Effect of Stimulus Contrast and Visual Attention on Spike-Gamma Phase Relationship in Macaque Primary Visual Cortex

Brain signals often show rhythmic activity in the so-called gamma range (30-80 Hz), whose magnitude and center frequency are modulated by properties of the visual stimulus such as size and contrast, as well as by cognitive processes such as attention. How gamma rhythm can potentially influence cortical processing remains unclear; previous studies have proposed a scheme called phase coding, in which the intensity of the incoming stimulus is coded in the position of the spike relative to the rhythm. Using chronically implanted microelectrode arrays in the primary visual cortex (area V1) of macaques engaged in an attention task while presenting stimuli of varying contrasts, we tested whether the phase of the gamma rhythm relative to spikes varied as a function of stimulus contrast and attentional state. A previous study had found no evidence of gamma phase coding for either contrast or attention in V1, but in that study spikes and local field potential (LFP) were recorded from the same electrode, due to which spike-gamma phase estimation could have been biased. Further, the filtering operation to obtain LFP could also have biased the gamma phase. By analyzing spikes and LFP from different electrodes, we found a weak but significant effect of attention, but not stimulus contrast, on gamma phase relative to spikes. The results remained consistent even after correcting the filter induced lags, although the absolute magnitude of gamma phase shifted by up to similar to 15 degrees. Although we found a significant effect of attention, we argue that a small magnitude of phase shift as well as the dependence of phase angles on gamma power and center frequency limits a potential role of gamma in phase coding in V1.