Displaying 1 - 27 of 27
  • Bonandrini, R., Gornetti, E., & Paulesu, E. (2024). A meta-analytical account of the functional lateralization of the reading network. Cortex, 177, 363-384. doi:10.1016/j.cortex.2024.05.015.

    Abstract

    The observation that the neural correlates of reading are left-lateralized is ubiquitous in the cognitive neuroscience and neuropsychological literature. Still, reading is served by a constellation of neural units, and the extent to which these units are consistently left-lateralized is unclear. In this regard, the functional lateralization of the fusiform gyrus is of particular interest, by virtue of its hypothesized role as a “visual word form area”. A quantitative Activation Likelihood Estimation meta-analysis was conducted on activation foci from 35 experiments investigating silent reading, and both a whole-brain and a bayesian ROI-based approach were used to assess the lateralization of the data submitted to meta-analysis. Perirolandic areas showed the highest level of left-lateralization, the fusiform cortex and the parietal cortex exhibited only a moderate pattern of left-lateralization, while in the occipital, insular cortices and in the cerebellum the lateralization turned out to be the lowest observed. The relatively limited functional lateralization of the fusiform gyrus was further explored in a regression analysis on the lateralization profile of each study. The functional lateralization of the fusiform gyrus during reading was positively associated with the lateralization of the precentral and inferior occipital gyri and negatively associated with the lateralization of the triangular portion of the inferior frontal gyrus and of the temporal pole. Overall, the present data highlight how lateralization patterns differ within the reading network. Furthermore, the present data highlight how the functional lateralization of the fusiform gyrus during reading is related to the degree of functional lateralization of other language brain areas.
  • Coopmans, C. W., Mai, A., & Martin, A. E. (2024). “Not” in the brain and behavior. PLOS Biology, 22: e3002656. doi:10.1371/journal.pbio.3002656.
  • Ding, R., Ten Oever, S., & Martin, A. E. (2024). Delta-band activity underlies referential meaning representation during pronoun resolution. Journal of Cognitive Neuroscience, 36(7), 1472-1492. doi:10.1162/jocn_a_02163.

    Abstract

    Human language offers a variety of ways to create meaning, one of which is referring to entities, objects, or events in the world. One such meaning maker is understanding to whom or to what a pronoun in a discourse refers to. To understand a pronoun, the brain must access matching entities or concepts that have been encoded in memory from previous linguistic context. Models of language processing propose that internally stored linguistic concepts, accessed via exogenous cues such as phonological input of a word, are represented as (a)synchronous activities across a population of neurons active at specific frequency bands. Converging evidence suggests that delta band activity (1–3 Hz) is involved in temporal and representational integration during sentence processing. Moreover, recent advances in the neurobiology of memory suggest that recollection engages neural dynamics similar to those which occurred during memory encoding. Integrating from these two research lines, we here tested the hypothesis that neural dynamic patterns, especially in delta frequency range, underlying referential meaning representation, would be reinstated during pronoun resolution. By leveraging neural decoding techniques (i.e., representational similarity analysis) on a magnetoencephalogram data set acquired during a naturalistic story-listening task, we provide evidence that delta-band activity underlies referential meaning representation. Our findings suggest that, during spoken language comprehension, endogenous linguistic representations such as referential concepts may be proactively retrieved and represented via activation of their underlying dynamic neural patterns.
  • Mai, A., Riès, S., Ben-Haim, S., Shih, J. J., & Gentner, T. Q. (2024). Acoustic and language-specific sources for phonemic abstraction from speech. Nature Communications, 15: 677. doi:10.1038/s41467-024-44844-9.

    Abstract

    Spoken language comprehension requires abstraction of linguistic information from speech, but the interaction between auditory and linguistic processing of speech remains poorly understood. Here, we investigate the nature of this abstraction using neural responses recorded intracranially while participants listened to conversational English speech. Capitalizing on multiple, language-specific patterns where phonological and acoustic information diverge, we demonstrate the causal efficacy of the phoneme as a unit of analysis and dissociate the unique contributions of phonemic and spectrographic information to neural responses. Quantitive higher-order response models also reveal that unique contributions of phonological information are carried in the covariance structure of the stimulus-response relationship. This suggests that linguistic abstraction is shaped by neurobiological mechanisms that involve integration across multiple spectro-temporal features and prior phonological information. These results link speech acoustics to phonology and morphosyntax, substantiating predictions about abstractness in linguistic theory and providing evidence for the acoustic features that support that abstraction.

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  • Meinhardt, E., Mai, A., Baković, E., & McCollum, A. (2024). Weak determinism and the computational consequences of interaction. Natural Language & Linguistic Theory, 42, 1191-1232. doi:10.1007/s11049-023-09578-1.

