Cortical mapping of gamma oscillations in areas V1 and V4 of the macaque monkey

To characterize the temporal and spatial parameters of gamma activity evoked by visual stimuli in areas V1 and V4 of the monkey cortex, we recorded the electrocorticogram (ECoG) with an implanted array of 28 and 31 subdural electrodes placed over the surface of the operculum in two anesthetized monkeys. This intermediate level of recordings should help to bridge the gap between multiunit and scalp recordings. Both averaged and single-trial responses to small flashed stimuli, for which we varied the retinotopic position, the luminance and the color, were analyzed in the time-frequency domain using a wavelet-based decomposition of the signal. Large gamma oscillations (40-55 Hz), not phase locked to stimulus onset, were observed during the whole stimulus presentation, whereas visual evoked potentials (VEPs) were present mainly at stimulus onset and offset. Cortical mapping showed that both activities were restricted in spatial extent and followed the retinotopic organization of area V1 on the operculum, thus strongly suggesting they were generated in the underlying cortex. Oscillatory burst detection in single trials showed that one to two bursts lasting from 100 ms to 500 ms occurred in the first 500 ms following stimulus onset, and that bursts occurring during the subsequent phases of the response had a smaller amplitude and duration. Finally, we showed that gamma activity was stronger with higher luminances and for red than for green, yellow, or white stimuli of same luminance. In one animal we recorded gamma activity over area V4. This was of lower magnitude than the activity recorded over V1 and was delayed by 40 ms with respect to the beginning of gamma activity in V1, in contrast with the VEPs that were delayed by 20 ms only. Both gamma oscillations and early VEP followed the retinotopic organization of V4 over the prelunate gyrus. The results show that gamma oscillations are dependent upon the same parameters as the VEPs (retinotopic position, luminance, and color). However, the differences in the time course of VEPs and gamma activity (transient vs. sustained) suggests that these two responses may reflect different cell populations, different networks, or different firing modes.