Biology subjects

Carreiras, M.

Publications and source records attributed to Carreiras, M..

At least 19 recordsLinked to original sources

bioRxiv2026

Low-frequency neural responses synchronize to distinct structural rather than lexical features during sentence comprehension

Language comprehension involves the integration of single words (lexical units) into phrases and sentences (multi-word structures). Previous frequency-tagging studies have found that low-frequency neural responses synchronize to the frequency of multi-word structures. However, it is currently unclear how exactly structural and lexical processes jointly impact these synchronization findings. The present magnetoencephalography experiment implemented the frequency-tagging paradigm in the visual modality with written words to investigate neural synchronization to multi-word sentences varying in internal structure (reversed word orders between verb-initial Spanish and verb-final Basque sentences) and in lexical content (real words and pseudo words). We find converging evidence that neural responses largely synchronize to structural rather than lexical features. This was observed as robust phase synchronization strength to the frequency of sentences containing reversed structures, with certain lexical modulations depending on language-specific structural features. Crucially, we also found shifted phase angle dynamics between the reversed structures of Spanish and Basque sentences independently of word-level lexical characteristics. Together, these findings suggest that neural synchronization to multi-word structures is largely driven by distinct structural features operating via two segregated neural dimensions: frequency coding for the coarser aspects (i.e., timescale/duration) and phase representing the finer-grained aspects (i.e., internal structure) of multi-word structures. Our findings thus advance key insights into the core components of the neural mechanisms supporting language comprehension. HighlightsO_LINeural synchronization to sentences is driven by structural (not lexical) features. C_LIO_LIRobust sentence-frequency synchronization across languages varying in structure. C_LIO_LIPhase angle is selectively sensitive to cross-linguistic structural differences. C_LIO_LILexical modulations depend on language-specific structure. C_LIO_LIStructure synchronization segregates into two dimensions: frequency and phase. C_LI

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bioRxiv2026

Young children rely on visual information to process degraded speech: Evidence from behavioural and neuroimaging measures

Young children acquire language in environments where speech is often acoustically degraded, yet little is known about how developing brains adapt to reduced speech intelligibility. Using a combination of eye-tracking and functional near-infrared spectroscopy (fNIRS), we investigated young childrens attentional allocation to a speaking face at varying levels of speech intelligibility and the brain activity supporting this behaviour during development. Infants (8-10 months) and toddlers (27-30 months) viewed videos of a speaker in three conditions: producing clear speech, spectrally degraded (vocoded) speech, and silent (audio muted) speech. Visual attention to the speakers mouth increased when speech was degraded relative to clear speech in both age groups, indicating an early-emerging compensatory strategy. However, this shared behavioural response was supported by brain activity that differed by age. Degraded speech elicited greater recruitment of prefrontal regions associated with effortful listening, particularly in infants, whereas toddlers showed stronger engagement of posterior temporal regions implicated in audiovisual integration. In response to silent speech, there was no evidence for increased visual attention to the mouth compared to the clear speech condition, but there was reduced temporal activation and increased prefrontal brain responses, especially in infants. Together, these findings suggest that experience with audiovisual correspondences and linguistic maturity contribute to a more efficient processing of speech, particularly relevant when speech is degraded. By combining behavioural and neuroimaging measures, this study advances mechanistic accounts of audiovisual speech processing and provides insights relevant to populations experiencing spectrally degraded input, such as children using cochlear implants.

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bioRxiv2026

Resting-state Compensatory Remapping in Patients with Brain Tumour Before and After Surgery

