Apart from their primary function as balance sensors, Hermissenda hair
cells are presynaptic neurons involved in the Ca2+-dependent neuronal
plasticity in postsynaptic B photoreceptors that accompanies classica
l conditioning. With a view to beginning to understand presynaptic mec
hanisms of plasticity in the vestibule-visual system, a locus for cond
itioning-induced neuronal plasticity, outward currents that may govern
the excitability of hair cells were recorded by means of a whole-cell
patch-clamp technique. Three K+ currents were characterized: a 4-amin
opyridine-sensitive transient outward K+ current (I-A), a tetraethyl a
mmonium-sensitive delayed rectifier K+ current (I-K,I-V), and a Ca2+-a
ctivated K+ current (I-K,I-Ca). I-A activates and decays rapidly; the
steady-state activation and inactivation curves of the current reveal
a window current close to the apparent resting voltage of the hair cel
ls, suggesting that the current is partially activated at rest. By mod
ulating firing frequency and perhaps damping membrane oscillations, I-
A may regulate synaptic release at baseline. In contrast, I-KV and I-K
,I-Ca have slow onset and exhibit little or no inactivation. These two
K+ currents may determine the duration of the repolarization phase of
hair-cell action potentials and hence synaptic release via Ca2+ influ
x through voltage-gated Ca2+ channels. In addition, I-K,I-Ca may be re
sponsible for the afterhyperpolarization of hair cell membrane voltage
following prolonged stimulation.