THE ACTIONS OF CALCIUM ON THE MECHANOELECTRICAL TRANSDUCER CURRENT OF TURTLE HAIR-CELLS

1. Mechano-electrical transducer currents evoked by deflections of the hair bundle were recorded in turtle isolated hair cells under whole-cell voltage clamp. The outcome of perfusing with solutions of reduced Ca2+ concentration was investigated. 2. The transducer current was roughly doubled by lowering the concentration of divalent cations from normal (2.2 mM-Mg2+, 2.8 mM-Ca2+) to 0 Mg2+, 0.5 mM-Ca2+. No significant effects on the current's kinetics or reversal potential, or on the current-displacement relationship, were noted. 3. If the Ca2+ concentration was lowered to 50-mu-M (with no Mg2+), there was about a threefold increase in the maximum current but other changes, including loss of adaptation and a decreased slope and negative shift in the current-displacement relationship, were also observed. As a result, more than half the peak transducer current became activated at the resting position of the hair bundle compared to about a tenth in the control solution. 4. The extra changes manifest during perfusion with 50-mu-M-Ca2+ had also been seen when the cell was held at positive potentials near the Ca2+ equilibrium potential. This supports the view that some consequences of reduced external Ca2+ stem from a decline in its intracellular concentration. 5. With 20-mu-M-Ca2+, a standing inward current developed and the cell became unresponsive to mechanical stimuli, which may be explained by the transducer channels being fully activated at the resting position of the bundle. 6. The results are interpreted in terms of a dual action of Ca2+: an external block of the transducer channel which reduces the maximum current, and an intracellular effect on the position and slope of the current-displacement relationship; the latter effect can be modelled by internal Ca2+ stabilizing one of the closed states of the channel. 7. During perfusion with 1-mu-M-Ca2+, the holding current transiently increased but then returned to near its control level. There was a concomitant irreversible loss of sensitivity to hair bundle displacements which we suggest is due to rupture of the mechanical linkages to the transducer channel. 8. Following treatment with 1-mu-M-Ca2+, single-channel currents with an amplitude of -9 pA at -85 mV were sometimes visible in the whole-cell recording. The probability of such channels being open could be modulated by small deflections of the hair bundle which indicates that they may be the mechano-electrical transducer channels of conductance about 100 pS. 9. Open- and closed-time distributions for the channel were fitted by single exponentials, the mean open time at rest being approximately 1 ms. The mean open time was increased and the mean closed time decreased for movements of the hair bundle towards the kinocilium. 10. The Ca2+ concentration in the endolymph of the turtle's cochlear duct was measured with a Ca2+ -sensitive microelectrode and found to be 65-mu-M. The significance of the composition of endolymph for mechano-electrical transduction by hair cells is discussed.