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Thut, G.

Publications and source records attributed to Thut, G..

17 recordsLinked to original sources

Working memory operations emerge from dynamic changes in neural subspace geometry

Working memory representations of multiple items are maintained within quasi-orthogonal neural subspaces, a mechanism thought to reduce interference between memory contents. Previous studies in both non-human and human primates have demonstrated the existence of such subspaces, but it remains unclear how cognitive operations are performed on the information they encode. One hypothesis proposes that these operations are implemented through dynamic changes in subspace geometry, yet direct empirical evidence for this idea in the context of working memory is lacking. Furthermore, the neural mechanisms underlying such geometric reconfigurations remain unknown, particularly the role of oscillatory and aperiodic neural activity. Here, we addressed these questions using simultaneous magneto- and electroencephalography (M/EEG) recordings from healthy human participants performing a multi-item, multi-feature delayed match-to-sample visual WM task. Neural subspaces were estimated using dimensionality reduction techniques, and their geometry was characterized by quantifying the orthogonality between subspaces, distances between memory representations and subspace shapes. Participants maintained orientation and shape information simultaneously while task demands required either deprioritizing one feature or updating the representation of the prioritized feature. We found that subspace geometry was flexibly reshaped according to task demands: prioritization expanded representational subspaces, whereas de-prioritization shrunk them. In addition, subspaces representing different feature domains were generally oblique and became more orthogonal only under high cognitive demands. Finally, these task-dependent changes in subspace geometry were largely driven by interactions between aperiodic neural activity and slow oscillations in the delta and theta frequency bands.

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Age differences in motivation drive performance during the sustained attention to response task

Young and older adults prioritise speed and accuracy differently during sustained attention tasks. While older adults generally show a preference of accuracy over speed, this is not always the case. The underlying factor behind this inconsistency may be motivational differences, with older participants compensating for a speed disadvantage with increased intrinsic motivation to perform well. We investigated this in a pre-registered study, using the Sustained aAttention to Response Task (SART) in young (n = 25, mean age = 19) and older adults (n = 25, mean age = 69.5). We matched participant accuracy by titrating response window length. Both groups achieved similar performance and strategy during the titration, enabling a comparison without confounds resulting from differences in default age-specific strategies. All participants were then monetarily encouraged to perform better in terms of accuracy. Both groups responded with an adaptive strategy of slowed reaction times and enhanced accuracy, but the young participants improved much more, outperforming older adults, and reversing the speed-accuracy strategies that are typically observed. In addition, older participants reported higher baseline levels of motivation alongside a reduced motivation to alter performance for money. So, while the older participants could match young participant performance in titration due to their higher baseline motivational levels, the young participants improved much more than older adults in response to the monetary incentive. From these findings we argue that older adults are intrinsically motivated to do well on tasks whereas younger age groups perform optimally only after incentivisation.

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EEG markers of vigilance, task-induced fatigue and motivation during sustained attention: Evidence for decoupled alpha- and beta-signatures

Reduced vigilance can be captured in measures of attentional lapses in sustained attention tasks, but just how these lapses relate to task-induced fatigue and motivation to maintain optimal performance is unclear. We used the sustained attention to response task (SART) to induce fatigue, and manipulated motivation levels for the last block of the task in young and older participants (N = 34), while recording EEG to track electrophysiological markers of vigilance change, fatigue and motivation. Despite significant increases in subjective fatigue and mind wandering over 45 minutes, no vigilance decline was observed. However, the age groups differed markedly in their response strategies from the outset (adopting distinct speed-accuracy trade-off strategies) with faster/more erroneous responses in the younger and slower/more accurate responses in the older participants. The subjective rises in fatigue/mind wandering were coupled with an increase in pre-stimulus alpha-power, whereas the post-stimulus activity revealed two distinguishable beta signatures: a fronto-central topography as a marker of response strategy and a fronto-parietal distribution modulated by motivation per se. Our results thus show three distinct neural patterns underpinning the effects of fatigue, response strategy and motivation and suggest a (motivational) cognitive control mechanism behind resetting of performance decrement, independent of persistent fatigue.

