Secondary adaptation of memory-guided saccades.

Adaptation of saccade gains in response to errors keeps vision and action co-registered in the absence of awareness or effort. Timing is key, as the visual error must be available shortly after the saccade is generated or adaptation does not occur. Here, we tested the hypothesis that when feedback is delayed, learning still occurs, but does so through small secondary corrective saccades. Using a memory-guided saccade task, we gave feedback about the accuracy of saccades that was falsely displaced by a consistent amount, but only after long delays. Despite the delayed feedback, over time subjects improved in accuracy toward the false feedback. They did so not by adjusting their primary saccades, but via directed corrective saccades made before feedback was given. We propose that saccade learning may be driven by different types of feedback teaching signals. One teaching signal relies upon a tight temporal relation with the saccade and contributes to obligatory learning independent of awareness. When this signal is ineffective due to delayed error feedback, a second compensatory teaching signal enables flexible adjustments to the spatial goal of saccades and helps maintain sensorimotor accuracy.