Spatial summation in the receptive fields of simple cells in the cat's striate cortex

1. We have examined the responses of simple cells in the cat's atriate cortex to visual patterns that were designed to reveal the extent to which these cells may be considered to sum light-evoked influences linearly across their receptive fields. We used one-dimensional luminance-modulated bars and grating as stimuli; their orientation was always the same as the preferred orientation of the neurone under study. The stimuli were presented on an oscilloscope screen by a digital computer, which also accumulated neuronal responses and controlled a randomized sequence of stimulus presentations. 2. The majority of simple cells respond to sinusoidal gratings that are moving or whose contrast is modulated in time in a manner consistent with the hypothesis that they have linear spatial summation. Their responses to moving gratings of all spatial frequencies are modulated in synchrony with the passage of the gratings' bars across their receptive fields, and they do not produce unmodulated responses even at the highest spatial frequencies. Many of these cells respond to temporally modulated stationary gratings simply by changing their response amplitude sinusoidally as the spatial phase of the grating the grating is varied. Nonetheless, their behavior appears to indicate linear spatial summation, since we show in an Appendix that the absence of a 'null' phase in a visual neurone need not indicate non-linear spatial summation, and further that a linear neurone lacking a 'null' phase should give responses of the form that we have observed in this type of simple cell. 3. A minority of simple cells appears to have significant non-linearities of spatial summation. These neurones respond to moving gratings of high spatial frequency with a partially or totally unmodulated elevation of firing rate. They have no 'null' phases when tested with stationary gratings, and reveal their non-linearity by giving responses to gratings of some spatial phases that are composed partly or wholly of even harmonics of the stimulus frequency ('on-off' responses). 4. We compared simple receptive fields with their sensitivity to sinusoidal gratings of different spatial frequencies. Qualitatively, the most sensitive subregions of simple cells' receptive fields are roughly the same width as the individual bars of the gratings to which they are most sensitive. Quantitatively, their receptive field profiles measured with thin stationary lines, agree well with predicted profiles derived by Fourier synthesis of their spatial frequency tuning curves.