    Abstract

    Recent work has claimed that (non-tonal) phonological patterns are subregular (Heinz 2011a,b, 2018; Heinz and Idsardi 2013), occupying a delimited proper subregion of the regular functions—the weakly deterministic (WD) functions (Heinz and Lai 2013; Jardine 2016). Whether or not it is correct (McCollum et al. 2020a), this claim can only be properly assessed given a complete and accurate definition of WD functions. We propose such a definition in this article, patching unintended holes in Heinz and Lai’s (2013) original definition that we argue have led to the incorrect classification of some phonological patterns as WD. We start from the observation that WD patterns share a property that we call unbounded semiambience, modeled after the analogous observation by Jardine (2016) about non-deterministic (ND) patterns and their unbounded circumambience. Both ND and WD functions can be broken down into compositions of deterministic (subsequential) functions (Elgot and Mezei 1965; Heinz and Lai 2013) that read an input string from opposite directions; we show that WD functions are those for which these deterministic composands do not interact in a way that is familiar from the theoretical phonology literature. To underscore how this concept of interaction neatly separates the WD class of functions from the strictly more expressive ND class, we provide analyses of the vowel harmony patterns of two Eastern Nilotic languages, Maasai and Turkana, using bimachines, an automaton type that represents unbounded bidirectional dependencies explicitly. These analyses make clear that there is interaction between deterministic composands when (and only when) the output of a given input element of a string is simultaneously dependent on information from both the left and the right: ND functions are those that involve interaction, while WD functions are those that do not.
  • Slaats, S., Meyer, A. S., & Martin, A. E. (2024). Lexical surprisal shapes the time course of syntactic structure building. Neurobiology of Language, 5(4), 942-980. doi:10.1162/nol_a_00155.

    Abstract

    When we understand language, we recognize words and combine them into sentences. In this article, we explore the hypothesis that listeners use probabilistic information about words to build syntactic structure. Recent work has shown that lexical probability and syntactic structure both modulate the delta-band (<4 Hz) neural signal. Here, we investigated whether the neural encoding of syntactic structure changes as a function of the distributional properties of a word. To this end, we analyzed MEG data of 24 native speakers of Dutch who listened to three fairytales with a total duration of 49 min. Using temporal response functions and a cumulative model-comparison approach, we evaluated the contributions of syntactic and distributional features to the variance in the delta-band neural signal. This revealed that lexical surprisal values (a distributional feature), as well as bottom-up node counts (a syntactic feature) positively contributed to the model of the delta-band neural signal. Subsequently, we compared responses to the syntactic feature between words with high- and low-surprisal values. This revealed a delay in the response to the syntactic feature as a consequence of the surprisal value of the word: high-surprisal values were associated with a delayed response to the syntactic feature by 150–190 ms. The delay was not affected by word duration, and did not have a lexical origin. These findings suggest that the brain uses probabilistic information to infer syntactic structure, and highlight an importance for the role of time in this process.

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    supplementary data
  • Ten Oever, S., & Martin, A. E. (2024). Interdependence of “what” and “when” in the brain. Journal of Cognitive Neuroscience, 36(1), 167-186. doi:10.1162/jocn_a_02067.

    Abstract

    From a brain's-eye-view, when a stimulus occurs and what it is are interrelated aspects of interpreting the perceptual world. Yet in practice, the putative perceptual inferences about sensory content and timing are often dichotomized and not investigated as an integrated process. We here argue that neural temporal dynamics can influence what is perceived, and in turn, stimulus content can influence the time at which perception is achieved. This computational principle results from the highly interdependent relationship of what and when in the environment. Both brain processes and perceptual events display strong temporal variability that is not always modeled; we argue that understanding—and, minimally, modeling—this temporal variability is key for theories of how the brain generates unified and consistent neural representations and that we ignore temporal variability in our analysis practice at the peril of both data interpretation and theory-building. Here, we review what and when interactions in the brain, demonstrate via simulations how temporal variability can result in misguided interpretations and conclusions, and outline how to integrate and synthesize what and when in theories and models of brain computation.
  • Ten Oever, S., Titone, L., te Rietmolen, N., & Martin, A. E. (2024). Phase-dependent word perception emerges from region-specific sensitivity to the statistics of language. Proceedings of the National Academy of Sciences of the United States of America, 121(3): e2320489121. doi:10.1073/pnas.2320489121.

