Generalized theory for current-source-density analysis in brain tissue

The current-source density (CSD) analysis is a widely used method in brain electrophysiology, but this method rests on a series of assumptions, namely that the surrounding extracellular medium is resistive and uniform, and in some versions of the theory, that the current sources are exclusively made by dipoles. Because of these assumptions, this standard model does not correctly describe the contributions of monopolar sources or of nonresistive aspects of the extracellular medium. We propose here a general framework to model electric fields and potentials resulting from current source densities, without relying on the above assumptions. We develop a mean-field formalism that is a generalization of the standard model and that can directly incorporate nonresistive (nonohmic) properties of the extracellular medium, such as ionic diffusion effects. This formalism recovers the classic results of the standard model such as the CSD analysis, but in addition, we provide expressions to generalize the CSD approach to situations with nonresistive media and arbitrarily complex multipolar configurations of current sources. We found that the power spectrum of the signal contains the signature of the nature of current sources and extracellular medium, which provides a direct way to estimate those properties from experimental data and, in particular, estimate the possible contribution of electric monopoles.