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Working memory enhancement using real-time phase-tuned transcranial alternating current stimulation

BackgroundPrior work has shown that transcranial alternating current stimulation (tACS) of parietooccipital alpha oscillations (8 - 14 Hz) can modulate working memory (WM) performance as a function of the phase lag to endogenous oscillations. However, leveraging this effect using real-time phase-tuned tACS was not feasible so far due to stimulation artifacts. Objectives/HypothesisWe aimed to develop a system that tracks and adapts the phase lag between tACS and ongoing parietooccipital alpha oscillations in real-time. We hypothesized that such real-time phase-tuned tACS enhances working memory performance, depending on the phase lag. MethodsWe developed real-time phase-tuned closed-loop amplitude-modulated tACS (CLAM-tACS) targeting parietooccipital alpha oscillations. CLAM-tACS was applied at six different phase lags relative to ongoing alpha oscillations while participants (N = 21) performed a working memory task. To exclude that behavioral effects of CLAM-tACS were mediated by other factors such as sensory co-stimulation, a second group of participants (N = 25) received equivalent stimulation of the forehead. ResultsWM accuracy improved in a phase lag dependent manner (p < 0.05) in the group receiving parietooccipital stimulation, with the strongest enhancement observed at 330{degrees} phase lag between tACS and ongoing alpha oscillations (p < 0.01, d = 0.976). Moreover, across participants, modulation of frontoparietal alpha oscillations correlated both in amplitude (p < 0.05) and phase (p < 0.05) with the modulation of WM accuracy. No such effects were observed in the control group receiving frontal stimulation. ConclusionsOur results demonstrate the feasibility and efficacy of real-time phase-tuned CLAM-tACS in modulating both brain activity and behavior, thereby paving the way for further investigation into brain-behavior relationships and the exploration of innovative therapeutic applications.

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Top-down and bottom-up interactions rely on nested brain oscillations.

Adaptive visual processing is enabled through the dynamic interplay between top-down and bottom-up (feedback/feedforward) information exchange, presumably propagated through brain oscillations. Here we causally tested for the oscillatory mechanisms governing this interaction in the visual system. Using concurrent TMS-EEG, we emulated top-down signals by a single TMS-pulse over the Frontal Eye Field (right-FEF), while manipulating the strength of sensory input through the presentation of moving concentric gratings (compared to a control-TMS site). FEF-TMS without sensory input led to a top-down controlled occipital phase-realignment, alongside higher fronto-occipital phase-connectivity, in the alpha/beta-band. Sensory input in the absence of FEF-TMS increased occipital gamma activity. Crucially, testing the interaction between top-down and bottom-up processes (FEF-TMS during sensory input) revealed an increased nesting of the bottom-up gamma activity in the alpha/beta-band cycles. This establishes a causal link between phase-to-power coupling and top-down modulation of feedforward signals, providing novel mechanistic insights into how attention interacts with sensory input at the neural level, shaping rhythmic sampling.

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Characterising time-on-task effects on oscillatory and aperiodic EEG components and their co-variation with visual task performance.

Research on brain-behaviour relationships often makes the implicit assumption that these derive from a co-variation of stochastic fluctuations in brain activity and performance across trials of an experiment. However, challenging this assumption, oscillatory brain activity, as well as indicators of performance, such as response speed, can show systematic trends with time on task. Here we tested whether time-on-task trends explain a range of relationships between oscillatory brain activity and response speed, accuracy as well as decision confidence. Thirty-six participants performed 900 trials of a two-alternative forced choice visual discrimination task with confidence ratings. Pre- and post-stimulus spectral power (1-40Hz) and aperiodic (i.e., non-oscillatory) components were compared across blocks of the experimental session and tested for relationships with behavioural performance. We found that time-on-task effects on oscillatory EEG activity were primarily localised within the alpha band, with alpha power increasing and peak alpha frequency decreasing over time, even when controlling for aperiodic contributions. Aperiodic, broadband activity on the other hand did not show time-on-task effects in our data set. Importantly, time-on-task effects in alpha frequency and power explained variability in single-trial reaction times, and controlling for time-on-task effectively removed these relationships. However, time-on-task effects did not affect other EEG signatures of behavioural performance, including post-stimulus predictors of single-trial decision confidence. Our results dissociate alpha-band brain-behaviour relationships that can be explained away by time-on-task from those that remain after accounting for it - thereby further specifying the potential functional roles of alpha in human visual perception.

