Differentiation of VEP intermodulation and second harmonic components by dichoptic, monocular, and binocular stimulation

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Abstract
Modulation by two temporal frequencies differentiates visual processing at the fundamentals (1Fs), second harmonics (2Fs), and second-order intermodulation components (IMCs), the latter created neurally as the sum or difference of the two modulation frequencies. Steady-state VEPs were recorded while stereo-normal adults viewed luminance or grating stimuli modulated by up to three temporal frequencies under dichoptic, monocular, or ordinary (binocular) viewing conditions arranged using liquid crystal light shutters. In Experiment 1, modulation of luminance by a single temporal frequency produced strong 1F and 2F VEP components, but modulation of gratings produced only 2Fs. Modulation by two temporal frequencies resulted in IMCs, often in the absence of evoked activity in the EEC at the 1Fs. IMCs were generally larger during pattern as compared to luminance modulation. Amplitudes of 1Fs and IMCs were smaller, but 2Fs were larger, during dichoptic as compared to ordinary viewing. Although the 2F to a single modulation presented to one eye was not reduced when a second frequency was added to the opposite eye, monocular IMCs were diminished when a frequency was added to the opposite eye. We conclude that 2Fs and IMCs are associated with different neural substrates. Results are consistent with a two pathway model with one pathway having a nonlinear filter prior to binocular combination, the other pathway having a nonlinearity following binocular linear summation. Implications of these data for binocular function are discussed.