    Abstract

    Neural oscillations reflect fluctuations in excitability, which biases the percept of ambiguous sensory input. Why this bias occurs is still not fully understood. We hypothesized that neural populations representing likely events are more sensitive, and thereby become active on earlier oscillatory phases, when the ensemble itself is less excitable. Perception of ambiguous input presented during less-excitable phases should therefore be biased toward frequent or predictable stimuli that have lower activation thresholds. Here, we show such a frequency bias in spoken word recognition using psychophysics, magnetoencephalography (MEG), and computational modelling. With MEG, we found a double dissociation, where the phase of oscillations in the superior temporal gyrus and medial temporal gyrus biased word-identification behavior based on phoneme and lexical frequencies, respectively. This finding was reproduced in a computational model. These results demonstrate that oscillations provide a temporal ordering of neural activity based on the sensitivity of separable neural populations.
  • Weissbart, H., & Martin, A. E. (2024). The structure and statistics of language jointly shape cross-frequency neural dynamics during spoken language comprehension. Nature Communications, 15: 8850. doi:10.1038/s41467-024-53128-1.

    Abstract

    Humans excel at extracting structurally-determined meaning from speech despite inherent physical variability. This study explores the brain’s ability to predict and understand spoken language robustly. It investigates the relationship between structural and statistical language knowledge in brain dynamics, focusing on phase and amplitude modulation. Using syntactic features from constituent hierarchies and surface statistics from a transformer model as predictors of forward encoding models, we reconstructed cross-frequency neural dynamics from MEG data during audiobook listening. Our findings challenge a strict separation of linguistic structure and statistics in the brain, with both aiding neural signal reconstruction. Syntactic features have a more temporally spread impact, and both word entropy and the number of closing syntactic constituents are linked to the phase-amplitude coupling of neural dynamics, implying a role in temporal prediction and cortical oscillation alignment during speech processing. Our results indicate that structured and statistical information jointly shape neural dynamics during spoken language comprehension and suggest an integration process via a cross-frequency coupling mechanism
  • Yang, J. (2024). Rethinking tokenization: Crafting better tokenizers for large language models. International Journal of Chinese Linguistics, 11(1), 94-109. doi:10.1075/ijchl.00023.yan.

    Abstract

    Tokenization significantly influences language models (LMs)’ performance. This paper traces the evolution of tokenizers from word-level to subword-level, analyzing how they balance tokens and types to enhance model adaptability while controlling complexity. Despite subword tokenizers like Byte Pair Encoding (BPE) overcoming many word tokenizer limitations, they encounter difficulties in handling non-Latin languages and depend heavily on extensive training data and computational resources to grasp the nuances of multiword expressions (MWEs). This article argues that tokenizers, more than mere technical tools, should drawing inspiration from the cognitive science about human language processing. This study then introduces the “Principle of Least Effort” from cognitive science, that humans naturally seek to reduce cognitive effort, and discusses the benefits of this principle for tokenizer development. Based on this principle, the paper proposes that the Less-is-Better (LiB) model could be a new approach for LLM tokenizer. The LiB model can autonomously learn an integrated vocabulary consisting of subwords, words, and MWEs, which effectively reduces both the numbers of tokens and types. Comparative evaluations show that the LiB tokenizer outperforms existing word and BPE tokenizers, presenting an innovative method for tokenizer development, and hinting at the possibility of future cognitive science-based tokenizers being more efficient.
  • Zhao, J., Martin, A. E., & Coopmans, C. W. (2024). Structural and sequential regularities modulate phrase-rate neural tracking. Scientific Reports, 14: 16603. doi:10.1038/s41598-024-67153-z.

    Abstract

    Electrophysiological brain activity has been shown to synchronize with the quasi-regular repetition of grammatical phrases in connected speech—so-called phrase-rate neural tracking. Current debate centers around whether this phenomenon is best explained in terms of the syntactic properties of phrases or in terms of syntax-external information, such as the sequential repetition of parts of speech. As these two factors were confounded in previous studies, much of the literature is compatible with both accounts. Here, we used electroencephalography (EEG) to determine if and when the brain is sensitive to both types of information. Twenty native speakers of Mandarin Chinese listened to isochronously presented streams of monosyllabic words, which contained either grammatical two-word phrases (e.g., catch fish, sell house) or non-grammatical word combinations (e.g., full lend, bread far). Within the grammatical conditions, we varied two structural factors: the position of the head of each phrase and the type of attachment. Within the non-grammatical conditions, we varied the consistency with which parts of speech were repeated. Tracking was quantified through evoked power and inter-trial phase coherence, both derived from the frequency-domain representation of EEG responses. As expected, neural tracking at the phrase rate was stronger in grammatical sequences than in non-grammatical sequences without syntactic structure. Moreover, it was modulated by both attachment type and head position, revealing the structure-sensitivity of phrase-rate tracking. We additionally found that the brain tracks the repetition of parts of speech in non-grammatical sequences. These data provide an integrative perspective on the current debate about neural tracking effects, revealing that the brain utilizes regularities computed over multiple levels of linguistic representation in guiding rhythmic computation.
  • Zioga, I., Zhou, Y. J., Weissbart, H., Martin, A. E., & Haegens, S. (2024). Alpha and beta oscillations differentially support word production in a rule-switching task. eNeuro, 11(4): ENEURO.0312-23.2024. doi:10.1523/ENEURO.0312-23.2024.