Brain tumours invade neural tissue, disrupting the functional organisation of neural networks. This disruption can trigger compensatory neuroplastic mechanisms that help preserve cognitive function despite pathological burden. Resting-state functional connectivity (rs-FC) provides a valuable approach for examining these alterations, yet the scope and trajectory of network-level changes remain unclear. In this study, we used rs-FC to characterise compensatory responses induced by left-hemisphere brain tumours affecting the integrity of the language network. The comparison of connectivity patterns at diagnosis and after surgery allowed us to track the temporal progression of these patterns. Relative to healthy participants, patients showed significant deviations in pre-surgical rs-FC, with widespread changes involving the dorsal attention, sensorimotor, frontoparietal, default mode, and cerebellar networks. These differences persisted post-surgically, with no significant longitudinal modifications. Network-level alterations also influenced topological properties: patients showed increased segregation while preserving global integration relative to controls. These properties changed significantly from pre- to postoperative period, with a postoperative increase in segregation and integration towards a pattern more closely resembling that of healthy controls. Furthermore, preoperative network segregation was identified as a predictor of postoperative cognitive recovery. These findings suggest that brain tumours induce network remapping, affecting functional connectivity far beyond the regions strictly related to the affected region. The observed changes in network topological organisation likely reflect the adaptive role of domain-general networks in facilitating homeostatic processes. Surgical strategies aimed at preserving these changes seem to promote sustained cognitive function, regardless of malignancy grade or lesion extent. Key pointsO_LIPresurgical patients show widespread inter- and intrahemispheric rs-FC alterations, reflecting tumour-driven network remapping. C_LIO_LINetwork integration and segregation show longitudinal recovery, approaching neurotypical topology four months post-surgery. C_LIO_LIPreoperative network segregation predicts postoperative cognitive recovery. C_LI Importance of the StudyBrain tumours disrupt large-scale brain networks, yet the extent, temporal dynamics, and cognitive relevance of these alterations remain poorly understood. Using resting-state functional connectivity, this study provides a longitudinal, network-level characterisation of tumour-induced neuroplasticity in patients with left-hemisphere lesions affecting language-related regions. Compared with prior work focused on local effects or single networks, our results demonstrate widespread inter- and intrahemispheric connectivity changes involving multiple domain-general resting-state networks, highlighting extensive network remapping beyond lesion boundaries. Importantly, we show that postoperative recovery is accompanied by a rebalancing of network integration and segregation toward a neurotypical configuration. Critically, preoperative network segregation emerges as a predictor of postoperative cognitive recovery, identifying a potential biomarker of resilience. These findings have direct translational relevance, suggesting that preserving and supporting large-scale network organisation during surgical planning may promote cognitive outcomes independently of tumour grade or lesion extent.

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bioRxiv2026

Alternative strategies of orthographic processing: the case of skilled deaf readers

The cognitive factors that enable us to be proficient readers can greatly vary across individuals. The case of skilled deaf readers is emblematic as it shows that high reading performances can be achieved even when lifelong acoustic experience is absent or minimal. Here we present a set of experiments investigating how alternative strategies of orthographic processing can lead to high levels of reading proficiency. Four EEG studies compared behavioral and brain correlates of orthographic processing in skilled deaf readers and matched hearing controls. Using single word recognition and priming paradigms, we investigated two pillars of orthographic processing: letter identity and letter position. Our findings show that, although both groups had similarly accurate reading performance, skilled deaf readers were faster, and they consistently differ from hearing controls in the way they process letter identity. This group difference was observed in both lexical and sublexical tasks and was specifically related to the identity of orthographic representations, regardless of the visual form of the written stimuli (such as character visual similarity and letter case). These findings uncover alternative strategies that make possible high reading performance, even in the absence of acoustic experience. Public Significance StatementThis research identifies alternative orthographic strategies that improve single-word reading efficiency and can potentially serve as effective compensatory tools when phonological processing is impaired.

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bioRxiv2025

Contralesional grey matter volume as an index of macrostructural plasticity in patients with brain tumors

This research challenges the traditional localizationist view that brain tumors affect only regions directly associated with the lesion, by examining whether they also induce macrostructural alterations in the contralesional hemisphere. We applied Voxel-Based Morphometry, linear regression, and Principal Component Analysis (PCA) to a cohort of 107 adults, including patients with gliomas in the language-dominant left hemisphere and healthy participants. Unlike previous studies, a subset of the clinical population was followed longitudinally for up to four months after oncological treatment, allowing us to describe the temporal progression of structural grey matter changes. Interestingly, a principal component model based on anomaly detection enabled robust differentiation between patients and controls. Patients exhibited significantly greater grey matter volume in the contralesional hemisphere compared to healthy participants, and these structural differences evolved over time, improving the models AUC-ROC metrics. Although exploratory, a correlation analysis revealed that these structural changes were negatively associated with postsurgical cognitive performance. Together with the PCA findings, these results suggest that brain tumors induce extensive and dynamic adaptive mechanisms in the contralateral, unaffected hemisphere, likely reflecting altered patterns of structural covariance rather than simple regional volume increases. Understanding whether these changes could represent potential predictors of postoperative cognitive recovery is crucial for developing comprehensive clinical strategies. Key PointsO_LILeft-hemisphere tumors induce contralesional grey matter increases pre- and post-sugery C_LIO_LIPCA detects altered structural covariance and distinguishes patients from healthy participants C_LIO_LIContralesional grey matter volume changes correlate with postoperative cognitive performance C_LI Importance of the StudyThis study challenges the traditional view that brain tumors cause only localized effects by demonstrating widespread macrostructural alterations in the contralesional hemisphere. We reveal increased grey matter volume and altered patterns of structural covariance outside the tumor region. Longitudinal follow-up after surgery shows these changes are dynamic and evolve over time. Further, we identify moderate associations between contralesional grey matter alterations and cognitive performance, suggesting a link between large-scale neuroplastic responses and functional outcomes. These findings offer new insights into tumor-related neuroplasticity and position structural covariance as a promising marker for tracking brain-wide adaptation in this population. The study has translational relevance for developing predictive tools to monitor recovery and guide personalized rehabilitation.

