THE EFFECTIVENESS OF THE LONGITUDINAL FIELD, COUPLED WITH DEPOLARIZATION IN ACTIVATING FROG TWITCH MUSCLES

Effective excitation, preceding the mechanical response in frog twitch muscles, involves two distinct events: depolarization of the excitable membrane and the flow of internal currents. To distinguish between the effects of these two potential factors in activation of the contractile machinery, experiments ought to be conducted in which one or the other is excluded. Our experiments are designed to distinguish between these effects by indirect methods. Depolarization in a longitudinal electric field can be expected to be greatest at the ends where current leaves the muscle fibers (space constant at [K] = 16 mM/liter is >1 mm.), whereas the internal longitudinal current is known to be greatest in the middle portion. Depolarization, therefore, should affect the ends more strongly and internal current the middle portion. Our experiments show that non-propagating frog twitch muscles shorten, during isotonic work, along their whole length and not only at their ends, when effectively stimulated in a longitudinal A.C. field. At a field strength about twice the new threshold value (at [K](o) = 16 mM) shortening is distinctly greater in the middle portion of the muscle than at the ends. The muscles, although temporarily non-propagating, remain intact throughout the experiment, as demonstrated by complete recovery after repolarization. These findings may be taken as an indication that internal currents are more directly linked to activation than is depolarization, but the latter is an essential priming step, which must precede or coincide with effective current flow.