Fast-acting compressive and facilitatory nonlinearities in light-adapted fly photoreceptors

Light-adapted fly photoreceptor cells were stimulated with brief positive and negative contrast flashes (contrast = delta I/I, I = intensity). Membrane potential responses to a wide range of flash intensities were well-fitted by a static nonlinearity followed by a compartmental model represented by a gamma function. However, the agreement improved if one parameter of the gamma function, n, varied quadratically with input light intensity. Response amplitude and time to peak were estimated from the fitted parameters. Response amplitude varied approximately linearly with contrast but showed nonlinear compression with the largest negative flashes. Reducing the background light level by 3 decades or hyperpolarizing the cell electrically produced stronger nonlinear compression with both contrast polarities. This is probably due to fast voltage-activated K+ channels. Responses to double flashes with varying time separations were well-fitted by summed gamma functions, allowing separation of the individual flash responses. There was no detectable time-dependent interaction between paired positive flashes at all separations. However, the response to two negative flashes was greater than the linear prediction at short separations, and this facilitatory nonlinearity decayed with a time constant of about 1 msec. The facilitation is probably related to resonant behavior in light-adapted photoreceptors and may be due to an IP3-induced intracellular Ca2+ release.