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Melloni, L.

Publications and source records attributed to Melloni, L..

At least 19 recordsLinked to original sources

Learned Audiovisual Predictability Modulates Depth-Resolved Hippocampal-Cortical Functional Coupling

Perception reflects the dynamic integration of prior expectations, uncertainty, and incoming sensory evidence, but the neural circuitry supporting these computations remains debated. Current theories implicate coordinated interactions between the hippocampus and neocortex, potentially at the level of hippocampal subfields and cortical layers. Evidence in humans has come primarily from the visual domain, while less is known about how these interactions support learned auditory predictions. Using submillimetre 7T fMRI, we investigated depth-resolved functional coupling between hippocampal subfields and temporal and frontal cortical regions during audiovisual associative prediction. Participants learned deterministic associations between visual cues and auditory syllables, allowing us to compare auditory processing in predictable and unpredictable contexts. Predictability-related connectivity changes showed a spatially structured pattern involving DG/CA3-pSTG and CA1-inferior frontal regions, with significant edge-wise effects resolved across cortical depths. These findings provide fine-grained evidence that learned auditory predictability modulates hippocampal-cortical functional interactions across temporo-frontal systems.

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Long-timescale dynamics shape visual conscious awareness

Consciousness is mostly studied as if it were composed of independent snapshots, aggregating reports over trials and experimental contexts, disregarding any temporal aspect. However, almost nothing is known about how conscious experience spontaneously fluctuates in time. To shed light on the temporal dynamics of conscious awareness, we evaluated the temporal autocorrelation of subjective visibility/confidence reports and objective performance across 20 datasets (N= 335). Two widely used measures of consciousness dissociate in temporal autocorrelation under full spatial attention: subjective measures followed an exponential decay function with long timescales (> 5s), whereas objective accuracy exhibited a slow linear decay function. But both measures share a common underlying temporal fluctuation dynamics, suggesting that the strong autocorrelation in subjective measures cannot be fully attributed to response bias or criterion shift, but might reflect a genuinely aperiodic fluctuation of conscious awareness. Causal transcranial magnetic stimulation (TMS) suggests that this temporal dependency of subjective measures is read out in the dorsolateral prefrontal cortex (DLPFC), where a similarly long neural timescale has been demonstrated. Together, these findings establish a temporal constraint for neural models of conscious access: candidate mechanisms should explain not only whether a stimulus becomes consciously accessible, but also how the probability of conscious access evolves over time.

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Dynamic construction of subjective time through statistical learning of event structure

The perception of time is elastic, often deviating from physical intervals depending on how experience is structured. Yet, what determines how subjective time is constructed remains debated. Here, we tested whether perceived ongoing time is actively constructed from learned event representations rather than a dedicated internal clock. Using a novel pause-adjustment task across three statistical learning experiments, we measured temporal distortions during continuous listening to structured versus unstructured syllable streams. The presence of event structure systematically warped time: pauses were perceived as longer between pseudowords and shorter within pseudowords. This bidirectional temporal warping emerged online and remained stable across pause durations. Enriching these events with semantic meaning eliminated boundary-related dilation while preserving within-event compression. Moreover, physiological tracking of event structure, indexed by pupil dynamics, dissociated from the magnitude of temporal warping. These findings show that subjective time is constructed from hierarchical event representations and depends not only on where events are segmented, but also on how they are represented.

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Multiple task-demands flexibly optimize neural geometry in human ventral temporal cortex

A hallmark of human intelligence is the ability to perform multiple tasks immediately upon instruction. Yet, the neural processes that implement such flexibility remain unclear. Using intracranial electrophysiology in epilepsy patients, we examined how representational geometries evolve as participants switched among three tasks - individuation, categorization, and conceptualization - on a trial-by-trial basis. Although tasks differed in the required representational geometries, neural representations across cortex were not immediately optimized following task cues. Instead, task-tailored geometries emerged gradually over successive stimulus repetitions within a trial. Ventral temporal cortex was the only region to exhibit task-specific adjustments for all three tasks, dynamically transforming within- and between-category distances in line with task demands. Importantly, these gradual representational changes were behaviorally relevant, predicting trial-level performance. Our results show that behavioral flexibility is supported by incremental, task-dependent refinement of representational geometry already within sensory cortex - far earlier in the processing hierarchy than previously thought.

