Gaze characteristics of freely walking blowflies Calliphora vicina in a goal-directed task

In contrast to flying flies, walking flies experience relatively strong rotational gaze shifts, even during overall straight phases of locomotion. These gaze shifts are caused by the walking apparatus and modulated by the stride frequency. Accordingly, even during straight walking phases, the retinal image flow is composed of both translational and rotational optic flow, which might affect spatial vision, as well as fixation behavior. We addressed this issue for an orientation task where walking blowflies approached a black vertical bar. The visual stimulus was stationary, or either the bar or the background moved horizontally. The stride-coupled gaze shifts of flies walking toward the bar had similar amplitudes under all visual conditions tested. This finding indicates that these shifts are an inherent feature of walking, which are not even compensated during a visual goal fixation task. By contrast, approaching flies showed a frequent stop-and-go behavior that was affected by the stimulus conditions. As sustained image rotations may impair distance estimation during walking, we propose a hypothesis that explains how rotation-independent translatory image flow containing distance information can be determined. The algorithm proposed works without requiring differentiation at the behavioral level of the rotational and translational flow components. By contrast, disentangling both has been proposed to be necessary during flight. By comparing the retinal velocities of the edges of the goal, its rotational image motion component can be removed. Consequently, the expansion velocity of the goal and, thus, its proximity can be extracted, irrespective of distance-independent stride-coupled rotational image shifts.