Evidence for nonretinal feedback in combined version-vergence eye movements.

Recently, a quantitative model for the general of rapid eye movements in direction and depth was proposed. In this scheme, the saccadic and the vergence system share a common initiation system and are controlled by local feedback loops based on efference copy signals. We have used a remembered-target double-step paradigm to test the idea that both subsystems are guided by extraretinal signals. The subject was instructed to move the binocular point of fixation to the remembered positions indicated by a double-step movement of the target, in direction and depth. Since both binocular refixations were made in complete darkness, correct execution of this task requires information about both the stored visual coordinates of the final target and the coordinates of the first movement. Binocular eye movements from five subjects were compared with predictions from two feed-forward models and a feedback model. Analysis of the pooled direction data showed that the feedback model performed best and fitted well. Qualitatively the same result was obtained in the vergence component, but in this case the goodness of fit was considerably less. These results, confirmed in each individual subject, show that the saccadic and vergence subsystem can use nonretinal information about a prior movement in direction and depth. Further analysis showed that the gain of the direction response of the second movement was, on average, roughly correct. By contrast, the vergence component of these responses was only about 60% of the required amplitude. Since the fit procedure gave the same weighting factors to the second target and to the first movement, we propose that the low vergence gain reflects mechanisms operating after the calculation of the motor error signal, possibly at the execution stage. Finally, we discuss the possibility of a central control stage keeping track of the ongoing movement sequence, based on a comparison of desired and current eye position signals.