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Correlations of neural predictability and information transfer in cortex and their relation to predictive coding

Predictive-coding like theories agree in describing top-down communication through the cortical hierarchy as a transmission of predictions generated by internal models of the inputs. With respect to the bottom-up connections, however, these theories differ in the neural processing strategies suggested for updating the internal model. Some theories suggest a coding strategy where unpredictable inputs, i.e., those not captured by the internal model, are passed on through the cortical hierarchy, whereas others claim that the predictable part of the inputs is passed on. Here, we addressed which neural coding strategy is employed in cortico-cortical connections using an information-theoretic approach. Our framework allows for quantifying two core aspects of both strategies, namely, predictability of inputs and information transfer, through local active information storage and local transfer entropy, respectively. A previous study on the neural processing of retinal ganglion cells connected to the lateral geniculate nucleus showed a coding for predictable information, captured by an increase in the information transfer with the predictability of inputs. Here, we further investigate predictive coding strategies at the cortical level. In particular, we analyzed LFP activity obtained from intracranial EEG recordings in humans and spike recordings from mouse cortex. We detected cortico-cortical connections with increasing information transfer with the predictability of inputs in recorded channels from frontal, parietal and temporal areas in human cortex. In the mouse visual system, we detected connections exhibiting both an increase and decrease in the information transfer with input predictability, although the former was predominant. Our evidence supports the presence of both predictive coding strategies at the cortical level, with a potential predominance of encoding for predictable information. SummaryThe ability of the brain to infer the hidden causes of sensory experiences has been conceptualized within the computational framework of predictive coding. This framework explains perceptual inference and learning as a process of constantly updating an internal model of the world. Predictive coding describes cortical activity as a communication of sensory evidence and predictions generated from prior expectations. While different views of predictive coding agree on the communication of prior expectations throughout cortex, they differ in how internal expectations are updated. One view states that, to update the internal model, the cortex propagates the mismatch between the expected neural activity and the actual neural response to sensory stimuli. In contrast, another view suggests that the cortex propagates the match between the expected neural activity and the actual neural response. In this work, we were able to tease apart these two views, both in human cortex and the mouse visual system, using information theory. We observed that the brain predominantly propagates expected information, i.e., the match between prior expectations and incoming sensory inputs.

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The "Podcast" ECoG dataset for modeling neural activity during natural language comprehension

Naturalistic electrocorticography (ECoG) data are a rare but essential resource for studying the brains linguistic capabilities. ECoG offers a high temporal resolution suitable for investigating processes at multiple temporal timescales and frequency bands. It also provides broad spatial coverage, often along critical language areas. Here, we share a dataset of nine ECoG participants with 1,330 electrodes listening to a 30-minute audio podcast. The richness of this naturalistic stimulus can be used for various research endeavors, from auditory perception to semantic integration. In addition to the neural data, we extract linguistic features of the stimulus ranging from phonetic information to large language model word embeddings. We use these linguistic features in encoding models that relate stimulus properties to neural activity. Finally, we provide detailed tutorials for preprocessing raw data, extracting stimulus features, and running encoding analyses that can serve as a pedagogical resource or a springboard for new research.

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Fast-timescale hippocampal processes bridge between slowly unfurling neocortical states during memory search

Prior behavioral work showed that event structure supports our ability to search through continuous naturalistic memories. We hypothesized that, neurally, this memory search process involves a division of labor between slowly unfurling neocortical states representing event knowledge and fast hippocampal-neocortical communication supporting retrieval of new information at transitions between events. In a sample of ten intracranial EEG patients we tracked slow neural state-patterns as patients viewed a movie and then searched their memories in a structured naturalistic interview. As patients answered questions ("after A, when does B happen next?"), state-patterns from movie-viewing were reinstated in neocortex. During the critical memory-search period, states unfurled in a forward direction, realizing a trajectory from A to B in memory. Punctate moments of state-transition during memory-search were marked by low-frequency power decreases in cortex and preceded by power decreases in hippocampus that correlated with reinstatement. Connectivity-analysis revealed information-flow from hippocampus to cortex underpinning state-transitions.

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Predictive acoustical processing in human cortical layers

In our dynamic environments, predictive processing is vital for auditory perception and its associated behaviors. Predictive coding formalizes inferential processes by implementing them as information exchange across cortical layers and areas. With laminar-specific blood oxygenation level dependent we measured responses to a cascading oddball paradigm, to ground predictive auditory processes on the mesoscopic human cortical architecture. We show that the violation of predictions are potentially hierarchically organized and associated with responses in superficial layers of the planum polare and middle layers of the lateral temporal cortex. Moreover, we relate the updating of the brains internal model to changes in deep layers. Using a modeling approach, we derive putative changes in neural dynamics while accounting for draining effects. Our results support the role of temporal cortical architecture in the implementation of predictive coding and highlight the ability of laminar fMRI to investigate mesoscopic processes in a large extent of temporal areas.

