Language experience shapes predictive coding of rhythmic sound sequences.

Perceptual systems heavily rely on prior knowledge and predictions to make sense of the environment. Predictions can originate from multiple sources of information, including contextual short-term priors, based on isolated temporal situations, and context-independent long-term priors, arising from extended exposure to statistical regularities. While the effects of short-term predictions on auditory perception have been well-documented, how long-term predictions shape early auditory processing is poorly understood. To address this, we recorded magnetoencephalography data from native speakers of two languages with different word orders (Spanish: functor-initial vs Basque: functor-final) listening to simple sequences of binary sounds alternating in duration with occasional omissions. We hypothesized that, together with contextual transition probabilities, the auditory system uses the characteristic prosodic cues (duration) associated with the native language's word order as an internal model to generate long-term predictions about incoming non-linguistic sounds. Consistent with our hypothesis, we found that the amplitude of the mismatch negativity elicited by sound omissions varied orthogonally depending on the speaker's linguistic background and was most pronounced in the left auditory cortex. Importantly, listening to binary sounds alternating in pitch instead of duration did not yield group differences, confirming that the above results were driven by the hypothesized long-term 'duration' prior. These findings show that experience with a given language can shape a fundamental aspect of human perception - the neural processing of rhythmic sounds - and provides direct evidence for a long-term predictive coding system in the auditory cortex that uses auditory schemes learned over a lifetime to process incoming sound sequences.

Early language experience modulates the tradeoff between acoustic-temporal and lexico-semantic cortical tracking of speech.

Cortical tracking of speech is relevant for the development of speech perception skills. However, no study to date has explored whether and how cortical tracking of speech is shaped by accumulated language experience, the central question of this study. In 35 bilingual children (6-year-old) with considerably bigger experience in one language, we collected electroencephalography data while they listened to continuous speech in their two languages. Cortical tracking of speech was assessed at acoustic-temporal and lexico-semantic levels. Children showed more robust acoustic-temporal tracking in the least experienced language, and more sensitive cortical tracking of semantic information in the most experienced language. Additionally, and only for the most experienced language, acoustic-temporal tracking was specifically linked to phonological abilities, and lexico-semantic tracking to vocabulary knowledge. Our results indicate that accumulated linguistic experience is a relevant maturational factor for the cortical tracking of speech at different levels during early language acquisition.

On the speech envelope in the cortical tracking of speech.

The synchronization between the speech envelope and neural activity in auditory regions, referred to as cortical tracking of speech (CTS), plays a key role in speech processing. The method selected for extracting the envelope is a crucial step in CTS measurement, and the absence of a consensus on best practices among the various methods can influence analysis outcomes and interpretation. Here, we systematically compare five standard envelope extraction methods the absolute value of Hilbert transform (absHilbert), gammatone filterbanks, heuristic approach, Bark scale, and vocalic energy), analyzing their impact on the CTS. We present performance metrics for each method based on the recording of brain activity from participants listening to speech in clear and noisy conditions, utilizing intracranial EEG, MEG and EEG data. As expected, we observed significant CTS in temporal brain regions below 10 Hz across all datasets, regardless of the extraction methods. In general, the gammatone filterbanks approach consistently demonstrated superior performance compared to other methods. Results from our study can guide scientists in the field to make informed decisions about the optimal analysis to extract the CTS, contributing to advancing the understanding of the neuronal mechanisms implicated in CTS.

Early language dissociation in bilingual minds: magnetoencephalography evidence through a machine learning approach.

Does neural activity reveal how balanced bilinguals choose languages? Despite using diverse neuroimaging techniques, prior studies haven't provided a definitive solution to this problem. Nonetheless, studies involving direct brain stimulation in bilinguals have identified distinct brain regions associated with language production in different languages. In this magnetoencephalography study with 45 proficient Spanish-Basque bilinguals, we investigated language selection during covert picture naming and word reading tasks. Participants were prompted to name line drawings or read words if the color of the stimulus changed to green, in 10% of trials. The task was performed either in Spanish or Basque. Despite similar sensor-level evoked activity for both languages in both tasks, decoding analyses revealed language-specific classification ~100 ms post-stimulus onset. During picture naming, right occipital-temporal sensors predominantly contributed to language decoding, while left occipital-temporal sensors were crucial for decoding during word reading. Cross-task decoding analysis unveiled robust generalization effects from picture naming to word reading. Our methodology involved a fine-grained examination of neural responses using magnetoencephalography, offering insights into the dynamics of language processing in bilinguals. This study refines our understanding of the neural underpinnings of language selection and bridges the gap between non-invasive and invasive experimental evidence in bilingual language production.

