Coherence fails to reliably capture inter-areal interactions in bidirectional neural systems with transmission delays

Accurately measuring and quantifying the underlying interactions between brain areas is crucial for understanding the flow of information in the brain. Of particular interest in the field of electrophysiology is the analysis and characterization of the spectral properties of these interactions. Coherence and Granger-Geweke causality are well-established, commonly used methods for quantifying inter-areal interactions, and are thought to reflect the strength of inter-areal interactions. Here we show that the application of both methods to bidirectional systems with transmission delays is problematic, especially for coherence. Under certain circumstances, coherence can be completely abolished despite there being a true underlying interaction. This problem occurs due to interference caused in the computation of coherence, and is an artifact of the method. We motivate an understanding of the problem through computational modelling and numerical simulations. In addition, we have developed two methods that can recover the true bidirectional interactions in the presence of transmission delays.