Dynamics of Normalization Underlying Masking in Human Visual Cortex

Stimulus visibility can be reduced by other stimuli that overlap the same region of visual space, a process known as masking. Here we studied the neural mechanisms of masking in humans using source-imaged steady state visual evoked potentials and frequency-domain analysis over a wide range of relative stimulus strengths of test and mask stimuli. Test and mask stimuli were tagged with distinct temporal frequencies and we quantified spectral response components associated with the individual stimuli (self terms) and responses due to interaction between stimuli (intermodulation terms). In early visual cortex, masking alters the self terms in a manner consistent with a reduction of input contrast. We also identify a novel signature of masking: a robust intermodulation term that peaks when the test and mask stimuli have equal contrast and disappears when they are widely different. We fit all of our data simultaneously with family of a divisive gain control models that differed only in their dynamics. Models with either very short or very long temporal integration constants for the gain pool performed worse than a model with an integration time of ∼30 ms. Finally, the absolute magnitudes of the response were controlled by the ratio of the stimulus contrasts, not their absolute values. This contrast–contrast invariance suggests that many neurons in early visual cortex code relative rather than absolute contrast. Together, these results provide a more complete description of masking within the normalization framework of contrast gain control and suggest that contrast normalization accomplishes multiple functional goals.