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 pathways shaped by numeracy and literacy in the human brain, partially supporting the hypothesis of distinct perceptual neural circuits involved in the visual processing of the two categories. In this study, we aim to investigate the temporal dynamics for number and letter processing. We present magnetoencephalography (MEG) data from two experiments (N = 25 each). In the first experiment, single numbers, letters, 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 correlates 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 visual processing.

Increased top-down semantic processing in natural speech linked to better reading in dyslexia.

Early research proposed that individuals with developmental dyslexia use contextual information to facilitate lexical access and compensate for phonological deficits. Yet at present there is no corroborating neuro-cognitive evidence. We explored this with a novel combination of magnetoencephalography (MEG), neural encoding and grey matter volume analyses. We analysed MEG data from 41 adult native Spanish speakers (14 with dyslexic symptoms) who passively listened to naturalistic sentences. We used multivariate Temporal Response Function analysis to capture online cortical tracking of both auditory (speech envelope) and contextual information. To compute contextual information tracking we used word-level Semantic Surprisal derived using a Transformer Neural Network language model. We related online information tracking to participants' reading scores and grey matter volumes within the reading-linked cortical network. We found that right hemisphere envelope tracking was related to better phonological decoding (pseudoword reading) for both groups, with dyslexic readers performing worse overall at this task. Consistently, grey matter volume in the superior temporal and bilateral inferior frontal areas increased with better envelope tracking abilities. Critically, for dyslexic readers only, stronger Semantic Surprisal tracking in the right hemisphere was related to better word reading. These findings further support the notion of a speech envelope tracking deficit in dyslexia and provide novel evidence for top-down semantic compensatory mechanisms.

Theta-gamma phase-amplitude coupling in auditory cortex is modulated by language proficiency.

The coordination between the theta phase (3-7 Hz) and gamma power (25-35 Hz) oscillations (namely theta-gamma phase-amplitude coupling, PAC) in the auditory cortex has been proposed as an essential neural mechanism involved in speech processing. However, it has not been established how this mechanism is related to the efficiency with which a listener processes speech. Speech processing in a non-native language offers a useful opportunity to evaluate if theta-gamma PAC is modulated by the challenges imposed by the reception of speech input in a non-native language. The present study investigates how auditory theta-gamma PAC (recorded with magnetoencephalography) is modulated in both native and non-native speech reception. Participants were Spanish native (L1) speakers studying Basque (L2) at three different levels: beginner (Grade 1), intermediate (Grade 2), and advanced (Grade 3). We found that during L2 speech processing (i) theta-gamma PAC was more highly coordinated for intelligible compared to unintelligible speech; (ii) this coupling was modulated by proficiency in Basque being lower for beginners, higher for intermediate, and highest for advanced speakers (no difference observed in Spanish); (iii) gamma power did not differ between languages and groups. These findings highlight how the coordinated theta-gamma oscillatory activity is tightly related to speech comprehension: the stronger this coordination is, the more the comprehension system will proficiently parse the incoming speech input.

"Neural dynamics supporting longitudinal plasticity of action naming across languages: MEG evidence from bilingual brain tumor patients".

Previous evidence suggests that distinct ventral and dorsal streams respectively underpin the semantic processing of object and action knowledge. Recently, we found that brain tumor patients with dorsal gliomas in frontoparietal hubs show a selective longitudinal compensation (post-vs. pre-surgery) during the retrieval of lexico-semantic information about actions (but not objects), indexed by power increases in beta rhythms (13-28 Hz). Here, we move one-step further and ask whether a similar organizational principle also stands across the different languages a bilingual speaks. To test this hypothesis, we combined a picture-naming task with MEG recordings and evaluated highly proficient Spanish-Basque bilinguals undergoing surgery for tumor resection in left frontoparietal regions. We assessed patients before and three months after surgery. At the behavioral level, we observed a similar performance across sessions irrespectively of the language at use, suggesting overall successful function preservation. At the oscillatory level, we found longitudinal selective power increases in beta for action naming in Spanish and Basque. Nevertheless, tumor resection triggered a differential reorganization of the L1 and the L2, with the latter one additionally recruiting the right hemisphere. Overall, our results provide evidence for (i) the specific involvement of frontoparietal regions in the semantic retrieval/representation of action knowledge across languages; (ii) a key role of beta oscillations as a signature of language compensation and (iii) the existence of divergent plasticity trajectories in L1 and L2 after surgery. By doing so, they provide new insights into the spectro-temporal dynamics supporting postoperative recovery in the bilingual brain.