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"What" and "when" predictions jointly modulate speech processing

Adaptive behavior rests on forming predictions based on previous statistical regularities encountered in the environment. Such regularities pertain not only to the contents of the stimuli ("what") but also their timing ("when"), and both interactively modulate sensory processing. In speech streams, predictions can be formed at multiple hierarchical levels, both in terms of contents (e.g. single syllables vs. words) and timing (e.g., faster vs. slower time scales). Whether and how these hierarchies map onto each other in terms of integrating "what" and "when" predictions remains unknown. Under one hypothesis neural hierarchies may link "what" and "when" predictions within sensory processing areas: with lower cortical regions mediating interactions for smaller units e.g., syllables, and higher cortical areas mediating interactions for larger units e.g., words. Alternatively, interactions between "what" and "when" predictions might rest on a generic, sensory-independent mechanism, mediated by common attention-related (e.g., frontoparietal) networks. To address those questions, we manipulated "what" and "when" predictions at two levels - single syllables and disyllabic pseudowords - while recording neural activity using magnetoencephalography (MEG) in healthy volunteers (N=22). We studied how syllable and/or word deviants are modulated by "when" predictability, both analyzing event-related fields and using source reconstruction and dynamic causal modeling to explain the observed effects in terms of the underlying effective connectivity. "When" predictions modulated "what" mismatch responses in a specific way with regards to speech hierarchy, such that mismatch responses to deviant words (vs. syllables) were amplified by temporal predictions at a slower (vs. faster) time scale. However, these modulations were source-localized to a shared network of cortical regions, including frontal and parietal sources. Effective connectivity analysis showed that, while mismatch responses to violations of "what" predictions modulated connectivity between regions, the integration of "what" and "when" predictions selectively modulated connectivity within regions, consistent with gain effects. These results suggest that the brain integrates "what" and "when" predictions that are congruent with respect to their hierarchical level, but this integration is mediated by a shared and distributed cortical network. This contrasts with recent studies indicating separable networks for different levels of hierarchical speech processing.

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A shared linguistic space for transmitting our thoughts from brain to brain in natural conversations

Effective communication hinges on a mutual understanding of word meaning in different contexts. The embedding space learned by large language models can serve as an explicit model of the shared, context-rich meaning space humans use to communicate their thoughts. We recorded brain activity using electrocorticography during spontaneous, face-to-face conversations in five pairs of epilepsy patients. We demonstrate that the linguistic embedding space can capture the linguistic content of word-by-word neural alignment between speaker and listener. Linguistic content emerged in the speakers brain before word articulation, and the same linguistic content rapidly reemerged in the listeners brain after word articulation. These findings establish a computational framework to study how human brains transmit their thoughts to one another in real-world contexts.

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An adversarial collaboration to critically evaluate theories of consciousness

Different theories explain how subjective experience arises from brain activity1,2. These theories have independently accrued evidence, yet, confirmation bias and dependence on design choices hamper progress in the field3. Here, we present an open science adversarial collaboration which directly juxtaposes Integrated Information Theory (IIT)4,5 and Global Neuronal Workspace Theory (GNWT)6-10, employing a theory-neutral consortium approach11,12. We investigate neural correlates of the content and duration of visual experience. The theory proponents and the consortium developed and preregistered the experimental design, divergent predictions, expected outcomes, and their interpretation12. 256 human subjects viewed suprathreshold stimuli for variable durations while neural activity was measured with functional magnetic resonance imaging, magnetoencephalography, and electrocorticography. We find information about conscious content in visual, ventro-temporal and inferior frontal cortex, with sustained responses in occipital and lateral temporal cortex reflecting stimulus duration, and content-specific synchronization between frontal and early visual areas. These results confirm some predictions of IIT and GNWT, while substantially challenging both theories: for IIT, a lack of sustained synchronization within posterior cortex contradicts the claim that network connectivity specifies consciousness. GNWT is challenged by the general lack of ignition at stimulus offset and limited representation of certain conscious dimensions in prefrontal cortex. Beyond challenging the theories themselves, we present an alternative approach to advance cognitive neuroscience through a principled, theory-driven, collaborative effort. We highlight the challenges to change peoples mind 13 and the need for a quantitative framework integrating evidence for systematic theory testing and building.

