Perception of illusory contours forms intermodulation responses of steady state visual evoked potentials as a neural signature of spatial integration

Perception of illusory contours was shown to be a consequence of neural activity related to spatial integration in early visual areas. Candidates for such filling-in phenomena are long-range horizontal connections of neurons in V1/V2, and feedback from higher order visual areas. To get a direct measure of spatial integration in early visual cortex, we presented two differently flickering inducers, which evoked steady-state visual evoked potentials (SSVEPs) while manipulating the formation of an illusory rectangle. As a neural marker of integration we tested differences in amplitudes of intermodulation frequencies i.e. linear combinations of the driving frequencies. These were significantly increased when an illusory rectangle was perceived. Increases were neither due to changes of any of the two driving frequencies nor in the frequency that tagged the processing of the compound object, indicating that results are not a consequence of paying more attention to inducers when the illusory rectangle was visible.