Responses of simple and complex cells to compound sine-wave gratings

We have studied the responses of simple and complex cells in the primary visual cortex of the cat to rigidly drifting compound sine-wave gratings as a function of the phase offset between fundamental and harmonic frequencies that both fell within the passband of the cell. Simple cells show phase-dependent increases and decreases in peak and mean response which are predictable on the basis of a cell's line weighting function. However, the amplitudes and phases of the base and harmonic frequencies in the response are, in general, not well predicted by the relationships of these same components in the compound grating stimuli. These distortions are shown to be largely a consequence of the rectification that follows linear summation at the simple cell stage. Such distortions are, in principle, correctable when the responses of a second simple cell, as part of a 180 deg phase pair, are taken into account. Complex cells typically showed a strong nonlinear response component at the difference frequency of drifting compound gratings. This was sometimes accompanied by a linear response component at one, or both, of the separate stimulus frequencies. Information about the absolute phases of the frequency components of a compound grating is not preserved in the nonlinear response of complex cells; however, information about the local phase difference between the gratings is preserved. In effect, the nonlinear component of the complex cell response is proportional to the time-varying signal envelope that results from the mutual interference of stimulus frequencies that fall in the cell's spatial receptive field and frequency passband.