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"What" and "When" Predictions Modulate Auditory Processing in a Contextually Specific Manner

Extracting regularities from ongoing stimulus streams to form predictions is crucial for adaptive behavior. Such regularities exist in terms of the content of the stimuli (i.e., "what" it is) and their timing (i.e., "when" it will occur), both of which are known to interactively modulate sensory processing. In real-world stimulus streams, regularities also occur contextually - e.g. predictions of individual notes vs. melodic contour in music. However, it is unknown whether the brain integrates predictions in a contextually congruent manner (e.g., if slower "when" predictions selectively interact with complex "what" predictions), and whether integrating predictions of simple vs. complex features rely on dissociable neural correlates. To address these questions, our study employed "what" and "when" violations at different levels - single tones (elements) vs. tone pairs (chunks) - within the same stimulus stream, while neural activity was recorded using electroencephalogram (EEG) in participants (N=20) performing a repetition detection task. Our results reveal that "what" and "when" predictions interactively modulated stimulus-evoked response amplitude in a contextually congruent manner, but that these modulations were shared between contexts in terms of the spatiotemporal distribution of EEG signals. Effective connectivity analysis using dynamic causal modeling showed that the integration of "what" and "when" prediction selectively increased connectivity at relatively late cortical processing stages, between the superior temporal gyrus and the fronto-parietal network. Taken together, these results suggest that the brain integrates different predictions with a high degree of contextual specificity, but in a shared and distributed cortical network. Significance statementPredictions of stimulus features, present in different statistically-regular contexts in the environment, are crucial to forming adaptive behavior. However, it is unknown if the brain integrates predictions selectively according to such contextual differences. By recording human electroencephalography during experimental manipulations of time-based and content-based predictions, we found that those predictions interactively modulated neural activity in a contextually congruent manner, such that local (vs. global) time-based predictions modulated content-based predictions of sequence elements (vs. chunks). These modulations were shared between contextual levels in terms of the spatiotemporal distribution of neural activity. This suggests that the brain integrates different predictions with a high degree of contextual specificity, but in a shared and distributed cortical network.

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Simultaneous Mnemonic and Predictive Representations in the Auditory Cortex

Recent studies have shown that stimulus history can be decoded via the use of broadband sensory impulses to reactivate mnemonic representations. It has also been shown that predictive mechanisms in the auditory system demonstrate similar tonotopic organization of neural activity as that elicited by the perceived stimuli. However, it remains unclear if the mnemonic and predictive information can be decoded from cortical activity simultaneously and from overlapping neural populations. Here, we recorded neural activity using electrocorticography (ECoG) in the auditory cortex of anesthetized rats while exposed to repeated stimulus sequences, where events within the sequence were occasionally replaced with a broadband noise burst or omitted entirely. We show that both stimulus history and predicted stimuli can be decoded from neural responses to broadband impulse at overlapping latencies but linked to largely independent neural populations. We also demonstrate that predictive representations are learned over the course of stimulation at two distinct time scales, reflected in two dissociable time windows of neural activity. These results establish a valuable tool for investigating the neural mechanisms of passive sequence learning, memory encoding, and prediction mechanisms within a single paradigm, and provide novel evidence for learning predictive representations even under anaesthesia.

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Tracking transitional probabilities and segmenting auditory sequences are dissociable processes in adults and neonates

Since speech is a continuous stream with no systematic boundaries between words, how do pre-verbal infants manage to discover words? A proposed solution is that they might use the transitional probability between adjacent syllables, which drops at word boundaries. Here, we tested the limits of this mechanism by increasing the size of the word-unit to 4 syllables, and its automaticity by testing asleep neonates. Using markers of statistical learning in neonates EEG, compared to adult behavioral performances in the same task, we confirmed that statistical learning is automatic enough to be efficient even in sleeping neonates. But we also revealed that: 1) Successfully tracking transition probabilities in a sequence is not sufficient to segment it 2) Prosodic cues, as subtle as subliminal pauses, enable to recover segmenting capacities 3) Adults and neonates capacities are remarkably similar despite the difference of maturation and expertise. Finally, we observed that learning increased the similarity of neural responses across infants, providing a new neural marker to monitor learning. Thus, from birth, infants are equipped with adult-like tools, allowing to extract small coherent word-like units within auditory streams, based on the combination of statistical analyses and prosodic cues.

