Modules in connectomes of phase-synchronization comprise anatomically contiguous, functionally related regions

Modules in brain functional connectomes are essential to balancing segregation and integration of neuronal activity. Connectomes are the complete set of pairwise connections between brain regions. Non-invasive Electroencephalography (EEG) and Magnetoencephalography (MEG) have been used to identify modules in connectomes of phase-synchronization. However, their resolution is suboptimal because of spurious phase-synchronization due to EEG volume conduction or MEG field spread. Here, we used invasive, intracerebral recordings from stereo-electroencephalography (SEEG, N = 67), to identify modules in connectomes of phase-synchronization. To generate SEEG-based group-level connectomes affected only minimally by volume conduction, we used submillimeter accurate localization of SEEG contacts and referenced electrode contacts in cortical gray matter to their closest contacts in white matter. Combining community detection methods with consensus clustering, we found that the connectomes of phase-synchronization were characterized by distinct and stable modules at multiple spatial scales, across frequencies from 3 to 320 Hz. These modules were highly similar within canonical frequency bands. Unlike the distributed brain systems identified with functional Magnetic Resonance Imaging (fMRI), modules up to the high-gamma frequency band comprised only anatomically contiguous regions. Notably, the identified modules comprised cortical regions involved in shared repertoires of sensorimotor and cognitive functions including memory, language and attention. These results suggest that the identified modules represent functionally specialised brain systems, which only partially overlap with the brain systems reported with fMRI. Hence, these modules might regulate the balance between functional segregation and functional integration through phase-synchronization.