Neurodevelopmental oscillatory basis of speech processing in noise.

Humans' extraordinary ability to understand speech in noise relies on multiple processes that develop with age. Using magnetoencephalography (MEG), we characterize the underlying neuromaturational basis by quantifying how cortical oscillations in 144 participants (aged 5-27 years) track phrasal and syllabic structures in connected speech mixed with different types of noise. While the extraction of prosodic cues from clear speech was stable during development, its maintenance in a multi-talker background matured rapidly up to age 9 and was associated with speech comprehension. Furthermore, while the extraction of subtler information provided by syllables matured at age 9, its maintenance in noisy backgrounds progressively matured until adulthood. Altogether, these results highlight distinct behaviorally relevant maturational trajectories for the neuronal signatures of speech perception. In accordance with grain-size proposals, neuromaturational milestones are reached increasingly late for linguistic units of decreasing size, with further delays incurred by noise.

Oscillatory dynamics underlying noun and verb production in highly proficient bilinguals.

Words representing objects (nouns) and words representing actions (verbs) are essential components of speech across languages. While there is evidence regarding the organizational principles governing neural representation of nouns and verbs in monolingual speakers, little is known about how this knowledge is represented in the bilingual brain. To address this gap, we recorded neuromagnetic signals while highly proficient Spanish-Basque bilinguals performed a picture-naming task and tracked the brain oscillatory dynamics underlying this process. We found theta (4-8 Hz) power increases and alpha-beta (8-25 Hz) power decreases irrespectively of the category and language at use in a time window classically associated to the controlled retrieval of lexico-semantic information. When comparing nouns and verbs within each language, we found theta power increases for verbs as compared to nouns in bilateral visual cortices and cognitive control areas including the left SMA and right middle temporal gyrus. In addition, stronger alpha-beta power decreases were observed for nouns as compared to verbs in visual cortices and semantic-related regions such as the left anterior temporal lobe and right premotor cortex. No differences were observed between categories across languages. Overall, our results suggest that noun and verb processing recruit partially different networks during speech production but that these category-based representations are similarly processed in the bilingual brain.

Maintenance cost in the processing of subject-verb dependencies.

Although research in sentence comprehension has suggested that processing long-distance dependencies involves maintenance between the elements that form the dependency, studies on maintenance of long-distance subject-verb (SV) dependencies are scarce. The few relevant studies have delivered mixed results using self-paced reading or phoneme-monitoring tasks. In the current study, we used eye tracking during reading to test whether maintaining a long-distance SV dependency results in a processing cost on an intervening adverbial clause. In Experiment 1, we studied this question in Spanish and found that both go-past reading times and regressions out of an adverbial clause to the previous regions were significantly increased when the clause interrupts a SV dependency compared to when the same clause doesn't interrupt this dependency. We then replicated these findings in English (Experiment 2), observing significantly increased go-past reading times on a clause interrupting a SV dependency. The current study provides the first eye-tracking data showing a maintenance cost in the processing of SV dependencies cross-linguistically. Sentence comprehension models should account for the maintenance cost generated by SV dependency processing, and future research should focus on the nature of the maintained representation. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Temporal uncertainty enhances suppression of neural responses to predictable visual stimuli.

