VOLTAGE-GATED NA-SHAPED AND SPIKE-SHAPED CONDITIONING DEPOLARIZATIONSOF RETINAL GANGLION-CELLS( CURRENT AVAILABILITY AFTER STEP)

We used two conditioning voltage protocols to assess inactivation of v oltage-gated Na+ current in retinal ganglion cells. The first protocol tested the possibility, raised by published activation and steady-sta te inactivation curves, that Na+ ions carry a ''window'' current in th ese cells. The second protocol was used, because these cells spike rep etitively in situ, to measure the Na+ current available for activation following spikes. Na+ current activated at test potentials more posit ive than -65 mV. At test potentials more positive than -55 mV,Naf curr ent peaked and then declined along a time course that could be fit by the sum of a large, rapidly decaying component, a small, slowly decayi ng component and a non-decaying component. Both step- and spike-shaped conditioning depolarizations reduced the amount of current available for subsequent activation, sparing the non-decaying ''persistent'' com ponent. Most of the Na+ current recovered from this inactivation along a rapid exponential time course (tau=3 ms). The remaining recovery wa s complete within at least 4 s (at -70 mV). Our use of step depolariza tions has identified a current component not anticipated from previous measurements of steady-state inactivation in retinal ganglion cells. Our use of spike-shaped depolarizations shows that Na+ current density at 1 ms after a single spike is roughly 25% of that activated by the conditioning spike, and that recovery from inactivation is 50-90% comp lete within 10 ms thereafter. Na+ current amplitude declines during sp ikes repeated at relatively low frequencies, consistent with a slow co mponent of full recovery from inactivation.