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Probing sustained attention and fatigue across the lifespan

Trait fatigues reflects tiredness that persists throughout a prolonged period, whereas state fatigue is defined to be short term after "intense and/or prolonged effort". We investigated the impact of prolonged sustained attention (using the SART task) on both trait and state fatigue levels in the general population. A JsPsych online version of the SART task was undertaken by 115 participants, stratified across the whole adult lifespan. While pre-task trait fatigue was a strong indicator of the initial state fatigue levels, undergoing the task itself induced an increase in reported subjective state fatigue, as well as reduced energy. Consistent with this finding, greater subjective state fatigue levels were associated with reduced accuracy. In addition, age was the best predictor of inter-participant accuracy (the older the participants, the greater the accuracy), and learning (i.e., task duration reducing reaction times). Moreover, a ceiling effect occurred where participants with higher trait fatigue did not experience greater state fatigue changes relative to those with low trait scores. In summary, we found improved accuracy in older adults, as well as a tight coupling between state fatigue and SART performance decline (in an online environment). The findings warrant further investigation into fatigue as a dynamic, task-dependent state and into SART task performance as an objective measure and inducer of fatigue.

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Developmental changes in individual alpha frequency: Recording EEG data during public engagement events

Statistical power in cognitive neuroimaging experiments is often very low. Low sample size can reduce the likelihood of detecting real effects (false negatives) and increase the risk of detecting non-existing effects by chance (false positives). Here we document our experience of leveraging a relatively unexplored method of collecting a large sample size for simple electroencephalography (EEG) studies: by recording EEG in the community during public engagement and outreach events. We collected data from 346 participants (189 females, age range 6-76 years) over 6 days, totalling 29 hours, at local science festivals. Alpha activity (6-15 Hz) was filtered from 30 seconds of signal, recorded from a single electrode placed between the occipital midline (Oz) and inion (Iz) while participants rested with their eyes closed. A total of 289 good quality datasets were obtained. Using this community-based approach, we were able to replicate controlled, lab-based findings: IAF increased during childhood, reaching a peak frequency of 10.28 Hz at 28.1 years old, and slowed again in middle and older age. Total alpha power decreased linearly, but the aperiodic-adjusted alpha power did not change over the lifespan. Aperiodic slopes and intercepts were highest in the youngest participants. There were no associations between these EEG indexes and self-reported fatigue, measured by the Multidimensional Fatigue Inventory. Finally, we present a set of important considerations for researchers who wish to collect EEG data within public engagement and outreach environments.

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The Timing of Cortical Activation in Associator Grapheme-Colour Synaesthetes using MEG