Contextual information triggers predictions about the content ("what") of environmental stimuli to update an internal generative model of the surrounding world. However, visual information dynamically changes across time, and temporal predictability ("when") may influence the impact of internal predictions on visual processing. In this magnetoencephalography (MEG) study, we investigated how processing feature specific information ("what") is affected by temporal predictability ("when"). Participants (N = 16) were presented with four consecutive Gabor patches (entrainers) with constant spatial frequency but with variable orientation and temporal onset. A fifth target Gabor was presented after a longer delay and with higher or lower spatial frequency that participants had to judge. We compared the neural responses to entrainers where the Gabor orientation could, or could not be temporally predicted along the entrainer sequence, and with inter-entrainer timing that was constant (predictable), or variable (unpredictable). We observed suppression of evoked neural responses in the visual cortex for predictable stimuli. Interestingly, we found that temporal uncertainty increased expectation suppression. This suggests that in temporally uncertain scenarios the neurocognitive system invests less resources in integrating bottom-up information. Multivariate pattern analysis showed that predictable visual features could be decoded from neural responses. Temporal uncertainty did not affect decoding accuracy for early visual responses, with the feature specificity of early visual neural activity preserved across conditions. However, decoding accuracy was less sustained over time for temporally jittered than for isochronous predictable visual stimuli. These findings converge to suggest that the cognitive system processes visual features of temporally predictable stimuli in higher detail, while processing temporally uncertain stimuli may rely more heavily on abstract internal expectations.

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 children's 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.

One Way or Another: Cortical Language Areas Flexibly Adapt Processing Strategies to Perceptual And Contextual Properties of Speech.

Cortical circuits rely on the temporal regularities of speech to optimize signal parsing for sound-to-meaning mapping. Bottom-up speech analysis is accelerated by top-down predictions about upcoming words. In everyday communications, however, listeners are regularly presented with challenging input-fluctuations of speech rate or semantic content. In this study, we asked how reducing speech temporal regularity affects its processing-parsing, phonological analysis, and ability to generate context-based predictions. To ensure that spoken sentences were natural and approximated semantic constraints of spontaneous speech we built a neural network to select stimuli from large corpora. We analyzed brain activity recorded with magnetoencephalography during sentence listening using evoked responses, speech-to-brain synchronization and representational similarity analysis. For normal speech theta band (6.5-8 Hz) speech-to-brain synchronization was increased and the left fronto-temporal areas generated stronger contextual predictions. The reverse was true for temporally irregular speech-weaker theta synchronization and reduced top-down effects. Interestingly, delta-band (0.5 Hz) speech tracking was greater when contextual/semantic predictions were lower or if speech was temporally jittered. We conclude that speech temporal regularity is relevant for (theta) syllabic tracking and robust semantic predictions while the joint support of temporal and contextual predictability reduces word and phrase-level cortical tracking (delta).

Language Proficiency Entails Tuning Cortical Activity to Second Language Speech.

Cortical tracking of linguistic structures in speech, such as phrases (<3 Hz, delta band) and syllables (3-8 Hz, theta band), is known to be crucial for speech comprehension. However, it has not been established whether this effect is related to language proficiency. Here, we investigate how auditory cortical activity in second language (L2) learners tracked L2 speech. Using magnetoencephalography, we recorded brain activity from participants listening to Spanish and Basque. Participants were Spanish native (L1) language speakers studying Basque (L2) at the same language center at three different levels: beginner (Grade 1), intermediate (Grade 2), and advanced (Grade 3). We found that 1) both delta and theta tracking to L2 speech in the auditory cortex were related to L2 learning proficiency and that 2) top-down modulations of activity in the left auditory regions during L2 speech listening-by the left inferior frontal and motor regions in delta band and by the left middle temporal regions in theta band-were also related to L2 proficiency. Altogether, these results indicate that the ability to learn an L2 is related to successful cortical tracking of L2 speech and its modulation by neuronal oscillations in higher-order cortical regions.

Speech-brain phase coupling is enhanced in low contextual semantic predictability conditions.

Semantic prediction and cortical entrainment to the acoustic landmarks of the speech envelope are two fundamental yet qualitatively different mechanisms that facilitate speech comprehension. However, it is not clear how and to what extent those mechanisms interact with each other. On the one hand, richer semantic context could enhance the perceptual representation of a predictable stimulus, thus improving speech entrainment. On the other hand, pre-activating an upcoming item could inhibit further bottom-up analyses to minimize processing costs, thus weakening speech entrainment. To test these competing hypotheses, we recorded EEG activity from 27 participants while they listened to a 14-min recording of text reading. The passage contained target words presented twice: once in a highly constraining and once in a minimally constraining context. First, we measured event related potentials on target words in the two conditions. In line with previous research, we showed that semantic predictability modulated the N400 amplitude: words in minimally constraining contexts elicited larger negative amplitudes than words in highly constraining contexts between 250 and 450 ms. Second, we evaluated speech entrainment effects by analyzing phase alignment between neural activity and the envelope of target words. Importantly, we found increased speech entrainment for words in minimally constraining compared to highly constraining contexts between 400 and 450 ms. Both effects were located in central electrodes and were significantly correlated. Our results indicate a trade-off between semantic pre-activation and cortical entrainment to speech and support the cost minimization hypothesis.

