Functional but not obligatory link between microsaccades and neural modulation by covert spatial attention

Covert spatial attention is associated with spatially specific modulation of neural activity as well as with directional biases in fixational eye-movements known as microsaccades. Recently, this link has been suggested to be obligatory, such that modulation of neural activity by covert spatial attention occurs only when paired with microsaccades toward the attended location. Here we revisited this link between microsaccades and neural modulation by covert spatial attention in humans. We investigated spatial modulation of 8-12 Hz EEG alpha activity and microsaccades in a context with no incentive for overt gaze behaviour: when attention is directed internally within the spatial layout of visual working memory. In line with a common attentional origin, we show that spatial modulations of alpha activity and microsaccades co-vary: alpha lateralisation is stronger in trials with microsaccades toward compared to away from the memorised location of the to-be-attended item and occurs earlier in trials with earlier microsaccades toward this item. Critically, however, trials without attention-driven microsaccades nevertheless showed clear spatial modulation of alpha activity - comparable to the neural modulation observed in trials with attention-driven microsaccades. Thus, directional biases in microsaccades are correlated with neural signatures of covert spatial attention, but they are not a prerequisite for neural modulation by covert spatial attention to be manifest.