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ten Oever, S.

Publications and source records attributed to ten Oever, S..

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Predictable Timing Enhances the Use of Predictable Content

As events unfold, they carry structure in both their timing and content, with these dimensions often interrelated. Most research has treated these dimensions separately, although processing them jointly may provide important computational benefits. It is an open question whether the brain processes a sequence's timing and predictable content structure independently, or integrates these two sources of information to guide prediction. We hypothesised that predictable timing would allow listeners to better exploit structure in the content, with this effect increasing as content structure becomes stronger. To test this, we presented tone sequences with pitch trajectories following a 1/f^{beta} structure, where {beta} determines how strongly the content of each tone persists, while independently varying the temporal periodicity of the sequence. Performance was quantified using response errors relative to outcome predictions of a Gaussian Process Regression model and a model-free measure of how strongly preceding tones influenced responses. Response error decreased and reliance on preceding tones increased with higher {beta} and periodic timing. Moreover, the influence of preceding tones showed an interaction between timing and content, such that the effect of periodicity grew with increasing {beta}. The response error showed no such interaction, although the effect of periodic timing was reliable only for the most structured sequences. Our findings suggest an interdependency between content and timing, where predictable timing helps listeners make greater use of predictable content.

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tACS induces state-dependent shifts in inhibitory phase preference consistent with phase precession

Phase-dependent modulation of motor inhibition suggests that oscillatory timing plays a key role in inhibitory control. Here we show that rhythmic brain stimulation can selectively alter this timing in a manner consistent with phase precession. Participants performed an anticipatory stop-signal task while receiving 20 Hz transcranial alternating current stimulation (tACS) over the pre-supplementary motor area, with stop signals presented at defined phases of the stimulation waveform. An additional control experiment with no tACS was conducted, in which the stop signal phase was derived post-hoc from the ongoing EEG. In both tACS and control sessions, stopping efficacy varied sinusoidally with beta phase, confirming that inhibitory control is intrinsically phase-dependent. During tACS, however, the phase associated with the strongest inhibition advanced progressively across the session by about 180{degrees}, and this effect was specific to failed-stop trials. No comparable shift occurred without stimulation. These findings indicate that rhythmic drive can bias the temporal phase at which inhibition is most effective when the brain is in an error-prone state, revealing a state-dependent, precession-like reorganization of inhibitory timing. We propose that tACS interacts with error-related plasticity to promote gradual phase advancement, placing the system within spike-timing-dependent learning windows, whereas intrinsic fluctuations alone appear insufficient to facilitate cumulative precession. More broadly, these results suggest that phase precession may represent a general coding principle extending beyond hippocampal circuits, enabling the motor system to adaptively adjust inhibitory control through oscillatory timing mechanisms. HighlightsO_LIBeta tACS over preSMA induces shifts in phase preference during motor inhibition C_LIO_LIPrecession-like phase shifts emerge selectively during failed but not successful stops C_LIO_LIPhase precession was absent in a control experiment without tACS C_LIO_LItACS might provide a method to probe phase precession mechanisms in cognition C_LI

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Pronoun resolution via reinstatement of referent-related activity in the delta band

Human language offers a variety of ways to create meaning, one of which is referring to entities, objects, or events in the world. One such meaning maker is understanding to whom or to what a pronoun in a discourse refers to. To understand a pronoun, the brain must access matching entities or concepts that have been encoded in memory from previous linguistic context. Models of language processing propose that internally stored linguistic concepts, accessed via exogenous cues such as phonological input of a word, are represented as (a)synchronous activities across a population of neurons active at specific frequency bands. Converging evidence suggests that delta band activity (1-3Hz) is involved in temporal and representational integration during sentence processing. Moreover, recent advances in the neurobiology of memory suggest that recollection engages reinstatement of neural dynamics that occurred during memory encoding. Integrating from these two research lines, we here predicted that neural dynamic patterns, especially in delta frequency range, underlying referential meaning representation would be reinstated during pronoun resolution. By leveraging neural decoding techniques (i.e., representation similarity analysis) on a magnetoencephalogram (MEG) dataset acquired during a naturalistic story-listening task, we provide evidence that delta-band activity underlies referential meaning representation. Our findings suggest that, during spoken language comprehension, endogenous linguistic representations such as referential concepts may be retrieved and represented via reinstatement of dynamic neural patterns.

