages (cross-bridges) between two different kinds of filament. The main proteins involved, in the respective filaments of all types of muscle, are actin and myosin; and in all muscles the process is powered by breakdown of adenosine triphosphate, during which chemical energy is converted by the interactions between these proteins into the mechanical energy of contraction. To initiate the process, muscle cells require excitation, which leads to contraction by a sequence that crucially involves an increase in the concentration of free calcium ions inside the cell — a sequence termed excitation– contraction coupling.

There are three main types of muscle in the body: skeletal, cardiac, and smooth. When skeletal muscles contract they either move parts of the body via their attachments to bones, or produce tension to oppose stretch or even to allow controlled lengthening. Cardiac muscle and smooth muscle, by shortening, reduce the capacity of hollow organs and tubes: thus cardiac muscle ejects blood from the heart; smooth muscle ejects urine from the bladder or the fetus from the uterus, moves the contents of the gut along, and influences the flow of blood to different regions by varying the diameter of blood vessels.

Skeletal and cardiac are together known as striated muscles, because their fibres have a striped appearance under the microscope, due to the orderly arrangement of alternating ranks of interdigitating actin and myosin filaments within their cytoplasm. Smooth (unstriated) muscle does not show this: the two types of filament are mingled throughout the cytoplasm of the cells. Whilst cardiac and skeletal muscle have a structural resemblance, skeletal muscle can be under conscious control and is therefore also known as voluntary muscle whereas cardiac muscle and smooth muscle share the designation involuntary because their actions are never under direct conscious control. (In certain contemplative regimes, the subtle influence which may be achieved — such as on the heart rate — is an indirect consequence of a profoundly disciplined emotional state.)

The voluntary/involuntary distinction implies differences also in control of the three types of muscle. Skeletal muscle is controlled through pathways in the nervous system that can be consciously activated, cardiac and smooth by the involuntary or ‘autonomic’ pathways. Each skeletal muscle fibre is called into action by release of transmitter from a terminal branch of a single axon from a motor neuron in the spinal cord; the point at which this nerve terminal contacts the muscle fibre is a specialized synapse, the neuromuscular junction. All muscle fibres controlled by this nerve are recruited together, and the grouping of a motor neuron plus its family of muscle fibres is said to comprise a ‘motor unit’. When transmitter is not being released, the muscle fibres are relaxed. Individual cardiac muscle cells by contrast are activated by electrical transmission of excitation from their neighbours; this excitation originates rhythmically at a pacemaker, even in the absence of nerve action, although normally the rate of firing is modulated by the release, close to the pacemaker site, of transmitters from autonomic nerves. Smooth muscles differ again: in some, notably in the uterus at term, excitation is electrical, starting at pacemaker sites, much as in the heart. In others, such as those controlling the diameter of a large blood vessel, excitation is by neurotransmitters released from autonomic nerve endings close to the cells, but not with structured synapses. The contraction/relaxation state of smooth muscle can also be modified by chemical agents other than neurotransmitters, released from neighbouring cells or circulating in the blood. In the autonomic control of involuntary muscle, there is at many sites the possibility of either excitatory or inhibitory neural action, according to the particular transmitter released, resulting in a two-way control system analogous to accelerator and brake. The heart, for instance, is slowed by one transmitter, yet speeded up by another; the stomach wall is contracted by one and relaxed by another.

Neil Spurway

Sheila Jennett

See musculo-skeletal system.See also autonomic nervous system; cardiac muscle; motor neurons; skeletal muscle; smooth muscle.