Sustained spatial attention accounts for the direction bias of human microsaccades

Small ballistic eye movements, so called microsaccades, occur even while foveating an object. Previous studies using covert attention tasks have shown that shortly after a symbolic spatial cue, specifying a behaviorally relevant location, microsaccades tend to be directed toward the cued location. This suggests that microsaccades can serve as an index for the covert orientation of spatial attention. However, this hypothesis faces two major challenges: First, effects associated with visual spatial attention are hard to distinguish from those that associated with the contemplation of foveating a peripheral stimulus. Second, it is less clear whether endogenously sustained attention alone can bias microsaccade directions without a spatial cue on each trial. To address the first issue, we investigated the direction of microsaccades in human subjects while they attended to a behaviorally relevant location and prepared a response eye movement either toward or away from this location. We find that directions of microsaccades are biased toward the attended location rather than towards the saccade target. To tackle the second issue, we verbally indicated the location to attend before the start of each block of trials, to exclude potential visual cue-specific effects on microsaccades. Our results indicate that sustained spatial attention alone reliably produces the microsaccade direction effect. Overall, our findings demonstrate that sustained spatial attention alone, even in the absence of saccade planning or a spatial cue, is sufficient to explain the direction bias observed in microsaccades.