Abstract
The functional relevance of brain oscillations in the alpha frequency range (8–13 Hz) has been repeatedly investigated through the use of rhythmic visual stimulation. The underlying mechanism of the steady-state visual evoked potential (SSVEP) measured in EEG during rhythmic stimulation, however, is not known. There are two hypotheses on the origin of SSVEPs: entrainment of brain oscillations and superposition of event-related responses (ERPs). The entrainment but not the superposition hypothesis justifies rhythmic visual stimulation as a means to manipulate brain oscillations, because superposition assumes a linear summation of single responses, independent from ongoing brain oscillations. Here, we stimulated participants with a rhythmic flickering light of different frequencies and intensities. We measured entrainment by comparing the phase coupling of brain oscillations stimulated by rhythmic visual flicker with the oscillations induced by arrhythmic jittered stimulation, varying the time, stimulation frequency, and intensity conditions. In line with a theoretical concept of entrainment (the so called Arnold tongue), we found the phase coupling to be more pronounced with increasing stimulation intensity as well as at stimulation frequencies closer to each participant's intrinsic frequency. Only inside the Arnold tongue did the conditions significantly differ from the jittered stimulation. Furthermore, even in a single sequence of an SSVEP, we found non-linear features (intermittency of phase locking) that contradict the linear summation of single responses, as assumed by the superposition hypothesis. Our findings provide unequivocal evidence that visual rhythmic stimulation entrains brain oscillations, thus validating the approach of rhythmic stimulation as a manipulation of brain oscillations.
Highlights
A relationship between ongoing alpha amplitudes and perception has been a subject of investigation since Berger’s findings in the late 1920s (Berger, 1929)
Outside the Arnold tongue, the two hypotheses were indiscernible, as they both predict that no entrainment would occur, and here we found that phase locking showed no significant difference
We showed for the first time in a systematic analysis that visual flickering stimulation results in entrainment of brain oscillations
Summary
A relationship between ongoing alpha amplitudes and perception has been a subject of investigation since Berger’s findings in the late 1920s (Berger, 1929). Alpha phase angles were found to correlate with behavior (Hanslmayr et al, 2007; Klimesch et al, 2007) These findings support the key role of brain oscillations in the perception process (Hanslmayr et al, 2005; Lakatos et al, 2008; Busch et al, 2009; Jensen and Mazaheri, 2010). It is yet unclear, whether oscillations reflect a fundamental mechanism of information processing or rather appear as an epiphenomenon (Buzsáki and Draguhn, 2004; Herrmann et al, 2015)
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