Grapheme-colour synaesthetes experience an anomalous form of perception in which graphemes systematically induce specific colour concurrents in their minds eye ("associator" type). Although grapheme-colour synaesthesia has been well characterized behaviourally, its neural mechanisms remain largely unresolved. There are currently several competing models, which can primarily be distinguished according to the anatomical and temporal predictions of synaesthesia-inducing neural activity. The first main model (Cross-Activation/ Cascaded Cross-Tuning) posits early recruitment of occipital colour areas in the initial feed-forward sweep of brain activity. The second (Disinhibited Feedback) posits: (i) later involvement of a multisensory convergence zone (for example, in parietal cortices) after graphemes have been processed in their entirety; and (ii) subsequent feedback to early visual areas (i.e., occipital colour areas). In this study, we examine both the timing and anatomical correlates of associator grapheme-colour synaesthetes (n=6) using MEG. Using unbiased analysis methods with little a priori assumptions, we applied Independent Component Analysis (ICA) on a single-subject level to identify the dominant patterns of the induced, synaesthetic percept. We observed evoked activity that significantly dissociates between synaesthesia-inducing and non-inducing conditions at approximately 190 ms following grapheme presentation. This effect is present in grapheme-colour synaesthetes, but not in matched controls, and exhibits an occipito-parietal topology localised consistently across individuals to extrastriate visual cortices and to superior parietal lobes. Due to the observed timing of this evoked activity and its localisation, our results support a model predicting relatively late synaesthesia-inducing activity, more akin to the Disinhibited Feedback model. In light of previous findings, our study suggest that differential timing and neural mechanisms may account for associator grapheme-colour synaesthetes, as compared to projectors. Highlights- first MEG study to-date examining associator grapheme-colour synaesthetes - we use single-subj. ICA to dissociate synaesthesia-inducing vs. non-inducing activity - we observe evoked activity [~]190 ms following grapheme presentation effects localize consistently to extrastriate visual and superior parietal cortices - the observed timing is consistent with a model of synaesthesia supporting relatively late activity of synaesthesia-inducing stimuli - our findings suggest that differential mechanisms may underlie different sub-types of grapheme-colour synaesthesia

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Coupling of pupil- and neuronal population dynamics reveals diverse influences of arousal on cortical processing

Fluctuations in arousal, controlled by subcortical neuromodulatory systems, continuously shape cortical state, with profound consequences for information processing. Yet, how arousal signals influence cortical population activity in detail has so far only been characterized for a few selected brain regions. Traditional accounts conceptualize arousal as a homogeneous modulator of neural population activity across the cerebral cortex. Recent insights, however, point to a higher specificity of arousal effects on different components of neural activity and across cortical regions. Here, we provide a comprehensive account of the relationships between fluctuations in arousal and neuronal population activity across the human brain. Exploiting the established link between pupil size and central arousal systems, we performed concurrent magnetoencephalographic (MEG) and pupillographic recordings in a large number of participants, pooled across three laboratories. We found a cascade of effects relative to the peak timing of spontaneous pupil dilations: Decreases in low-frequency (2-8 Hz) activity in temporal and lateral frontal cortex, followed by increased high-frequency (>64 Hz) activity in mid-frontal regions, followed by monotonic and inverted-U relationships with intermediate frequency-range activity (8-32 Hz) in occipito-parietal regions. Pupil-linked arousal also coincided with widespread changes in the structure of the aperiodic component of cortical population activity, indicative of changes in the excitation-inhibition balance in underlying microcircuits. Our results provide a novel basis for studying the arousal modulation of cognitive computations in cortical circuits.

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EEG Alpha Power Predicts the Temporal Sensitivity of Multisensory Integration

Pre-stimulus EEG oscillations, especially in the alpha range (8-13 Hz), can affect the integration of stimulus features into a coherent percept. The effects of alpha power are often explained in terms of alphas inhibitory functions, whereas effects of alpha frequency have bolstered theories of discrete perceptual cycles, where the length of a cycle, or window of integration, is determined by alpha frequency. Such studies typically employ visual detection paradigms with near-threshold or even illusory stimuli. It is unclear whether such results generalize to above-threshold stimuli. Here, we recorded electroencephalography, while measuring temporal discrimination sensitivity in a temporal order judgement task using above-threshold auditory and visual stimuli. We tested whether pre-stimulus oscillations predict audio-visual temporal discrimination sensitivity on a trial-by-trial basis. By applying a jackknife procedure to link single-trial pre-stimulus oscillatory power and instantaneous frequency to psychometric measures, we identified two highly overlapping clusters over posterior sites. One where lower alpha power was associated with higher temporal sensitivity of audiovisual discrimination, and another where higher instantaneous alpha-frequency predicted higher temporal discrimination sensitivity. A follow-up analysis revealed that these effects were not independent, and that the effect of instantaneous frequency could be explained by power modulations in the lower alpha band. These results suggest that temporal sensitivity for above-threshold multisensory stimuli changes spontaneously from moment to moment and is likely related to fluctuations in cortical excitability. Moreover, our results caution against interpreting instantaneous frequency effects as independent from power effects if these effects overlap in time and space.

