Receptive field organization of complex cells in the cat's striate cortex.

1. All complex cells in the cat's striate cortex exhibit gross non-linearities of spatial summation when tested with sinusoidal grating stimuli. Their responses to moving gratings of all but the lowest spatial frequencies are usually dominated by a component that is not modulated by the passage of the bars of the grating across the receptive field. They give responses to temporally modulated stationary gratings that consist mostly of even harmonics of the stimulus frequency and that vary little in amplitude or wave form as the spatial phase of the grating is varied. 2. We compared complex cells' receptive fields with their sensitivity to sinusoidal gratings of different spatial frequencies. Qualitatively, the receptive fields are usually two to five times wider than the bars of the gratings that stimulate them most effectively. Quantitatively, the receptive field profiles of complex cells are invariably broader than those predicted by Fourier synthesis of their spatial frequency tuning curves, and in particular lack predicted spatially antagonistic regions. 3. We further examined the receptive field organization of these cells, using pairs of stationary lines flashed synchronously on their receptive fields. If both lines are of the same polarity (bright or dark), complex cells respond to the paired stimulus much less well than they do to either of its component bars, unless the bars are separated by less than about one quarter of the width of the receptive field. If the lines are of opposite polarity, one bright and one dark, the opposite situation obtains: closely spaced bars elicit small responses, while paired bars of larger separation are much more effective. In either case, the results are independent in general character of the absolute positions of the stimuli within the receptive field; rather, they depend in a manner characteristic of each cell on the relative positions of the two bars. 4. The two-line interaction profile that plots the change in a complex cell's response to one bar as a function of the position of a second added bar corresponds closely to the receptive field profile predicted from Fourier synthesis of the cell's spatial frequency tuning curve. These profiles may thus reveal the spatial characteristics of subunits within complex cell-receptive fields. We examined the nature of the interaction between these subunits by performing several two-line interaction experiments in which the onset of the second bar was delayed some time after the onset of the first. The results suggest that neighbouring subunits interact in a facilitatory fashion: for an interval after the presentation of one bar, responses to neighbouring bars are enhanced. 5. The subunits of a complex receptive field may, by their spatial properties, determine the spatial selectivities of complex cells, while the nature of the interaction among the subunits may determine these cells' sensitivity and selectivity for moving visual stimuli...