Theta rhythms, ≈3–8 Hz, have been found in many different parts of the brain. They are predominant in the rodent hippocampus, yet have also been described in the neocortex, primarily in frontal and parietal areas in relation to executive functions. Here, we show a ≈4-Hz theta rhythm in awake macaque monkey area V4 and primary visual cortex. This theta rhythm was spatially coextensive with visually induced gamma-band activity, and gamma power was modulated by theta phase. The strength of theta and of theta-rhythmic gamma modulation was markedly reduced by selective attention. Theta rhythmicity has been observed in microsaccade sequences, and microsaccades influence early visual activity. Yet, removing (the effects of) microsaccades did not influence the results.
Theta rhythms govern rodent sniffing and whisking, and human language processing. Human psychophysics suggests a role for theta also in visual attention. However, little is known about theta in visual areas and its attentional modulation. We used electrocorticography (ECoG) to record local field potentials (LFPs) simultaneously from areas V1, V2, V4, and TEO of two macaque monkeys performing a selective visual attention task. We found a ≈4-Hz theta rhythm within both the V1–V2 and the V4–TEO region, and theta synchronization between them, with a predominantly feedforward directed influence. ECoG coverage of large parts of these regions revealed a surprising spatial correspondence between theta and visually induced gamma. Furthermore, gamma power was modulated with theta phase. Selective attention to the respective visual stimulus strongly reduced these theta-rhythmic processes, leading to an unusually strong attention effect for V1. Microsaccades (MSs) were partly locked to theta. However, neuronal theta rhythms tended to be even more pronounced for epochs devoid of MSs. Thus, we find an MS-independent theta rhythm specific to visually driven parts of V1–V2, which rhythmically modulates local gamma and entrains V4–TEO, and which is strongly reduced by attention. We propose that the less theta-rhythmic and thereby more continuous processing of the attended stimulus serves the exploitation of this behaviorally most relevant information. The theta-rhythmic and thereby intermittent processing of the unattended stimulus likely reflects the ecologically important exploration of less relevant sources of information.