The Mechanism of Ciliary Movement. I. Ciliary Reversal and Activation by Electric Current; the Ludloff Phenomenon in Terms of Core and Volume Conductors.*

Abstract

In Paramecium there is no known correlation between the direction of electric current through the membrane and of ciliary beat. One reason is that the Ludloff phenomenon, an anodal shift in the limit of the area of reversal with increased current strength, has seemed contradictory to most other data. However, by assuming Paramecium to be a core conductor immersed in a volume conductor and by applying the laws of polarizing currents it is possible to explain all existing data on reversal of normal ciliary action, and also on activation of cilia in immobilized specimens by electrical current. It is assumed that a threshold degree of depolarization of the normal membrane potential or of current density causes reversal. The Ludloff phenomenon is caused by anodal progression of this degree of depolarization with increasing membranecurrent. If it is also assumed that an increase in the membrane potential of immobilized specimens causes activation in the normal direction, one can predict anodal activation, progression of reversal with decrement in velocity, time course of development of excitation, ancdal stimulation upon “break.” stimulation by linearly rising currents, relative refractory and supernormal periods, effect of angle of orientation, and effect of acetylcholine and antiacetylchoiine esterase. Assumption of a neuromotor system is not needed. However, if available data are interpreted in the manner commonly used for nerve it can be concluded that an active accommodative process exists and possibly also a local excitatory state. A recent “dipolar” theory of galvanotaxis is not acceptable because it does not include ciliary reversal.