Propagating patterns of intrinsic activity along macroscale gradients coordinate functional connections across the whole brain

The intrinsic activity of the human brain, observed with resting-state fMRI (rsfMRI) and functional connectivity, exhibits macroscale spatial organization such as resting-state networks (RSNs) and functional connectivity gradients (FCGs). Dynamic analysis techniques have shown that the time-averaged maps captured by functional connectivity are mere summaries of time-varying patterns with distinct spatial and temporal characteristics. A better understanding of these patterns might provide insight into aspects of the brain’s intrinsic activity that cannot be inferred by functional connectivity, RSNs or FCGs. Here, we describe three spatiotemporal patterns of coordinated activity across the whole brain obtained by averaging similar ~20-second-long segments of rsfMRI timeseries. In each of these patterns, activity propagates along a particular macroscale FCG, simultaneously across the cortical sheet and in most other brain regions. In some areas, like the thalamus, the propagation suggests previously-undescribed FCGs. The coordinated activity across areas is consistent with known tract-based connections, and nuanced differences in the timing of peak activity between brain regions point to plausible driving mechanisms. The magnitude of correlation within and particularly between RSNs is remarkably diminished when these patterns are regressed from the rsfMRI timeseries, a quantitative demonstration of their significant role in functional connectivity. Taken together, our results suggest that a few recurring patterns of propagating intrinsic activity along macroscale gradients give rise to and coordinate functional connections across the whole brain.