    Abstract

    Research into the role of brain oscillations in basic perceptual and cognitive functions has suggested that the alpha rhythm reflects functional inhibition while the beta rhythm reflects neural ensemble (re)activation. However, little is known regarding the generalization of these proposed fundamental operations to linguistic processes, such as speech comprehension and production. Here, we recorded magnetoencephalography in participants performing a novel rule-switching paradigm. Specifically, Dutch native speakers had to produce an alternative exemplar from the same category or a feature of a given target word embedded in spoken sentences (e.g., for the word “tuna”, an exemplar from the same category—“seafood”—would be “shrimp”, and a feature would be “pink”). A cue indicated the task rule—exemplar or feature—either before (pre-cue) or after (retro-cue) listening to the sentence. Alpha power during the working memory delay was lower for retro-cue compared with that for pre-cue in the left hemispheric language-related regions. Critically, alpha power negatively correlated with reaction times, suggestive of alpha facilitating task performance by regulating inhibition in regions linked to lexical retrieval. Furthermore, we observed a different spatiotemporal pattern of beta activity for exemplars versus features in the right temporoparietal regions, in line with the proposed role of beta in recruiting neural networks for the encoding of distinct categories. Overall, our study provides evidence for the generalizability of the role of alpha and beta oscillations from perceptual to more “complex, linguistic processes” and offers a novel task to investigate links between rule-switching, working memory, and word production.
  • Bai, F., Meyer, A. S., & Martin, A. E. (2022). Neural dynamics differentially encode phrases and sentences during spoken language comprehension. PLoS Biology, 20(7): e3001713. doi:10.1371/journal.pbio.3001713.

    Abstract

    Human language stands out in the natural world as a biological signal that uses a structured system to combine the meanings of small linguistic units (e.g., words) into larger constituents (e.g., phrases and sentences). However, the physical dynamics of speech (or sign) do not stand in a one-to-one relationship with the meanings listeners perceive. Instead, listeners infer meaning based on their knowledge of the language. The neural readouts of the perceptual and cognitive processes underlying these inferences are still poorly understood. In the present study, we used scalp electroencephalography (EEG) to compare the neural response to phrases (e.g., the red vase) and sentences (e.g., the vase is red), which were close in semantic meaning and had been synthesized to be physically indistinguishable. Differences in structure were well captured in the reorganization of neural phase responses in delta (approximately <2 Hz) and theta bands (approximately 2 to 7 Hz),and in power and power connectivity changes in the alpha band (approximately 7.5 to 13.5 Hz). Consistent with predictions from a computational model, sentences showed more power, more power connectivity, and more phase synchronization than phrases did. Theta–gamma phase–amplitude coupling occurred, but did not differ between the syntactic structures. Spectral–temporal response function (STRF) modeling revealed different encoding states for phrases and sentences, over and above the acoustically driven neural response. Our findings provide a comprehensive description of how the brain encodes and separates linguistic structures in the dynamics of neural responses. They imply that phase synchronization and strength of connectivity are readouts for the constituent structure of language. The results provide a novel basis for future neurophysiological research on linguistic structure representation in the brain, and, together with our simulations, support time-based binding as a mechanism of structure encoding in neural dynamics.
  • Bai, F. (2022). Neural representation of speech segmentation and syntactic structure discrimination. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Coopmans, C. W., De Hoop, H., Kaushik, K., Hagoort, P., & Martin, A. E. (2022). Hierarchy in language interpretation: Evidence from behavioural experiments and computational modelling. Language, Cognition and Neuroscience, 37(4), 420-439. doi:10.1080/23273798.2021.1980595.

    Abstract

    It has long been recognised that phrases and sentences are organised hierarchically, but many computational models of language treat them as sequences of words without computing constituent structure. Against this background, we conducted two experiments which showed that participants interpret ambiguous noun phrases, such as second blue ball, in terms of their abstract hierarchical structure rather than their linear surface order. When a neural network model was tested on this task, it could simulate such “hierarchical” behaviour. However, when we changed the training data such that they were not entirely unambiguous anymore, the model stopped generalising in a human-like way. It did not systematically generalise to novel items, and when it was trained on ambiguous trials, it strongly favoured the linear interpretation. We argue that these models should be endowed with a bias to make generalisations over hierarchical structure in order to be cognitively adequate models of human language.
  • Coopmans, C. W., De Hoop, H., Hagoort, P., & Martin, A. E. (2022). Effects of structure and meaning on cortical tracking of linguistic units in naturalistic speech. Neurobiology of Language, 3(3), 386-412. doi:10.1162/nol_a_00070.

