50, P = 0.04, two-tailed). Interestingly, there is no difference between no TMS and TMS applied in an intermediate time window (t = 0.95, P = 0.37, two-tailed). Next, we explored the relationship between performance (i.e., correctly perceiving a stack as a stack) and late neural signaling in occipital cortex. We expected stack–frame differences to increase by excluding error trials, as these errors trials involved the mix-up of stack and frame stimuli (see Fig. 3). We therefore performed the same analysis as described above (Fig. 7A), but now excluding all error trials. Figure 7B shows that by excluding error trials, we were able to observe an enhancement (trending) Inhibitors,research,lifescience,medical of the stack–frame difference (collapsed

across TMS conditions, correct-all trials: t = 1.60, P = 0.07, one-tailed). Comparing different TMS conditions for correct-only trials resulted in a significant difference between the no TMS and early TMS condition (t = 2.62, P = 0.03, two-tailed). Interestingly, although behaviorally all EEG trials were equal (correct-only trials), we are still Inhibitors,research,lifescience,medical able

to observe a difference between the different TMS conditions (Fig. 7B). It therefore seems that TMS is able to influence neural signaling, without Inhibitors,research,lifescience,medical necessarily leading to overt behavioral effects. Figure 7 Transcranial magnetic stimulation (TMS) modulation of stack–frame difference. (A) Early TMS reduced the difference in activity evoked by stack and activity evoked by frame stimuli in comparison

with the no TMS condition (t = 2.97, P = 0.01, two-tailed) Inhibitors,research,lifescience,medical … Discussion By briefly disrupting activity in early visual cortex during a discrimination task, we were able to causally link activity in areas V1/V2 to different stages in figure–ground segregation. The present findings show that the role of early visual cortex is not limited to low-level computations, but reveal that areas V1/V2 are also necessary later in time when Inhibitors,research,lifescience,medical surface PD98059 datasheet segregation emerges. Here, we observed that disruption of V1/V2 activity in the late TMS time window resulted in reduced performance scores selectively for stack stimuli. In order to correctly discriminate a stack stimulus (from a frame stimulus) surface segregation is necessary, therefore causally linking activity in early visual cortex in this relatively late period to surface segregation. In addition, disruption of early visual cortex in this late time window selectively made participants erroneously see more stacks CYTH4 as frame stimuli supporting the claim that specifically surface segregation was affected in this time window (as frames are identical to stack stimuli except for a different amount of figure surface, see “Task design”). The necessity of early visual cortex in this late period during figure–ground segregation demonstrates that late V1/V2 activity is not epiphenomenal or merely a by-product of activity in higher (visual) areas.