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The Consciousness Theories Studies (ConTraSt) database: analyzing and comparing empirical studies of consciousness theories

Understanding how consciousness arises from neural activity remains one of the biggest challenges for neuroscience. Numerous theories have been proposed in recent years, each gaining independent empirical support. Currently, there is no comprehensive, quantitative and theory-neutral overview of the field that enables an evaluation of how theoretical frameworks interact with empirical research. We provide a birds eye view on studies that interpreted their findings in light of at least one of four leading neuroscientific theories of consciousness (N=412 experiments), asking how methodological choices of the researchers might affect the final conclusions. We found that supporting a specific theory can be predicted solely from methodological choices, irrespective of findings. Furthermore, most studies interpret their findings post-hoc, rather than a-priori testing critical predictions of the theories. Our results highlight challenges for the field and provide researchers with a unique, open-access website to further analyze trends in the neuroscience of consciousness.

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What canonical online and offline measures of statistical learning can and cannot tell us

Statistical learning (SL) allows individuals to rapidly detect regularities in the sensory environment. We replicated previous findings showing that adult participants become sensitive to the implicit structure in a continuous speech stream of repeating tri-syllabic pseudowords within minutes, as measured by standard tests in the SL literature: a target detection task and a 2AFC word recognition task. Consistent with previous findings, we found only a weak correlation between these two measures of learning, leading us to question whether there is overlap between the information captured by these two tasks. Representational similarity analysis on reaction times measured during the target detection task revealed that reaction time data reflect sensitivity to transitional probability, triplet position, word grouping, and duplet pairings of syllables. However, individual performance on the word recognition task was not predicted by similarity measures derived for any of these four features. We conclude that online detection tasks provide richer and multi-faceted information about the SL process, as compared with 2AFC recognition tasks, and may be preferable for gaining insight into the dynamic aspects of SL.

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Spontaneous neural oscillations influence behavior and sensory representations by suppressing neuronal excitability

The ability to process and respond to external input is critical for adaptive behavior. Why, then, do neural and behavioral responses vary across repeated presentations of the same sensory input? Spontaneous fluctuations of neuronal excitability are currently hypothesized to underlie the trial-by-trial variability in sensory processing. To test this, we capitalized on invasive electrophysiology in neurosurgical patients performing an auditory discrimination task with visual cues: specifically, we examined the interaction between prestimulus alpha oscillations, excitability, task performance, and decoded neural stimulus representations. We found that strong prestimulus oscillations in the alpha+ band (i.e., alpha and neighboring frequencies), rather than the aperiodic signal, correlated with a low excitability state, indexed by reduced broadband high-frequency activity. This state was related to slower reaction times and reduced neural stimulus encoding strength. We propose that the alpha+ rhythm modulates excitability, thereby resulting in variability in behavior and sensory representations despite identical input.

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The influence of auditory attention on rhythmic speech entrainment

Language comprehension relies on integrating words into progressively more complex structures, like phrases and sentences. This hierarchical structure building is reflected in rhythmic neural activity across multiple timescales in E/MEG (Ding et al., 2016, 2017). How does selective attention across levels of the hierarchy influence the expression of these rhythms? We investigated these questions in an EEG study of 72 healthy human volunteers listening to streams of monosyllabic isochronous English words that were either unrelated (scrambled condition) or composed of four-word-sequences building meaningful sentences (sentential condition). Importantly, there were no physical cues between four-word-sentences but boundaries were marked by syntactic structure and thematic role assignment. Participants were divided into three attention groups: from passive listening (passive group) to attending to individual words (word group) or sentences (sentence group). The passive and word group were naive to the sentential structure of the stimulus material, while the sentence group were not. We found significant entrainment at word- and sentence rate across all three groups, with sentence entrainment linked to left middle temporal gyrus and right superior temporal gyrus. Goal-directed attention to words did not enhance word-rate-entrainment suggesting that word entrainment relies on largely automatic mechanisms. Importantly, goal-directed attention to sentences relative to words significantly increased sentence-rate-entrainment over left inferior frontal gyrus. This attentional modulation of rhythmic EEG activity at the sentential level highlights the role of attention in integrating individual words into complex linguistic structures. SIGNIFICANCE STATEMENTNeural activity is known to entrain to physical characteristics of auditory stimuli. However, entrainment also occurs with structures lacking physical cues but rather require comprehension of the stimulus meaning - for example, entrainment to sentences in speech even without acoustic gaps separating these higher linguistic structures. We investigated how goal-directed attention to low-level (words) and high-level (sentences) linguistic structures influences entrainment strength. Whilst sentence entrainment occurred independently of selective attention, it increased with goal-directed attention towards sentences. Conversely, no such attentional effect was found for word entrainment. While goal-directed attention towards sentences strengthens entrainment, it is no prerequisite for it to occur, suggesting that low attentional effort is required for sentence comprehension, potentially reflecting the importance of speech in humans.

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