Nonlinearity of spatial summation in simple cells of areas 17 and 18 of cat visual cortex.

1. Nonlinearity of spatial summation in areas 17 and 18 of cat visual cortex was compared with the type of spatial nonlinearity that differentiates X and Y cells in the lateral geniculate nucleus (LGN) and retina. The comparisons were made to examine to what extent the information from X and Y cells may remain separated in higher visual centers. 2. Responses of simple cells in areas 17 and 18 were recorded while stationary, optimally oriented sinewave gratings were sinusoidally modulated within the receptive field of the cell. Both the spatial frequency and spatial phase of the stimulus were varied. 3. Y cells in the retina and LGN are defined by the presence of a specific form of spatial nonlinearity. When tested with contrast-modulated sinewave gratings of spatial frequencies about three-fold greater than the optimal, their responses are dominated by a frequency-doubled component. The amplitude of the frequency-doubled component is not dependent on the spatial phase of the stimulus. 4. Many simple cells in the cortex showed a form of spatial nonlinearity similar to the defining nonlinearity found in retinal and geniculate Y cells. A frequency-doubled response dominated at spatial frequencies more than threefold greater than the optimal spatial frequency. When this response was present, it was phase independent. 5. More than 50% of the simple cells in area 18 showed the Y-like spatial nonlinearity. Fewer than 10% of the simple cells in area 17 showed the Y-like spatial nonlinearity. 6. The virtual absence of Y-like nonlinearity in area 17 and its relative abundance in area 18 suggest that the functional separation between the parallel X and Y pathways remains distinct within areas 17 and 18 of cat visual cortex.