Transmission of stimulus-locked responses in two oscillators with bistable coupling

Numerous electroencephalography (EEG) and magnetoencephalography (MEG) studies aim at identifying the chronological order of activation of brain areas. This paper demonstrates that the timing sequence obtained with the gold standard for EEG/MEG analysis (averaging across trials) may not correlate at all with the actual transmission of a stimulus' effect within a pathway formed by connected brain areas. This is shown by studying transmission of stimulus-locked responses in a model that shares basic features with stimulated neuronal rhythms: in two phase oscillators with bistable coupling and noise one oscillator is stimulated. The model presents a mechanism that causes a response clustering, i.e., a switching between two different responses across trials, without extinction of the averaged response (calculated over all trials). Transmission times are calculated for all trials as well as for the two clusters separately with standard averaged responses and with a stochastic phase resetting analysis. The stochastic phase resetting analysis provides reliable estimates of the transmission time. In contrast, transmission times calculated by averaging across trials correspond to the phase difference in the different stable synchronized states (when calculated for the two clusters separately) or their weighted superposition (when calculated over all trials). The standard method does not detect the time elapsing during the transmission of the stimulus' action. The results presented here call into question many findings reported in the evoked response literature.