Adaptation of response transients in fly motion vision. I: Experiments

Two types of transient responses have been investigated in fly motion-sensitive neurons in the past: the impulse and the step response. In response to a brief motion pulse, cells show a sudden rise in activity followed by an exponential decay ('impulse response'). In response to the onset of a constant velocity stimulus, cells exhibit transient oscillations before settling to a steady-state value ('step response'). Since the impulse response has been shown to shorten when tested after presentation of an adapting motion stimulus, we investigated whether adaptation also occurs during the step response. We tested this hypothesis by recording extracellularly the response of the H1-cell in the lobula plate of the blowfly Calliphora vicina to gratings of varying pattern contrasts and drift velocity. We found that the transient oscillations of the step response strongly depend on the pattern contrast: at low contrasts, oscillations lasted for several seconds, whereas at high contrasts, they settled within fractions of a second. This suggests that motion adaptation occurs during the initial period of the stimulus presentation and is dependent on the contrast of the motion stimulus. Using identical stimulus parameters (contrast and temporal frequency) for the adapting stimulus and testing the impulse response afterwards, we found that the impulse response and the transient period in the step response shortened in a similar way. We then analyzed the dynamic of the transients oscillations produced by ongoing motion of a square wave pattern in the anti-preferred direction (null direction) of H1. As observed for preferred direction motion, we found that the duration and amplitude of those transients shortened as the contrast and the velocity of the pattern increased, and that the oscillations disappeared when a blank screen instead of a pattern was presented before the onset of motion. Under both stimulus conditions, i.e. grating and blank screen before motion onset, the steady-state response level showed the same dependence on the contrast and temporal frequency of the pattern. When we analyzed the responses of the cell to pattern of various sizes and contrasts moving in the preferred direction of the cell, we found that increments in the size affected the overall amplitude of both the transient oscillations and the steady-state response level, whereas the duration of the oscillations only depended on the local pattern contrast. We also tested the impulse response before and after the presentation of an adapting stimulus presented in either the same or a different location of the visual field. The response shortened only when both the adapting and the test stimuli were presented at the same location. These last experiments demonstrate a strictly local mechanism of adaptation affecting the response transients of both the impulse and the step response.