IONIC SELECTIVITY OF I(H) CHANNELS OF ROD PHOTORECEPTORS IN TIGER SALAMANDERS

Ionic selectivity of I(h) channels of tiger salamander rod photoreceptors was investigated using whole-cell voltage clamp. Measured reversal potentials and the Goldman-Hodgkin-Katz voltage equation were used to calculate permeability ratios with 20 mM K+ as a reference. In the absence of external K+, I(h) is small and hard to discern. Hence, we defined I(h) as the current blocked by 2 mM external Cs+. Some small amines permeate I(h) channels, with the following permeability ratios (P(X)/P(K)):NH4+, 0.17; methylammonium, 0.06; and hydrazine, 0.04. Other amines are tially impermeant: dimethylammonium (< 0.02), ethylammonium (< 0.01), and tetramethylammonium (< 0.01). When K+ is the only eternal permeant ion and its concentration is varied, the reversal potential of I(h) follows the Nernst potential for a K+ electrode. I(h) channels are also permeable to other alkali metal cations (P(X)/P(K)): Tl+, > 1.55; K+, l; Rb+, > 0.55; Na+, 0.33; Li+, 0.02. Except for Na+, the relative slope conductance had a similar sequence (G(X)/(G(K)): Tl+, 1.07; K+, l; Rb+, 0.37; NH4+, 0.07; Na+, 0.02. Based on permeabilities to organic cations, the narrowest part of the pore has a diameter between 4.0 and 4.6 angstrom. Some permeant cations have large effects on the gating kinetics of I(h) channels; however, permeant cations appear to have little effect on the steady-state activation curve of I(h) channels. Lowering K+ or replacing K+ with Na+ reduces the maximal conductance of I(h) but does not shift or change the steepness of its voltage dependence. With ammonium or methylammonium replacing K+ a similar pattern is seen, except that there is a small positive shift of approximately 10 mV in the voltage dependence.