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bioRxiv2025

NEUROLINGUA: A Neuroimaging Database Tailored to Unravel the Complexity of Multilingual Comprehension

The neural mechanisms underlying language processing involve a well-defined brain network, including mainly left perisylvian areas. Yet, the extent of individual variability remains largely unexplored, particularly in bilingual and multilingual contexts. Differences in linguistic profiles (e.g., age of acquisition, exposure, proficiency) provide an opportunity to assess how network topology is shaped by sociolinguistic factors. To address this, we developed NEUROLINGUA, a comprehensive database of functional and structural MRI data, enriched with sociodemographic, sociolinguistic, and behavioral information. It includes 725 healthy individuals aged 18-82 immersed in a Basque-Spanish multilingual environment, ranging from near-monolinguals to highly proficient multilinguals. Participants completed a functional MRI language localizer with both auditory and visual comprehension tasks, enabling cross-modal comparisons, as well as sentences involving arithmetic problem-solving. Exploratory analyses confirmed associations between structural MRI, sociodemographic, and cognitive measures. We demonstrate that NEUROLINGUAs functional MRI data localize the language comprehension network and thus capture linguistic profile effects. This integrative dataset offers an unparalleled resource to investigate factors influencing language network adaptability and variability in diverse sociolinguistic contexts.

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bioRxiv2024

Electrophysiological correlates of cognitive reserve in healthy older adults at different cognitive states

High cognitive reserve (CR) is associated with a set of lifestyle factors such as high education level and occupational attainment. Studies suggest that CR is linked to enhanced executive control functions. Nevertheless, it is unclear what are the functional neural activity patterns linked to high CR and to what extent they are steady across different cognitive states. The aim of the present study was to investigate electrophysiological differences between low CR (LCR) and high CR (HCR) at rest and during the performance of two cognitive control tasks with different difficulty level. Thus, 67 older adults performed an experimental session consisting of two spatial stimulus-response compatibility (SRC) tasks (i.e., the Simon task, and the more demanding spatial Stroop task), and a resting state during an electroencephalogram (EEG) recording. Cognitive control was better in HCR than LCR group, as revealed by higher accuracy in HCR compared to LCR group in incongruent conditions of the tasks. Moreover, event-related brain potentials (ERP) showed earlier latencies (P300) in HCR than LCR and time/frequency analysis showed higher alpha activity in HCR than LCR during the performance of the tasks, while no differences were observed at rest within any of the analyzed frequency bands. Higher P300 frontalization was observed in LCR than HCR in the more difficult task but it was not related to frontalization within a specific frequency band. Overall, results showed that electrophysiological differences between high and low CR depend on the cognitive state, with greater CR-related differences emerging at increased task demands.

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bioRxiv2024

Functional brain adaptations during speech processing in 4-month-old bilingual infants

Language learning is influenced by both neural development and environmental experiences. This work investigates the influence of early bilingual experience on the neural mechanisms underlying speech processing in 4-month-old infants. We study how an early environmental factor such as bilingualism interacts with neural development by comparing monolingual and bilingual infants brain responses to speech. We used functional near-infrared spectroscopy (fNIRS) to measure 4-month-old Spanish-Basque bilingual and Spanish monolingual infants brain responses while they listened to forward (FW) and backward (BW) speech stimuli in Spanish. We reveal distinct neural signatures associated with bilingual adaptations, including increased engagement of bilateral inferior frontal and temporal regions during speech processing in bilingual infants, as opposed to left hemispheric specialization observed in monolingual infants. This study provides compelling evidence of bilingualism-induced brain adaptations during speech processing in infants as young as four months. These findings emphasize the role of early language experience in shaping neural plasticity during infancy suggesting that bilingual exposure at this young age profoundly influences the neural mechanisms underlying speech processing. Research HighlightsO_LIThis research sheds light on the intricate relationship between language learning, neural maturation, and environmental factors in early infancy. C_LIO_LIBilingual infants exhibit unique functional brain adaptations during speech processing as early as 4 months of age. C_LIO_LIThis study underscores the critical role of language experience in shaping the developing brain. C_LIO_LIThese findings have implications for our understanding of language acquisition and neuroplasticity in infants growing up in bilingual environments. C_LI

