Functional integration across oscillation frequencies by cross-frequency phase synchronization.

Neuronal oscillations and their inter-areal synchronization may be instrumental in regulating neuronal communication in distributed networks. Several lines of research have, however, shown that cognitive tasks engage neuronal oscillations simultaneously in multiple frequency bands that have distinct functional roles in cognitive processing. Gamma oscillations (30-120 Hz) are associated with bottom-up processing, while slower oscillations in delta (1-4 Hz), theta (4-7 Hz), alpha (8-14 Hz) and beta (14-30 Hz) frequency bands may have roles in executive or top-down controlling functions, although also other distinctions have been made. Identification of the mechanisms that integrate such spectrally distributed processing and govern neuronal communication among these networks is crucial for understanding how cognitive functions are achieved in neuronal circuits. Cross-frequency interactions among oscillations have been recognized as a likely candidate mechanism for such integration. We advance here the hypothesis that phase-phase synchronization of neuronal oscillations in two different frequency bands, cross-frequency phase synchrony (CFS), could serve to integrate, coordinate and regulate neuronal processing distributed into neuronal assemblies concurrently in multiple frequency bands. A trail of studies over the past decade has revealed the presence of CFS among cortical oscillations and linked CFS with roles in cognitive integration. We propose that CFS could connect fast and slow oscillatory networks and thereby integrate distributed cognitive functions such as representation of sensory information with attentional and executive functions.