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Subjective perceptual experience tracks the neural signature of sensory evidence accumulation during decision formation

How neural representations of low-level visual information are accessed by higher-order processes to inform decisions and give rise to conscious experience is a longstanding question. Research on perceptual decision making has revealed a late event-related EEG potential (the Centro-Parietal Positivity, CPP) to be a correlate of the accumulation of sensory evidence. We tested to what extent this evidence accumulation signal is driven by externally presented (physical) versus internally experienced (subjective) sensory evidence. The results show that the known relationship between external evidence and the evidence accumulation signal (reflected in the CPP amplitude) is mediated by the level of subjective awareness. Additionally, the CPP closely tracks the subjective perceptual evidence during both correct and incorrect trials. Hence, a remarkably close relationship exists between the evidence accumulation process (i.e. CPP) and subjective perceptual experience, suggesting that neural decision processes and components of conscious experience are tightly linked.

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Stimulus-driven brain rhythms within the alpha band: The attentional-modulation conundrum

Two largely independent research lines use rhythmic sensory stimulation to study visual processing. Despite the use of strikingly similar experimental paradigms, they differ crucially in their notion of the stimulus-driven periodic brain responses: One regards them mostly as synchronised (entrained) intrinsic brain rhythms; the other assumes they are predominantly evoked responses (classically termed steady-state responses, or SSRs) that add to the ongoing brain activity. This conceptual difference can produce contradictory predictions about, and interpretations of, experimental outcomes. The effect of spatial attention on brain rhythms in the alpha-band (8 - 13 Hz) is one such instance: alpha-range SSRs have typically been found to increase in power when participants focus their spatial attention on laterally presented stimuli, in line with a gain control of the visual evoked response. In nearly identical experiments, retinotopic decreases in entrained alpha-band power have been reported, in line with the inhibitory function of intrinsic alpha. Here we reconcile these contradictory findings by showing that they result from a small but far-reaching difference between two common approaches to EEG spectral decomposition. In a new analysis of previously published human EEG data, recorded during bilateral rhythmic visual stimulation, we find the typical SSR gain effect when emphasising stimulus-locked neural activity and the typical retinotopic alpha suppression when focusing on ongoing rhythms. These opposite but parallel effects suggest that spatial attention may bias the neural processing of dynamic visual stimulation via two complementary neural mechanisms.\n\nSIGNIFICANCE STATEMENTAttending to a visual stimulus strengthens its representation in visual cortex and leads to a retinotopic suppression of spontaneous alpha rhythms. To further investigate this process, researchers often attempt to phase-lock, or entrain, alpha through rhythmic visual stimulation under the assumption that this entrained alpha retains the characteristics of spontaneous alpha. Instead, we show that the part of the brain response that is phase-locked to the visual stimulation increased with attention (in line with steady-state evoked potentials), while the typical suppression was only present in non-stimulus-locked alpha activity. The opposite signs of these effects suggest that attentional modulation of dynamic visual stimulation relies on two parallel cortical mechanisms - retinotopic alpha suppression and increased temporal tracking.

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Predictive entrainment of natural speech through two fronto-motor top-down channels

Natural communication between interlocutors is enabled by the ability to predict upcoming speech in a given context. Previously we showed that these predictions rely on a fronto-motor top-down control of low-frequency oscillations in auditory-temporal brain areas that track intelligible speech. However, a comprehensive spatio-temporal characterisation of this effect is still missing. Here, we applied transfer entropy to source-localised MEG data during continuous speech perception. First, at low frequencies (1-4 Hz, brain delta phase to speech delta phase), predictive effects start in left fronto-motor regions and progress to right temporal regions. Second, at higher frequencies (14-18 Hz, brain beta power to speech delta phase), predictive patterns show a transition from left inferior frontal gyrus via left precentral gyrus to left primary auditory areas. Our results suggest a progression of prediction processes from higher-order to early sensory areas in at least two different frequency channels.