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bioRxiv2023

Early delta cortical tracking to non-verbal auditory stimuli predicts reading skills: A one-year longitudinal study

The precision of cortical tracking of auditory rhythmic stimuli in low frequency ranges coding for prosodic (delta) and syllabic (theta) amplitude modulations in the speech signal has been proposed to contribute to the development of phonological processing and reading acquisition. The present study investigates the role of low frequency cortical tracking of non-verbal auditory stimuli in reading acquisition through a longitudinal design from before the onset of formal reading instruction until one year afterwards. At time one, 40 prereading children performed a passive listening task of amplitude modulated white noise presented in the delta (2 Hz) and theta (4 Hz) frequency bands, while their neural activity was recorded via EEG. At time two, at the end of first grade, childrens reading skills were assessed. Results show significant cortical tracking in prereading children at both frequencies, with larger responses for theta than for delta rate non-verbal auditory stimuli. Importantly, only pre-reding cortical tracking measures for delta rate stimuli predicted reading acquisition one year later. These findings underscore the role of early neural synchronization to delta rate rhythmic auditory stimuli to reading acquisition and support the potential important role of early prosodic processing for the development of future reading skills.

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bioRxiv2023

Early language dissociation in the bilingual mind: Sensor-level MEG evidence

Is language selection in balanced bilinguals decodable from neural activity? Previous research employing various neuroimaging methods has not yielded a conclusive answer to this issue. However, direct brain stimulation studies in bilinguals have detected different brain regions related to language production in separate languages. In the present MEG study, we addressed this question in a group of proficient Spanish-Basque bilinguals (N=45), who performed two tasks (picture naming and word reading). They were asked to name the line drawing or read the word out loud, either in Basque or Spanish, if the ink turned from black to green after one second (randomly, in 10% of trials). Sensor-level evoked activity was similar and could not be differentiated for the two languages in either task. Crucially however, decoding analyses classified the language used in both tasks, starting [~]100 ms after stimulus onset. Searchlight analyses revealed that activity detected in the right occipital-temporal sensors contributed the most to language decoding in the picture naming task, while the left occipital-temporal sensors contributed the most to decoding in word reading. The cross-task decoding analysis highlighted robust generalization effects from the picture naming to the word reading task in a later time interval. The present findings bridge the gap between non-invasive and invasive experimental evidence on bilingual lexical processing and provide novel evidence about the role of the two hemispheres in activating each language for picture naming and word reading.

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bioRxiv2023

Cognitive reserve counteracts typical neural activity changes related to ageing

Studies have shown that older adults with high Cognitive Reserve (HCR) exhibit better executive functioning than their low CR (LCR) counterparts. However, the neural processes linked to those differences are unclear. This study investigates (1) the neural processes underlying enhanced executive functions in older adults with HCR and (2) how executive control differences are modulated by task difficulty. We recruited 74 participants, who performed two executive control tasks with different difficulty levels while recording the electroencephalogram. The accuracy on both tasks requiring inhibition of irrelevant information was better in the HCR than the LCR group. Also, in the more demanding task, event-related potentials (ERP) latencies related to inhibition and working memory update were faster in HCR than LCR. Moreover, the HCR, but not the LCR, showed larger P300 amplitude in parietal than frontal regions and in the left than right hemisphere, suggesting a posterior to anterior shift of activity and loss of inter-hemispheric asymmetries in LCR participants. These results suggest that high CR counteracts neural activity changes related to ageing. Thus, high levels of CR can be related to maintenance of neural activity patterns typically observed in young adults rather than to deployment of neural compensatory mechanisms.

