Nonlinear Origin of SSVEP Spectra—A Combined Experimental and Modeling Study

Steady state visual evoked potentials (SSVEPs) are steady state oscillatory potentials elicited in the electroencephalogram (EEG) by flicker stimulation. The frequency of these responses maches the frequency of the stimulation and of its harmonics and subharmonics. In this study, we investigated the origin of the harmonic and subharmonic components of SSVEPs, which are not well understood. We applied both sine and square wave visual stimulation at 5 and 15 Hz to human subjects and analyzed the properties of the fundamental responses and harmonically related components. In order to interpret the results, we used the well-established neural mass model that consists of interacting populations of excitatory and inhibitory cortical neurons. In our study, this model provided a simple explanation for the origin of SSVEP spectra, and showed that their harmonic and subharmonic components are a natural consequence of the nonlinear properties of neuronal populations and the resonant properties of the modeled network. The model also predicted multiples of subharmonic responses, which were subsequently confirmed using experimental data.