Brain Functional Connectivity Through Phase Coupling of Neuronal Oscillations: A Perspective From Magnetoencephalography

Magnetoencephalography has gained an increasing importance in systems neuroscience thanks to the possibility it offers of unraveling brain networks at time-scales relevant to behavior, i.e., frequencies in the 1-100 Hz range, with sufficient spatial resolution. In the first part of this review, we describe, in a unified mathematical framework, a large set of metrics used to estimate MEG functional connectivity at the same or at different frequencies. The different metrics are presented according to their characteristics: same-frequency or cross-frequency, univariate or multivariate, directed or undirected. We focus on phase coupling metrics given that phase coupling of neuronal oscillations is a putative mechanism for inter-areal communication, and that MEG is an ideal tool to non-invasively detect such coupling. In the second part of this review, we present examples of the use of specific phase methods on real MEG data in the context of resting state, visuospatial attention and working memory. Overall, the results of the studies provide evidence for frequency specific and/or cross-frequency brain circuits which partially overlap with brain networks as identified by hemodynamic-based imaging techniques, such as functional Magnetic Resonance (fMRI). Additionally, the relation of these functional brain circuits to anatomy and to behavior highlights the usefulness of MEG phase coupling in systems neuroscience studies. In conclusion, we believe that the field of MEG functional connectivity has made substantial steps forward in the recent years and is now ready for bringing the study of brain networks to a more mechanistic understanding.