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bioRxiv2022

Decoding Numeracy and Literacy in the Human Brain: Insights from MEG and MVPA

Numbers and letters are the fundamental building blocks of our everyday social interactions. Previous studies have focused on determining the cortical networks shaped by numeracy and literacy in the human brain, partially supporting the hypothesis of distinct neural circuits involved the processing of the two categories. In this study, we aim to investigate the temporal dynamics for number and letter processing. We present MEG data from two experiments (N=20 each). In the first experiment, single numbers, letter, and their respective false fonts (false numbers and false letters) were presented, whereas, in the second experiment, numbers, letters, and their respective false fonts were presented as a string of characters. We used multivariate pattern analysis techniques (time-resolved decoding and temporal generalization) testing the strong hypothesis that the neural mechanisms supporting letter and number processing can be logistically classified as categorically separate. Our results show a very early dissociation ([~]100 ms) between numbers, and letters when compared to false fonts. Number processing can be dissociated with similar accuracy when presented as isolated items or strings of characters, while letter processing shows dissociable classification accuracy for single items compared to strings. These findings reinforce the evidence indicating that early visual processing can be differently shaped by the experience with numbers and letters; this dissociation is stronger for strings compared to single items, thus showing that combinatorial mechanisms for numbers and letters could be categorically distinguished and influence early brain activity.

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bioRxiv2022

Brain structure, phenotypic and genetic correlates of reading abilities

Reading is an evolutionary new development that recruits and tunes brain circuitry connecting visual- and language-processing regions. We investigated the structural correlates of reading and whether genetics influence brain-reading associations. First, we identified left hemisphere cortical surface area (CSA) and cortical thickness (CT) correlates of reading in the large ABCD dataset (N=9,013) of 9-to-10-year-olds. Next, the heritability of cognitive and brain measures of interest was examined through complementary approaches. Last, shared genetic effects between reading, reading-related cognitive traits and reading-associated brain measures were examined by computing genetic correlations and polygenic score analyses, and through mediation analyses. Our results support that morphometric brain measures are related to reading abilities, and that the total left CSA in general, and left superior temporal gyrus CSA in particular, contribute to reading partially through genetic factors.

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bioRxiv2021

Reading-related brain changes in audiovisual processing: cross-sectional and longitudinal MEG evidence

The ability to establish associations between visual objects and speech sounds is essential for human reading. Understanding the neural adjustments required for acquisition of these arbitrary audiovisual associations can shed light on fundamental reading mechanisms and help reveal how literacy builds on pre-existing brain circuits. To address these questions, the present longitudinal and cross-sectional MEG studies characterize the temporal and spatial neural correlates of audiovisual syllable congruency in children (4-9 years old, 22 males and 20 females) learning to read. Both studies showed that during the first years of reading instruction children gradually set up audiovisual correspondences between letters and speech sounds, which can be detected within the first 400 ms of a bimodal presentation and recruit the superior portions of the left temporal cortex. These findings suggest that children progressively change the way they treat audiovisual syllables as a function of their reading experience. This reading-specific brain plasticity implies (partial) recruitment of pre-existing brain circuits for audiovisual analysis. Significance StatementLinking visual and auditory linguistic representations is the basis for the development of efficient reading, while dysfunctional audiovisual letter processing predicts future reading disorders. Our developmental MEG project included a longitudinal and a cross-sectional study; both studies showed that childrens audiovisual brain circuits progressively change as a function of reading experience. They also revealed an exceptional degree of neuroplasticity in audiovisual neural networks, showing that as children develop literacy, the brain progressively adapts so as to better detect new correspondences between letters and speech sounds.

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bioRxiv2020

Monolingual and bilingual infants rely on the same brain networks: Evidence from resting-state functional connectivity

This study examines whether bilingual exposure has a profound effect on the functional organization of the developing human brain during infancy. Recent behavioural research attests that monolingual vs. bilingual experience affects cognitive and linguistic processes already during the first months of life. However, to what extent the intrinsic organization of the infant human brain adapts to monolingual vs. bilingual environments is unclear. We measured spontaneous hemodynamic brain activity using functional near-infrared spectroscopy (fNIRS) in a large cohort (N=99) of 4-month-old monolingual and bilingual infants. We implemented well-established analysis approaches of functional brain imaging that enabled us to reveal the functional organization of the infant brain in large-scale cortical networks, and to perform group-level comparisons (i.e., monolingual vs. bilingual groups) in a reliable manner. Our results revealed no differences between the intrinsic functional organization of the developing monolingual and bilingual infant brain at 4 months of age.

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bioRxiv2020

MULTIMAP: Multilingual visual naming test for the mapping of eloquent areas during awake surgeries.

