The frequency of alpha oscillations: Task-dependent modulation and its functional significance.

Power (amplitude) and frequency are two important characteristics of EEG alpha oscillations (8-12 Hz). There is an extensive literature showing that alpha power can be modulated in a goal-oriented manner to either enhance or suppress sensory information processing. Only a few studies to date have examined the task-dependent modulation of alpha frequency. Instead, alpha frequency is often viewed as a trait variable, and used to characterize individual differences in cognitive functioning. We performed two experiments to examine the task-dependent modulation of alpha frequency and its functional significance. In the first experiment, high-density EEG was recorded from 21 participants performing a Sternberg working memory task. The results showed that: (1) during memory encoding, alpha frequency decreased with increasing memory load, whereas during memory retention and retrieval, alpha frequency increased with increasing memory load, (2) higher alpha frequency prior to the onset of probe was associated with longer reaction time, and (3) higher alpha frequency prior to the onset of cue or probe was associated with weaker early cue-evoked or probe-evoked neural responses. In the second experiment, simultaneous EEG-fMRI was recorded from 59 participants during resting state. An EEG-informed fMRI analysis revealed that the spontaneous fluctuations of alpha frequency, but not alpha power, were inversely associated with BOLD activity in the visual cortex. Taken together, these findings suggest that alpha frequency is task-dependent, may serve as an indicator of cortical excitability, and along with alpha power, provides more comprehensive indexing of sensory gating.