Cortical layers, rhythms and BOLD signals.

This review investigates how laminar fMRI can complement insights into brain function derived from the study of rhythmic neuronal synchronization. Neuronal synchronization in various frequency bands plays an important role in neuronal communication between brain areas, and it does so on the backbone of layer-specific interareal anatomical projections. Feedforward projections originate predominantly in supragranular cortical layers and terminate in layer 4, and this pattern is reflected in inter-laminar and interareal directed gamma-band influences. Thus, gamma-band synchronization likely subserves feedforward signaling. By contrast, anatomical feedback projections originate predominantly in infragranular layers and terminate outside layer 4, and this pattern is reflected in inter-laminar and interareal directed alpha- and/or beta-band influences. Thus, alpha-beta band synchronization likely subserves feedback signaling. Furthermore, these rhythms explain part of the BOLD signal, with independent contributions of alpha-beta and gamma. These findings suggest that laminar fMRI can provide us with a potentially useful method to test some of the predictions derived from the study of neuronal synchronization. We review central findings regarding the role of layer-specific neuronal synchronization for brain function, and regarding the link between neuronal synchronization and the BOLD signal. We discuss the role that laminar fMRI could play by comparing it to invasive and non-invasive electrophysiological recordings. Compared to direct electrophysiological recordings, this method provides a metric of neuronal activity that is slow and indirect, but that is uniquely non-invasive and layer-specific with potentially whole brain coverage.