Oculomotor responses to gradual changes in target direction.

Smooth pursuit tracking of targets moving linearly (in one dimension) is well characterized by a model where retinal image motion drives eye acceleration. However, previous findings suggest that this model cannot be simply extended to two-dimensional (2D) tracking. To examine 2D pursuit, in the present study, human subjects tracked a target that moved linearly and then followed the arc of a circle. The subjects' gaze angular velocity accurately matched target angular velocity, but the direction of smooth pursuit always lagged behind the current target direction. Pursuit speed slowly declined after the onset of the curve (for about 500 ms), even though the target speed was constant. In a second experiment, brief perturbations were presented immediately prior to the beginning of the change in direction. The subjects' responses to these perturbations consisted of two components: (1) a response specific to the parameters of the perturbation and (2) a nonspecific response that always consisted of a transient decrease in gaze velocity. With the exception of this nonspecific response, pursuit behavior in response to the gradual changes in direction and to the perturbations could be explained by using retinal slip (image velocity) as the input signal. The retinal slip was parallel and perpendicular to the instantaneous direction of pursuit ultimately resulted in changes in gaze velocity (via gaze acceleration). Perhaps due to the subjects' expectations that the target will curve, the sensitivity to the image motion in the direction of pursuit was not as strong as the sensitivity to image motion perpendicular to gaze velocity.