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By Johann Caspar Rüegg M.D., Ph.D. (auth.)

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The latter, on the other hand, contains in addition to the T-tubular system, which is located near the borders of the A-band, a different type of tubules near the Zlines, the Z-tubules. These, however, do not seem to be involved in excitation-contraction coupling since, in contrast to stimulation of the A-band borderline region, depolarization of the cell membrane precisely over the Z-disc did not induce a local contraction (Peachey and Huxley 1964). With the exception of the body wall muscle of the nematode Ascaris lumbricoides (Rosenbluth 1965), the obliquely striated muscles of nematodes, annelids and molluscs do not contain T-tubules.

The possible chemical states and changes in free energy during the cycle are after Goody and Holmes (1983). Note that ATP binding detaches crossbridges which reattach after A TP hydrolysis to form an "energy-rich" complex of actin, myosin and the products of ATP hydrolysis. The release of products is associated with a large drop offree energy ( - 32 kJ) which may be converted into mechanical work cause statistically, at any given moment, there is the same number of bridges attached in the force-generating state.

In this coupling calcium ions are implicated as intracellular messengers linking the membrane events to the contractile machinery. Contraction may be characterized by the force that muscle develops under isometric conditions or by the extent and rate of shortening of a loaded muscle. In activated muscle "side projections" from the myosin filaments (crossbridges) attach to actin filaments, detach and reattach again in a cyclic fashion. One molecule of ATP is presumably hydrolyzed in each cyclic operation of the actin-myosin cross bridge so that, at the biochemical level, actomyosin-ATPase activity may be taken as an "easy to study" biochemical correlate of contraction.

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