    Abstract

    Recent research has established that cortical activity “tracks” the presentation rate of syntactic phrases in continuous speech, even though phrases are abstract units that do not have direct correlates in the acoustic signal. We investigated whether cortical tracking of phrase structures is modulated by the extent to which these structures compositionally determine meaning. To this end, we recorded electroencephalography (EEG) of 38 native speakers who listened to naturally spoken Dutch stimuli in different conditions, which parametrically modulated the degree to which syntactic structure and lexical semantics determine sentence meaning. Tracking was quantified through mutual information between the EEG data and either the speech envelopes or abstract annotations of syntax, all of which were filtered in the frequency band corresponding to the presentation rate of phrases (1.1–2.1 Hz). Overall, these mutual information analyses showed stronger tracking of phrases in regular sentences than in stimuli whose lexical-syntactic content is reduced, but no consistent differences in tracking between sentences and stimuli that contain a combination of syntactic structure and lexical content. While there were no effects of compositional meaning on the degree of phrase-structure tracking, analyses of event-related potentials elicited by sentence-final words did reveal meaning-induced differences between conditions. Our findings suggest that cortical tracking of structure in sentences indexes the internal generation of this structure, a process that is modulated by the properties of its input, but not by the compositional interpretation of its output.

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    supplementary information
  • Dingemanse, M., Liesenfeld, A., & Woensdregt, M. (2022). Convergent cultural evolution of continuers (mhmm). In A. Ravignani, R. Asano, D. Valente, F. Ferretti, S. Hartmann, M. Hayashi, Y. Jadoul, M. Martins, Y. Oseki, E. D. Rodrigues, O. Vasileva, & S. Wacewicz (Eds.), The Evolution of Language: Proceedings of the Joint Conference on Language Evolution (JCoLE) (pp. 160-167). Nijmegen: Joint Conference on Language Evolution (JCoLE). doi:10.31234/osf.io/65c79.

    Abstract

    Continuers —words like mm, mmhm, uhum and the like— are among the most frequent types of responses in conversation. They play a key role in joint action coordination by showing positive evidence of understanding and scaffolding narrative delivery. Here we investigate the hypothesis that their functional importance along with their conversational ecology places selective pressures on their form and may lead to cross-linguistic similarities through convergent cultural evolution. We compare continuer tokens in linguistically diverse conversational corpora and find languages make available highly similar forms. We then approach the causal mechanism of convergent cultural evolution using exemplar modelling, simulating the process by which a combination of effort minimization and functional specialization may push continuers to a particular region of phonological possibility space. By combining comparative linguistics and computational modelling we shed new light on the question of how language structure is shaped by and for social interaction.
  • Doumas, L. A. A., Puebla, G., Martin, A. E., & Hummel, J. E. (2022). A theory of relation learning and cross-domain generalization. Psychological Review, 129(5), 999-1041. doi:10.1037/rev0000346.

    Abstract

    People readily generalize knowledge to novel domains and stimuli. We present a theory, instantiated in a computational model, based on the idea that cross-domain generalization in humans is a case of analogical inference over structured (i.e., symbolic) relational representations. The model is an extension of the Learning and Inference with Schemas and Analogy (LISA; Hummel & Holyoak, 1997, 2003) and Discovery of Relations by Analogy (DORA; Doumas et al., 2008) models of relational inference and learning. The resulting model learns both the content and format (i.e., structure) of relational representations from nonrelational inputs without supervision, when augmented with the capacity for reinforcement learning it leverages these representations to learn about individual domains, and then generalizes to new domains on the first exposure (i.e., zero-shot learning) via analogical inference. We demonstrate the capacity of the model to learn structured relational representations from a variety of simple visual stimuli, and to perform cross-domain generalization between video games (Breakout and Pong) and between several psychological tasks. We demonstrate that the model’s trajectory closely mirrors the trajectory of children as they learn about relations, accounting for phenomena from the literature on the development of children’s reasoning and analogy making. The model’s ability to generalize between domains demonstrates the flexibility afforded by representing domains in terms of their underlying relational structure, rather than simply in terms of the statistical relations between their inputs and outputs.
  • Heesen, R., Fröhlich, M., Sievers, C., Woensdregt, M., & Dingemanse, M. (2022). Coordinating social action: A primer for the cross-species investigation of communicative repair. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 377(1859): 20210110. doi:10.1098/rstb.2021.0110.

