An Investigation Into the Nonlinear Coupling Between CA1 Layers and the Dentate Gyrus

Although the activity from the dentate gyrus is known to have strong connections with other hippocampal layers, the functionality of these connections, that is, the degree to which it drives activity in the downstream regions of the hippocampus, is not well understood. This question is particularly relevant for mesoscale localfield potential (LFP) rhythms such as gamma oscillations. Following the hypothesis that fundamental features of the LFP are consistent with turbulent dynamics, we investigate the crosslayer relationship between the CA1 layers and the dentate gyrus as a function of running speed. In agreement with previous studies, same-layer spectral and bispectral analyses show that increasing input (rat speed) results in an increase of power and nonlinearity (phase coupling) between theta and gamma. The effectiveness of the connection between the 2 regions is investigated using cross-bicoherence analysis. Based on the turbulence interpretation of the evolution of spectra and bispectra as a function of the power input rate, we propose a measure for estimating the strength of the cross-frequency, cross-layer nonlinear forcing, that compares the magnitude of bicoherence (same-layer) and cross-bicoherence (cross-layer). Our results suggest that at moderate speeds gamma in CA1 is mainly driven by local theta, while the coupling of the CA1 gamma to the dentate-gyrus gamma becomes significant. Overall, these data are consistent with the hypothesis of theta-to-gamma energy cascade model for the organization of hippocampal LFP, with theta playing the role of a global pacemaker across the entire hippocampus while gamma is a local oscillation generated by through local anatomical connections.