Saccadic compensation for smooth eye and head movements during head-unrestrained two-dimensional tracking.

Spatial updating is the ability to keep track of the position of world-fixed objects while we move. In the case of vision, this phenomenon is called spatial constancy and has been studied in head-restraint conditions. During head-restrained smooth pursuit, it has been shown that the saccadic system has access to extraretinal information from the pursuit system to update the objects' position in the surrounding environment. However, during head-unrestrained smooth pursuit, the saccadic system needs to keep track of three different motor commands: the ocular smooth pursuit command, the vestibuloocular reflex (VOR), and the head movement command. The question then arises whether saccades compensate for these movements. To address this question, we briefly presented a target during sinusoidal head-unrestrained smooth pursuit in darkness. Subjects were instructed to look at the flash as soon as they saw it. We observed that subjects were able to orient their gaze to the memorized (and spatially updated) position of the flashed target generally using one to three successive saccades. Similar to the behavior in the head-restrained condition, we found that the longer the gaze saccade latency, the better the compensation for intervening smooth gaze displacements; after about 400 ms, 62% of the smooth gaze displacement had been compensated for. This compensation depended on two independent parameters: the latency of the saccade and the eye contribution to the gaze displacement during this latency period. Separating gaze into eye and head contributions, we show that the larger the eye contribution to the gaze displacement, the better the overall compensation. Finally, we found that the compensation was a function of the head oscillation frequency and we suggest that this relationship is linked to the modulation of VOR gain. We conclude that the general mechanisms of compensation for smooth gaze displacements are similar to those observed in the head-restrained condition.