    Abstract

    Human joint action is inherently cooperative, manifested in the collaborative efforts of participants to minimize communicative trouble through interactive repair. Although interactive repair requires sophisticated cognitive abilities,
    it can be dissected into basic building blocks shared with non-human animal species. A review of the primate literature shows that interactionally contingent signal sequences are at least common among species of nonhuman great apes, suggesting a gradual evolution of repair. To pioneer a cross-species assessment of repair this paper aims at (i) identifying necessary precursors of human interactive repair; (ii) proposing a coding framework for its comparative study in humans and non-human species; and (iii) using this framework to analyse examples of interactions of humans (adults/children) and non-human great apes. We hope this paper will serve as a primer for cross-species comparisons of communicative breakdowns and how they are repaired.
  • Janssens, S. E. W., Sack, A. T., Ten Oever, S., & Graaf, T. A. (2022). Calibrating rhythmic stimulation parameters to individual electroencephalography markers: The consistency of individual alpha frequency in practical lab settings. European Journal of Neuroscience, 55(11/12), 3418-3437. doi:10.1111/ejn.15418.

    Abstract

    Rhythmic stimulation can be applied to modulate neuronal oscillations. Such ‘entrainment’ is optimized when stimulation frequency is individually calibrated based on magneto/encephalography markers. It remains unknown how consistent such individual markers are across days/sessions, within a session, or across cognitive states, hemispheres and estimation methods, especially in a realistic, practical, lab setting. We here estimated individual alpha frequency (IAF) repeatedly from short electroencephalography (EEG) measurements at rest or during an attention task (cognitive state), using single parieto-occipital electrodes in 24 participants on 4 days (between-sessions), with multiple measurements over an hour on 1 day (within-session). First, we introduce an algorithm to automatically reject power spectra without a sufficiently clear peak to ensure unbiased IAF estimations. Then we estimated IAF via the traditional ‘maximum’ method and a ‘Gaussian fit’ method. IAF was reliable within- and between-sessions for both cognitive states and hemispheres, though task-IAF estimates tended to be more variable. Overall, the ‘Gaussian fit’ method was more reliable than the ‘maximum’ method. Furthermore, we evaluated how far from an approximated ‘true’ task-related IAF the selected ‘stimulation frequency’ was, when calibrating this frequency based on a short rest-EEG, a short task-EEG, or simply selecting 10 Hz for all participants. For the ‘maximum’ method, rest-EEG calibration was best, followed by task-EEG, and then 10 Hz. For the ‘Gaussian fit’ method, rest-EEG and task-EEG-based calibration were similarly accurate, and better than 10 Hz. These results lead to concrete recommendations about valid, and automated, estimation of individual oscillation markers in experimental and clinical settings.
  • Janssens, S. E., Ten Oever, S., Sack, A. T., & de Graaf, T. A. (2022). “Broadband Alpha Transcranial Alternating Current Stimulation”: Exploring a new biologically calibrated brain stimulation protocol. NeuroImage, 253: 119109. doi:10.1016/j.neuroimage.2022.119109.

    Abstract

    Transcranial alternating current stimulation (tACS) can be used to study causal contributions of oscillatory brain mechanisms to cognition and behavior. For instance, individual alpha frequency (IAF) tACS was reported to enhance alpha power and impact visuospatial attention performance. Unfortunately, such results have been inconsistent and difficult to replicate. In tACS, stimulation generally involves one frequency, sometimes individually calibrated to a peak value observed in an M/EEG power spectrum. Yet, the ‘peak’ actually observed in such power spectra often contains a broader range of frequencies, raising the question whether a biologically calibrated tACS protocol containing this fuller range of alpha-band frequencies might be more effective. Here, we introduce ‘Broadband-alpha-tACS’, a complex individually calibrated electrical stimulation protocol. We band-pass filtered left posterior resting-state EEG data around the IAF (+/- 2 Hz), and converted that time series into an electrical waveform for tACS stimulation of that same left posterior parietal cortex location. In other words, we stimulated a brain region with a ‘replay’ of its own alpha-band frequency content, based on spontaneous activity. Within-subjects (N=24), we compared to a sham tACS session the effects of broadband-alpha tACS, power-matched spectral inverse (‘alpha-removed’) control tACS, and individual alpha frequency tACS, on EEG alpha power and performance in an endogenous attention task previously reported to be affected by alpha tACS. Broadband-alpha-tACS significantly modulated attention task performance (i.e., reduced the rightward visuospatial attention bias in trials without distractors, and reduced attention benefits). Alpha-removed tACS also reduced the rightward visuospatial attention bias. IAF-tACS did not significantly modulate attention task performance compared to sham tACS, but also did not statistically significantly differ from broadband-alpha-tACS. This new broadband-alpha tACS approach seems promising, but should be further explored and validated in future studies.

