Dance to the rhythm, cautiously: Isolating unique indicators of oscillatory entrainment

The idea of entrainment of neural oscillations to temporal structure has become a central theory for attentional selection in time [1,2]. In our study [3], we separated unique electroencephalogram (EEG) signatures of oscillatory entrainment to rhythmic streams from those (specifically, inter-trial phase coherence [ITPC]) that overlap with other mechanisms of prediction. Obleser, Henry, and Lakatos’s (OHL’s) comment [4] highlights important issues regarding the study of entrainment mechanisms. We share OHL’s main message: rhythms may be complex and not necessarily isochronous. In fact, finding a stream devoid of any regularity is challenging. This is probably why entrainment mechanisms are functionally important! Yet we should not see entrainment mechanisms as omnipresent, nor should we render them irrefutable. Below, we reflect on some of OHL’s concerns regarding our study. Oscillatory entrainment predicts increased ITPC for rhythms [2,5], as both our and OHL’s models show. However, taking ITPC as evidence for entrainment, without ruling out other explanations, would be making the logical fallacy of "affirming the consequence" (a!b does not imply that b!a). Our study [3] thus asked which electrophysiological phenomena are uniquely associated with entrainment to rhythms instead of with general temporal prediction mechanisms that also operate in rhythmic contexts [6]. We conjectured that if ITPC reflects entrainment it should be sensitive to the level of regularity. However, we found that reducing the level of temporal regularity of streams did not reduce ITPC when predictability was preserved but only did so if predictability was also reduced. OHL’s argument that this is explained by predictions-related climbing neuronal activity (CNA; e.g., the contingent negative variation) only recapitulates one of our main arguments [3]. Given this fact, can a repeated-CNA explanation be ruled out from previous ITPC findings, especially in designs with no warning signal? Using near-threshold stimuli [7] reduces onset-responses–driven ITPC but not anticipatory activity. OHL point out (like we did [3]) that our repeated-interval condition (RIC) was not arrhythmic, explaining observed ITPC levels. Critically though, the RIC was not designed to be arrhythmic but to be less periodic than the isochronous condition (IsoC), which should diminish entrainment signatures. For perceptual judgments evidence (cf. OHL’s demo) that this was achieved, see Fig 1. However, in accordance with OHL’s appropriate assertion that rhythmicity is ill defined, we worry that resorting to subjective judgment falls into the same ambiguity of periodicity/predictability that motivated our study. Preferably, the degree to which a sequence is conducive to entrainment of a given oscillation, well defined by frequency and phase, could be objectively quantified. In fact, OHL’s model, like ours, predicts a difference in phase (their figure 1b) and a lower ITPC (their figure 1c showing the effect of stimulus onset asynchrony