Picture naming tasks are currently the gold standard for identifying and preserving language-related areas during awake brain surgery. With multilingual populations increasing worldwide, patients frequently need to be tested in more than one language. There is still no reliable testing instrument, as the available batteries have been developed for specific languages. Heterogeneity in the selection criteria for stimuli leads to differences, for example, in the size, color, image quality, and even names associated with pictures, making direct cross-linguistic comparisons difficult. Here we present MULTIMAP, a new multilingual picture naming test for mapping eloquent areas during awake brain surgery. Recognizing that the distinction between nouns and verbs is necessary for detailed and precise language mapping, MULTIMAP consists of a database of 218 standardized color pictures representing both objects and actions. These images have been tested for name agreement with speakers of Spanish, Basque, Catalan, Italian, French, English, Mandarin Chinese, and Arabic, and have been controlled for relevant linguistic features in cross-language combinations. The MULTIMAP test for objects and verbs represents an alternative to the DO 80 monolingual pictorial set currently used in language mapping, providing an open-source, standardized set of up-to-date pictures, where relevant linguistic variables across several languages have been taken into account in picture creation and selection.

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bioRxiv2020

Linguistic input drives brain network configuration during language comprehension.

Assessing the synchrony and interplay between distributed neural regions is critical to understanding how language is processed. Here, we investigated possible neuro-functional links between form and meaning during sentence comprehension combining a classical whole-brain approach, which characterizes patterns of brain activation resulting from our experimental manipulation, and a novel multivariate network-based approach, which uses graph-theory measures to unravel the architectural configuration of the language system. Capitalizing on the Spanish gender agreement system, we experimentally manipulated formal and conceptual factors: whether the noun-adjective grammatical gender relationship was congruent or not and whether the noun gender type was semantically informative or strictly formal. Left inferior and middle frontal gyri, as well as left MTG/STG emerged as critical areas for the computation of grammatical relations. We demonstrate how the interface between formal and conceptual features depends on the synergic articulation of brain areas divided in three subnetworks that extend beyond the classical left-lateralized perisylvian language circuit. Critically, we isolated a subregion of the left angular gyrus showing a significant interaction between gender congruency and gender type. The functional interplay between the angular gyrus and left perisylvian language-specific circuit proves crucial for constructing coherent and meaningful messages. Importantly, using graph theory we show that this complex system is functionally malleable: the role each node plays within the network changes depending on the available linguistic cues. Significance StatementNeural networks can be described as graphs comprising distributed and interconnected nodes. These nodes share functional properties but also differ in terms of functional specialization and the number of interconnections mediating the efficient transfer of information. Previous work has shown functional connectivity differences based on graph-theory properties between typical and atypical samples. However, here we have used concepts from graph theory to characterize connectivity during language processing using task-related fMRI. This approach allowed us to demonstrate how linguistic input drives brain network configuration during language comprehension. This is the first evidence of functional flexibility in language networks: the communicative capacity of each hub changes depending on whether the linguistic input grants access to meaning or is purely formal.

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bioRxiv2019

Neocortical activity tracks syllable and phrasal structure of self-produced speech

To gain novel insights into how the human brain processes self-produced auditory information during reading aloud, we investigated the coupling between neuromagnetic activity and the temporal envelope of the heard speech sounds (i.e., speech brain tracking) in a group of adults who 1) read a text aloud, 2) listened to a recording of their own speech (i.e., playback), and 3) listened to another speech recording. Coherence analyses revealed that, during reading aloud, the readers brain tracked the slow temporal fluctuations of the speech output. Specifically, auditory cortices tracked phrasal structure (<1 Hz) but to a lesser extent than during the two speech listening conditions. Also, the tracking of syllable structure (4-8 Hz) occurred at parietal opercula during reading aloud and at auditory cortices during listening. Directionality analyses based on renormalized partial directed coherence revealed that speech brain tracking at <1 Hz and 4-8 Hz is dominated by speech-to-brain directional coupling during both reading aloud and listening, meaning that speech brain tracking mainly entails auditory feedback processing. Nevertheless, brain-to-speech directional coupling at 4- 8 Hz was enhanced during reading aloud compared with listening, likely reflecting speech monitoring before production. Altogether, these data bring novel insights into how auditory verbal information is tracked by the human brain during perception and self-generation of connected speech.\n\nHighlightsO_LIThe brain tracks phrasal and syllabic rhythmicity of self-produced (read) speech.\nC_LIO_LITracking of phrasal structures is attenuated during reading compared with listening.\nC_LIO_LISpeech rhythmicity mainly drives brain activity during reading and listening.\nC_LIO_LIBrain activity drives syllabic rhythmicity more during reading than listening.\nC_LI

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