    Additional information

    supplementary materials
  • Kemmerer, S. K., Sack, A. T., de Graaf, T. A., Ten Oever, S., De Weerd, P., & Schuhmann, T. (2022). Frequency-specific transcranial neuromodulation of alpha power alters visuospatial attention performance. Brain Research, 1782: 147834. doi:10.1016/j.brainres.2022.147834.

    Abstract

    Transcranial alternating current stimulation (tACS) at 10 Hz has been shown to modulate spatial attention. However, the frequency-specificity and the oscillatory changes underlying this tACS effect are still largely unclear. Here, we applied high-definition tACS at individual alpha frequency (IAF), two control frequencies (IAF+/-2Hz) and sham to the left posterior parietal cortex and measured its effects on visuospatial attention performance and offline alpha power (using electroencephalography, EEG). We revealed a behavioural and electrophysiological stimulation effect relative to sham for IAF but not control frequency stimulation conditions: there was a leftward lateralization of alpha power for IAF tACS, which differed from sham for the first out of three minutes following tACS. At a high value of this EEG effect (moderation effect), we observed a leftward attention bias relative to sham. This effect was task-specific, i.e., it could be found in an endogenous attention but not in a detection task. Only in the IAF tACS condition, we also found a correlation between the magnitude of the alpha lateralization and the attentional bias effect. Our results support a functional role of alpha oscillations in visuospatial attention and the potential of tACS to modulate it. The frequency-specificity of the effects suggests that an individualization of the stimulation frequency is necessary in heterogeneous target groups with a large variation in IAF.

    Additional information

    supplementary data
  • Kemmerer, S. K., De Graaf, T. A., Ten Oever, S., Erkens, M., De Weerd, P., & Sack, A. T. (2022). Parietal but not temporoparietal alpha-tACS modulates endogenous visuospatial attention. Cortex, 154, 149-166. doi:10.1016/j.cortex.2022.01.021.

    Abstract

    Visuospatial attention can either be voluntarily directed (endogenous/top-down attention) or automatically triggered (exogenous/bottom-up attention). Recent research showed that dorsal parietal transcranial alternating current stimulation (tACS) at alpha frequency modulates the spatial attentional bias in an endogenous but not in an exogenous visuospatial attention task. Yet, the reason for this task-specificity remains unexplored. Here, we tested whether this dissociation relates to the proposed differential role of the dorsal attention network (DAN) and ventral attention network (VAN) in endogenous and exogenous attention processes respectively. To that aim, we targeted the left and right dorsal parietal node of the DAN, as well as the left and right ventral temporoparietal node of the VAN using tACS at the individual alpha frequency. Every participant completed all four stimulation conditions and a sham condition in five separate sessions. During tACS, we assessed the behavioral visuospatial attention bias via an endogenous and exogenous visuospatial attention task. Additionally, we measured offline alpha power immediately before and after tACS using electroencephalography (EEG). The behavioral data revealed an effect of tACS on the endogenous but not exogenous attention bias, with a greater leftward bias during (sham-corrected) left than right hemispheric stimulation. In line with our hypothesis, this effect was brain area-specific, i.e., present for dorsal parietal but not ventral temporoparietal tACS. However, contrary to our expectations, there was no effect of ventral temporoparietal tACS on the exogenous visuospatial attention bias. Hence, no double dissociation between the two targeted attention networks. There was no effect of either tACS condition on offline alpha power. Our behavioral data reveal that dorsal parietal but not ventral temporoparietal alpha oscillations steer endogenous visuospatial attention. This brain-area specific tACS effect matches the previously proposed dissociation between the DAN and VAN and, by showing that the spatial attention bias effect does not generalize to any lateral posterior tACS montage, renders lateral cutaneous and retinal effects for the spatial attention bias in the dorsal parietal condition unlikely. Yet the absence of tACS effects on the exogenous attention task suggests that ventral temporoparietal alpha oscillations are not functionally relevant for exogenous visuospatial attention. We discuss the potential implications of this finding in the context of an emerging theory on the role of the ventral temporoparietal node.

    Additional information

    supplementary material
  • Ten Oever, S., Carta, S., Kaufeld, G., & Martin, A. E. (2022). Neural tracking of phrases in spoken language comprehension is automatic and task-dependent. eLife, 11: e77468. doi:10.7554/eLife.77468.

