Working memory readout varies with frontal theta rhythms.

Increasing evidence suggests that attention varies rhythmically, phase locked to ongoing cortical oscillations. Here, we report that the phase of theta oscillations (3-6 Hz) in the frontal eye field (FEF) is associated with the spatiotemporal variation of information readout from working memory (WM). Non-human primates were briefly shown a sample array of colored squares. A short time later, they viewed a test array and were rewarded for identifying which square changed color (the target). Behavioral performance varied systematically with theta phase at the time of test array onset, as well as with the target's location. This is consistent with theta "scanning" across the FEF and thus visual space from top to bottom. Theta was coupled, on opposing phases, to both spiking and beta (12-20 Hz). These results could be explained by a wave of activity that moves across the FEF, modulating the readout of information from WM.