Behavior of Delayed Current under Voltage Clamp in the Supramedullary Neurons of Puffer

Depolarizations applied to voltage-clamped cells bathed in the normal solution disclose an initial inward current followed by a delayed outward current. The maximum slope conductance for the peak initial current is about 30 times the leak conductance, but the maximum slope conductance for the delayed current is only about 10 times the leak conductance. During depolarizations for as long as 30 sec, the outward current does not maintain a steady level, but declines first exponentially with a time constant of about 6 msec; it then tends to increase for the next few seconds; finally, it declines slowly with a half-time of about 5 sec. Concomitant with the changes of the outward current, the membrane conductance changes, although virtually no change in electromotive force occurs. Thus, the changes in the membrane conductance represent two phases of K inactivation, one rapidly developing, the other slowly occurring, and a phase of K reactivation, which is interposed between the two inactivations. In isosmotic KCl solution after a conditioning hyperpolarization there occurs an increase in K permeability upon depolarization. When the depolarizations are maintained, the increase of K permeability undergoes changes similar to those observed in the normal medium. The significance of the K inactivation is discussed in relation to the after-potential of the nerve cells.