Dissociable roles of human frontal eye fields and early visual cortex in presaccadic attention - evidence from TMS.

Shortly before each saccadic eye movement, presaccadic attention improves visual sensitivity at the saccade target 1-5 at the expense of lowered sensitivity at non-target locations 6-11 . Some behavioral and neural correlates of presaccadic attention and covert attention -which likewise enhances sensitivity, but during fixation 12 -are similar 13 . This resemblance has led to the debatable 13-18 notion that presaccadic and covert attention are functionally equivalent and rely on the same neural circuitry 19-21 . At a broad scale, oculomotor brain structures (e.g., FEF) are also modulated during covert attention 22-24 - yet by distinct neuronal subpopulations 25-28 . Perceptual benefits of presaccadic attention rely on feedback from oculomotor structures to visual cortices 29,30 ( Fig. 1a ); micro-stimulation of FEF in non-human primates affects activity in visual cortex 31-34 and enhances visual sensitivity at the movement field of the stimulated neurons 35-37 . Similar feedback projections seem to exist in humans: FEF+ activation precedes occipital activation during saccade preparation 38,39 and FEF TMS modulates activity in visual cortex 40-42 and enhances perceived contrast in the contralateral hemifield 40 . We investigated presaccadic feedback in humans by applying TMS to frontal or visual areas during saccade preparation. By simultaneously measuring perceptual performance, we show the causal and differential roles of these brain regions in contralateral presaccadic benefits at the saccade target and costs at non-targets: Whereas rFEF+ stimulation reduced presaccadic costs throughout saccade preparation, V1/V2 stimulation reduced benefits only shortly before saccade onset. These effects provide causal evidence that presaccadic attention modulates perception through cortico-cortical feedback and further dissociate presaccadic and covert attention.