An objective signature for visual binding of face parts in the human brain

Whether and how the parts of a visual object are grouped together to form an integrated ("holistic") representation is a central question in cognitive neuroscience. Although the face is considered to be the quintessential example of holistic representation, this issue has been the subject of much debate in face perception research. The implication of holistic processing is that the response to the whole cannot be predicted from the sum of responses to the parts. Here we apply techniques from nonlinear systems analysis to provide an objective measure of the nonlinear integration of parts into a whole, using the left and right halves of a face stimulus as the parts. High-density electroencephalogram (EEG) was recorded in 15 human participants presented with two halves of a face stimulus, flickering at different frequencies (5.88 vs. 7.14 Hz). Besides specific responses at these fundamental frequencies, reflecting part-based responses, we found intermodulation components (e.g., 7.14 - 5.88 = 1.26 Hz) over the right occipito-temporal hemisphere, reflecting nonlinear integration of the face halves. Part-based responses did not depend on the relative alignment of the two face halves, their spatial separation, or whether the face was presented upright or inverted. By contrast, intermodulations were virtually absent when the two halves were spatially misaligned and separated. Inversion of the whole face configuration also reduced specifically the intermodulation components over the right occipito-temporal cortex. These observations indicate that the intermodulation components constitute an objective, configuration-specific signature of an emergent neural representation of the whole face that is distinct from that generated by the parts themselves.