Slow membrane currents in bursting pace-maker neurones of Tritonia

1. Membrane ionic currents in bursting pace-maker neurones of the marine mollusc Tritonia were studied in voltage-clamp experiments with emphasis on slow tail current relaxations after depolarizing pulses. 2. The slow tail current undergoes a complex transition from an initially inward current to an initially outward current as the duration of the depolarizing pulse is lengthened. It was found that the slow tail current is the sum of two separate and independent ionic currents. Methods were devised to study each current in isolation. 3. A slow inward tail current, termed IB, is activated by depolarization and decays exponentially on return to -55 mV with a time constant of 2-4 s. The voltage dependence and kinetics of IB activation were measured. Current amplitude is sensitive to removal of both Na+ and Ca2+ from the bathing medium but the current is not blocked by either tetrodotoxin (TTX) or replacement of Ca2+ by Co+. The amplitude of the current is independent of the external K+ concentration. 4. A slow outward tail current, termed IC, is also activated by depolarization. It is shown to be a K+ current whose activation results from an increase in the cytoplasmic Ca2+ concentration during depolarization. The decay of IC on repolarization requires more than 30 s to reach completion. 5. The slow rates of relaxation of IB and IC tail currents suggest that they are important determinants of the slow membrane potential variations characteristic of burst firing. IB activates more rapidly than IC during depolarization and is thought to be important for maintaining the depolarized phase of the burst cycle and for producing the depolarizing after-potential after each spike. IC activates more slowly but reaches greater amplitudes. It is thought to be important for adaptation in spike frequency during the burst, for burst termination, and for determining the duration of the interval between bursts.