Modeling the Generation of Phase-Amplitude Coupling in Cortical Circuits: From Detailed Networks to Neural Mass Models

Phase-amplitude coupling (PAC), the phenomenon where the amplitude of a high frequency oscillation is modulated by the phase of a lower frequency oscillation, is attracting an increasing interest in the neuroscience community due to its potential relevance for understanding healthy and pathological information processing in the brain. PAC is a diverse phenomenon, having been experimentally detected in at least ten combinations of rhythms: delta-theta, delta-alpha, delta-beta, delta-gamma, theta-alpha, theta-beta, theta-gamma, alpha-beta, alpha-gamma, and beta-gamma. However, a complete understanding of the biophysical mechanisms generating this diversity is lacking. Here we review computational models of PAC generation that range from detailed models of neuronal networks, where each cell is described by Hodgkin-Huxley-type equations, to neural mass models (NMMs) where only the average activities of neuronal populations are considered. We argue that NMMs are an appropriate mathematical framework (due to the small number of parameters and variables involved and the richness of the dynamics they can generate) to study the PAC phenomenon.