Bistable membrane potential of the ciliate Coleps hirtus

In normal recording solution, the swimming pattern of the freshwater ciliate Coleps hirtus, belonging to the class Prostomatea, consists of alternating periods of nearly linear forward swimming and circular swimming within a small area. Current-clamp recordings were performed to elucidate the mechanism for this behaviour. No members of this class have previously been studied using electrophysiological techniques. The ciliates were maintained in culture and fed on the planctonic alga Rhodomonas minuta. The membrane potential showed spontaneous shifts between a more negative (deep) level of approximately -50 mV and a less negative (shallow) level of approximately -30 mV. The input resistance and capacitance at the more negative level were approximately 400 M capomega and 120 pF respectively. C. hirtus displayed a pronounced inward rectification, which was virtually insensitive to 1 mmol l(-1) Cs(+) and almost completely blocked by 1 mmol l(-1) Ba(2+). Depolarising current injections failed to evoke graded, regenerative Ca(2+) spikes. However, current-induced depolarisations from the more negative potential level (-50 mV) showed a pronounced shoulder during the repolarising phase. Increased current injections prolonged the shoulder, which occasionally stabilised at the shallow membrane potential (-30 mV). The membrane potential could be shifted to the deep level by brief hyperpolarising current injections. Similar biphasic membrane properties have not been reported previously in any ciliate. The bistability of the membrane potential was abolished in Ca(2+)-free solution containing Co(2+) or Mg(2+). In Ca(2+)-free solution containing 1 mmol l(-1) Ba(2+), brief depolarising current injections at the deep potential level evoked all-or-nothing action potentials with a prolonged plateau coinciding with the shallow potential. We conclude that the deep membrane potential in C. hirtus corresponds to the traditional resting potential, whereas the shallow level is a Ca(2+)-dependent plateau potential. In normal solution, the direction of the ciliary beat was backwards at the deep potential level and forwards at the shallow membrane potential, probably reflecting the two main phases of the swimming pattern.