Impaired neural response to speech edges in dyslexia.

Speech comprehension has been proposed to critically rely on oscillatory cortical tracking, that is, phase alignment of neural oscillations to the slow temporal modulations (envelope) of speech. Speech-brain entrainment is readjusted over time as transient events (edges) in speech lead to speech-brain phase realignment. Auditory behavioral research suggests that phonological deficits in dyslexia are linked to difficulty in discriminating speech edges. Importantly, research to date has not specifically examined neural responses to speech edges in dyslexia. In the present study, we used MEG to record brain activity from normal and dyslexic readers while they listened to speech. We computed phase locking values (PLVs) to evaluate phase entrainment between neural oscillations and the speech envelope time-locked to edge onsets. In both groups, we observed that edge onsets induced phase resets in the auditory oscillations tracking speech, thereby enhancing their entrainment to speech. Importantly, dyslexic readers showed weaker PLVs compared to normal readers in left auditory regions from ~.15 sec to ~.65 sec after edge onset. Our results indicate that the neural mechanism that adapts cortical entrainment to the speech envelope is impaired in dyslexia. These findings here are consistent with the temporal sampling theory of developmental dyslexia.

Oscillatory and structural signatures of language plasticity in brain tumor patients: A longitudinal study.

Recent evidence suggests that damage to the language network triggers its functional reorganization. Yet, the spectro-temporal fingerprints of this plastic rearrangement and its relation to anatomical changes is less well understood. Here, we combined magnetoencephalographic recordings with a proxy measure of white matter to investigate oscillatory activity supporting language plasticity and its relation to structural reshaping. First, cortical dynamics were acquired in a group of healthy controls during object and action naming. Results showed segregated beta (13-28 Hz) power decreases in left ventral and dorsal pathways, in a time-window associated to lexico-semantic processing (~250-500 ms). Six patients with left tumors invading either ventral or dorsal regions performed the same naming task before and 3 months after surgery for tumor resection. When longitudinally comparing patients' responses we found beta compensation mimicking the category-based segregation showed by controls, with ventral and dorsal damage leading to selective compensation for object and action naming, respectively. At the structural level, all patients showed preoperative changes in white matter tracts possibly linked to plasticity triggered by tumor growth. Furthermore, in some patients, structural changes were also evident after surgery and showed associations with longitudinal changes in beta power lateralization toward the contralesional hemisphere. Overall, our findings support the existence of anatomo-functional dependencies in language reorganization and highlight the potential role of oscillatory markers in tracking longitudinal plasticity in brain tumor patients. By doing so, they provide valuable information for mapping preoperative and postoperative neural reshaping and plan surgical strategies to preserve language function and patient's quality of life.

Early dissociation of numbers and letters in the human brain.

Numbers and letters are culturally created symbols which are learned through repeated training. This experience leads to a functional specialization of the perceptual system of our brain. Recent evidence suggests a neural dissociation between these two symbols. While previous literature has shown that letters elicit a left lateralized neural response, new studies suggest that numbers elicit preferentially a bilateral or right lateralized response. However, the time course of the neural patterns that characterize this dissociation is still underspecified. In the present study, we investigated with magnetoencephalography (MEG) the spatio-temporal dynamics of the neural response generated by numbers, letters and perceptually matched false fonts presented visually. Twenty-five healthy adults were recorded while participants performed a dot detection task. By including two experiments, we were able to study the effects of single characters as well as those of strings of characters. The signal analysis was focused on the event related fields (ERF) of the MEG signal in the sensors and in the source space. The main results of our study showed an early (<200 msec) preferential dissociation between single numbers and single letters on occipito-temporal sensors. When comparing strings of numbers and pseudowords, they differed also over prefrontal regions of the brain. These data offer a new example of acquired category-specific responses in the human brain.