    Abstract

    Linguistic phrases are tracked in sentences even though there is no one-to-one acoustic phrase marker in the physical signal. This phenomenon suggests an automatic tracking of abstract linguistic structure that is endogenously generated by the brain. However, all studies investigating linguistic tracking compare conditions where either relevant information at linguistic timescales is available, or where this information is absent altogether (e.g., sentences versus word lists during passive listening). It is therefore unclear whether tracking at phrasal timescales is related to the content of language, or rather, results as a consequence of attending to the timescales that happen to match behaviourally relevant information. To investigate this question, we presented participants with sentences and word lists while recording their brain activity with magnetoencephalography (MEG). Participants performed passive, syllable, word, and word-combination tasks corresponding to attending to four different rates: one they would naturally attend to, syllable-rates, word-rates, and phrasal-rates, respectively. We replicated overall findings of stronger phrasal-rate tracking measured with mutual information for sentences compared to word lists across the classical language network. However, in the inferior frontal gyrus (IFG) we found a task effect suggesting stronger phrasal-rate tracking during the word-combination task independent of the presence of linguistic structure, as well as stronger delta-band connectivity during this task. These results suggest that extracting linguistic information at phrasal rates occurs automatically with or without the presence of an additional task, but also that IFG might be important for temporal integration across various perceptual domains.
  • Ten Oever, S., Kaushik, K., & Martin, A. E. (2022). Inferring the nature of linguistic computations in the brain. PLoS Computational Biology, 18(7): e1010269. doi:10.1371/journal.pcbi.1010269.

    Abstract

    Sentences contain structure that determines their meaning beyond that of individual words. An influential study by Ding and colleagues (2016) used frequency tagging of phrases and sentences to show that the human brain is sensitive to structure by finding peaks of neural power at the rate at which structures were presented. Since then, there has been a rich debate on how to best explain this pattern of results with profound impact on the language sciences. Models that use hierarchical structure building, as well as models based on associative sequence processing, can predict the neural response, creating an inferential impasse as to which class of models explains the nature of the linguistic computations reflected in the neural readout. In the current manuscript, we discuss pitfalls and common fallacies seen in the conclusions drawn in the literature illustrated by various simulations. We conclude that inferring the neural operations of sentence processing based on these neural data, and any like it, alone, is insufficient. We discuss how to best evaluate models and how to approach the modeling of neural readouts to sentence processing in a manner that remains faithful to cognitive, neural, and linguistic principles.
  • Van der Werf, O. J., Ten Oever, S., Schuhmann, T., & Sack, A. T. (2022). No evidence of rhythmic visuospatial attention at cued locations in a spatial cuing paradigm, regardless of their behavioural relevance. European Journal of Neuroscience, 55(11-12), 3100-3116. doi:10.1111/ejn.15353.

    Abstract

    Recent evidence suggests that visuospatial attentional performance is not stable over time but fluctuates in a rhythmic fashion. These attentional rhythms allow for sampling of different visuospatial locations in each cycle of this rhythm. However, it is still unclear in which paradigmatic circumstances rhythmic attention becomes evident. First, it is unclear at what spatial locations rhythmic attention occurs. Second, it is unclear how the behavioural relevance of each spatial location determines the rhythmic sampling patterns. Here, we aim to elucidate these two issues. Firstly, we aim to find evidence of rhythmic attention at the predicted (i.e. cued) location under moderately informative predictor value, replicating earlier studies. Secondly, we hypothesise that rhythmic attentional sampling behaviour will be affected by the behavioural relevance of the sampled location, ranging from non-informative to fully informative. To these aims, we used a modified Egly-Driver task with three conditions: a fully informative cue, a moderately informative cue (replication condition), and a non-informative cue. We did not find evidence of rhythmic sampling at cued locations, failing to replicate earlier studies. Nor did we find differences in rhythmic sampling under different predictive values of the cue. The current data does not allow for robust conclusions regarding the non-cued locations due to the absence of a priori hypotheses. Post-hoc explorative data analyses, however, clearly indicate that attention samples non-cued locations in a theta-rhythmic manner, specifically when the cued location bears higher behavioural relevance than the non-cued locations.
  • Woensdregt, M., Jara-Ettinger, J., & Rubio-Fernandez, P. (2022). Language universals rely on social cognition: Computational models of the use of this and that to redirect the receiver’s attention. In J. Culbertson, A. Perfors, H. Rabagliati, & V. Ramenzoni (Eds.), Proceedings of the 44th Annual Conference of the Cognitive Science Society (CogSci 2022) (pp. 1382-1388). Toronto, Canada: Cognitive Science Society.

    Abstract

    Demonstratives—simple referential devices like this and that—are linguistic universals, but their meaning varies cross-linguistically. In languages like English and Italian, demonstratives are thought to encode the referent’s distance from the producer (e.g., that one means “the one far away from me”),
    while in others, like Portuguese and Spanish, they encode relative distance from both producer and receiver (e.g., aquel means “the one far away from both of us”). Here we propose that demonstratives are also sensitive to the receiver’s focus of attention, hence requiring a deeper form of social cognition
    than previously thought. We provide initial empirical and computational evidence for this idea, suggesting that producers use
    demonstratives to redirect the receiver’s attention towards the intended referent, rather than only to indicate its physical distance.

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