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Task-dependent and automatic tracking of hierarchical linguistic structure

Linguistic phrases are tracked in sentences even though there is no clear acoustic phrasal marker in the physical signal. This phenomenon suggests an automatic tracking of abstract linguistic structure that is endogenously generated by the brain. However, all studies investigating linguistic tracking compare conditions where either relevant information at linguistic timescales is available, or where this information is absent altogether (e.g., sentences versus word lists during passive listening). It is therefore unclear whether tracking at these phrasal timescales is related to the content of language, or rather, is a consequence of attending to the timescales that happen to match behaviourally-relevant information. To investigate this question, we presented participants with sentences and word lists while recording their brain activity with MEG. Participants performed passive, syllable, word, and word-combination tasks corresponding to attending to rates they would naturally attend to, syllable-rates, word-rates, and phrasal-rates, respectively. We replicated overall findings of stronger phrasal-rate tracking measured with mutual information (MI) for sentences compared to word lists across the classical language network. However, in the inferior frontal gyrus (IFG) we found a task-effect suggesting stronger phrasal-rate tracking during the word-combination task independent of the presence of linguistic structure, as well as stronger delta-band connectivity during this task. These results suggest that extracting linguistic information at phrasal-rates occurs automatically with or without the presence of an additional task, but also that that IFG might be important for temporal integration across various perceptual domains.

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Frequency-specific transcranial neuromodulation of oscillatory alpha power alters and predicts human visuospatial attention performance

Unilateral transcranial alternating current stimulation (tACS) at alpha frequency modulates the locus of spatial attention. However, the neural mechanisms by which tACS influences spatial attention remain poorly understood. Here, we applied high-definition tACS at the individual alpha frequency (IAF), two control frequencies (IAF+/-2Hz) and sham to the left posterior parietal cortex and measured its effects on visuospatial attention performance as well as alpha power (using electroencephalography, EEG). Our results revealed a leftward lateralization of alpha power relative to sham. At a high value of leftward alpha lateralization, we also observed a leftward attention bias, which differed from sham. Moreover, the magnitude of the alpha lateralization effect predicted the attention bias. These effects occurred for tACS at IAF but not for the control frequencies. This suggests that tACS operates through oscillatory interactions with ongoing brain rhythms in line with the synchronization theory. Our results also highlight the importance of personalized stimulation protocols, especially in potential clinical settings.

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Does alpha phase modulate visual target detection? Three experiments with tACS phase-based stimulus presentation.

In recent years the influence of alpha (7-13 Hz) phase on visual processing has received a lot of attention. Magneto-/encephalography (M/EEG) studies showed that alpha phase indexes visual excitability and task performance. If occipital alpha phase is functionally relevant, the phase of occipital alpha-frequency transcranial alternating current stimulation (tACS) could modulate visual processing. Visual stimuli presented at different pre-determined, experimentally controlled, phases of the entraining tACS signal should then result in an oscillatory pattern of visual performance. We studied this in a series of experiments. In experiment one, we applied 10 Hz tACS to right occipital cortex (O2) and used independent psychophysical staircases to obtain contrast thresholds for detection of visual gratings in left or right hemifield, in six equidistant tACS phase conditions. In experiments two and three, tACS was at EEG-based individual peak alpha frequency. In experiment two, we measured detection rates for gratings with (pseudo-)fixed contrast levels. In experiment three, participants detected brief luminance changes in a custom-built LED device, at eight equidistant alpha phases. In none of the experiments did the primary outcome measure over phase conditions consistently reflect a one-cycle sinusoid as predicted. However, post-hoc analyses of reaction times (RT) suggested that tACS alpha phase did modulate RT in both experiments 1 and 2 (not measured in experiment 3). This observation is in line with the idea that alpha phase causally gates visual inputs through cortical excitability modulation.

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Left parietal tACS at alpha frequency induces a shift of visuospatial attention

BackgroundVoluntary shifts of visuospatial attention are associated with a lateralization of occipitoparietal alpha power (7-13Hz), i.e. higher power in the hemisphere ipsilateral and lower power contralateral to the locus of attention. Recent noninvasive neuromodulation studies demonstrated that alpha power can be experimentally increased using transcranial alternating current stimulation (tACS).\n\nObjective/HypothesisWe hypothesized that tACS at alpha frequency over the left parietal cortex induces shifts of attention to the left hemifield. However, spatial attention shifts not only occur voluntarily (endogenous), but also stimulus-driven (exogenous). In order to study the task-specificity of the potential effects of tACS on attentional processes, we administered three conceptually different spatial attention tasks.\n\nMethods36 healthy volunteers were recruited from an academic environment. In two seperate sessions, we applied either high-density tACS at 10Hz, or sham tACS, for 35-40 minutes to their left parietal cortex. We systematically compared performance on endogenous attention, exogenous attention, and stimulus detection tasks.\n\nResultsIn the Endogenous attention task, we found a greater leftward bias in reaction times during left parietal 10Hz tACS as compared to sham. There were no stimulation effects in the exogenous attention or stimulus detection task.\n\nConclusionThe study shows that high-density tACS at 10Hz can be used to modulate visuospatial attention performance. The tACS effect is task-specific, indicating that not all forms of attention are equally susceptible to the stimulation.

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