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Frequency and power of human alpha oscillations drift systematically and independently with time-on-task

Oscillatory neural activity is a fundamental characteristic of the mammalian brain spanning multiple levels of spatial and temporal scale. Current theories of neural oscillations and analysis techniques employed to investigate their functional significance are based on an often implicit assumption: In the absence of experimental manipulation, the spectral content of any given EEG- or MEG-recorded neural oscillator remains approximately stationary over the course of a typical experimental session (~1 hour), spontaneously fluctuating only around its dominant frequency. Here, we examined this assumption for ongoing neural oscillations in the alpha-band (8:13 Hz). We found that alpha peak frequency systematically decreased over time, while alpha-power increased. Intriguingly, these systematic changes showed partial independence of each other: Statistical source separation (independent component analysis) revealed that while some alpha components displayed concomitant power increases and peak frequency decreases, other components showed either unique power increases or frequency decreases. Interestingly, we also found these components to differ in frequency. Components that showed mixed frequency/power changes oscillated primarily in the lower alpha-band (~8-10Hz), while components with unique changes oscillated primarily in the higher alpha-band (~9-13Hz). Our findings provide novel clues on the time-varying intrinsic properties of large-scale neural networks as measured by M/EEG, with implications for the analysis and interpretation of studies that aim at identifying functionally relevant oscillatory networks or at driving them through external stimulation.

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No changes in parieto-occipital alpha during neural phase locking to visual quasi-periodic theta-, alpha-, and beta-band stimulation

Recent studies have probed the role of the parieto-occipital alpha rhythm (8 - 12 Hz) in human visual perception through attempts to drive its neural generators. To that end, paradigms have used high-intensity strictly-periodic visual stimulation that created strong predictions about future stimulus occurrences and repeatedly demonstrated perceptual consequences in line with an entrainment of parieto-occipital alpha. Our study, in turn, examined the case of alpha entrainment by non-predictive low-intensity quasi-periodic visual stimulation within theta-(4 - 7 Hz), alpha-(8 - 13 Hz) and beta (14 - 20 Hz) frequency bands, i.e. a class of stimuli that resemble the temporal characteristics of naturally occurring visual input more closely. We have previously reported substantial neural phase-locking in EEG recording during all three stimulation conditions. Here, we studied to what extent this phase-locking reflected an entrainment of intrinsic alpha rhythms in the same dataset. Specifically, we tested whether quasi-periodic visual stimulation affected several properties of parieto-occipital alpha generators. Speaking against an entrainment of intrinsic alpha rhythms by non-predictive low-intensity quasi-periodic visual stimulation, we found none of these properties to show differences between stimulation frequency bands. In particular, alpha band generators did not show increased sensitivity to alpha band stimulation and Bayesian inference corroborated evidence against an influence of stimulation frequency. Our results set boundary conditions for when and how to expect effects of entrainment of alpha generators and suggest that the parieto-occipital alpha rhythm may be more inert to external influences than previously thought.

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Entrained audiovisual speech integration implemented by two independent computational mechanisms: Redundancy in left posterior superior temporal gyrus and Synergy in left motor cortex

Information integration is fundamental to many aspects of human behavior, and yet its neural mechanism remains to be understood. For example, during face-to-face communication we know that the brain integrates the auditory and visual inputs but we do not yet understand where and how such integration mechanisms support speech comprehension. Here we show that two independent mechanisms forge audiovisual representations for speech comprehension in different brain regions. With a novel information theoretic measure, we found that theta (3-7 Hz) oscillations in the posterior superior temporal gyrus/sulcus (pSTG/S) code speech information that is common (i.e. redundant) to the auditory and visual inputs whereas the same oscillations in left motor and inferior temporal cortex code synergistic information between the same inputs. Importantly, redundant coding in the left pSTG/S and synergistic coding in the left motor cortex predict behavior - i.e. speech comprehension performance. Our findings therefore demonstrate that processes classically described as integration effectively reflect independent mechanisms that occur in different brain regions to support audiovisual speech comprehension.

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