Peter Hagoort

Publications

Displaying 1 - 44 of 44
  • 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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  • Dai, B., McQueen, J. M., Terporten, R., Hagoort, P., & Kösem, A. (2022). Distracting Linguistic Information Impairs Neural Tracking of Attended Speech. Current Research in Neurobiology, 3: 100043. doi:10.1016/j.crneur.2022.100043.

    Abstract

    Listening to speech is difficult in noisy environments, and is even harder when the interfering noise consists of intelligible speech as compared to unintelligible sounds. This suggests that the competing linguistic information interferes with the neural processing of target speech. Interference could either arise from a degradation of the neural representation of the target speech, or from increased representation of distracting speech that enters in competition with the target speech. We tested these alternative hypotheses using magnetoencephalography (MEG) while participants listened to a target clear speech in the presence of distracting noise-vocoded speech. Crucially, the distractors were initially unintelligible but became more intelligible after a short training session. Results showed that the comprehension of the target speech was poorer after training than before training. The neural tracking of target speech in the delta range (1–4 Hz) reduced in strength in the presence of a more intelligible distractor. In contrast, the neural tracking of distracting signals was not significantly modulated by intelligibility. These results suggest that the presence of distracting speech signals degrades the linguistic representation of target speech carried by delta oscillations.
  • Giglio, L., Ostarek, M., Weber, K., & Hagoort, P. (2022). Commonalities and asymmetries in the neurobiological infrastructure for language production and comprehension. Cerebral Cortex, 32(7), 1405-1418. doi:10.1093/cercor/bhab287.

    Abstract

    The neurobiology of sentence production has been largely understudied compared to the neurobiology of sentence comprehension, due to difficulties with experimental control and motion-related artifacts in neuroimaging. We studied the neural response to constituents of increasing size and specifically focused on the similarities and differences in the production and comprehension of the same stimuli. Participants had to either produce or listen to stimuli in a gradient of constituent size based on a visual prompt. Larger constituent sizes engaged the left inferior frontal gyrus (LIFG) and middle temporal gyrus (LMTG) extending to inferior parietal areas in both production and comprehension, confirming that the neural resources for syntactic encoding and decoding are largely overlapping. An ROI analysis in LIFG and LMTG also showed that production elicited larger responses to constituent size than comprehension and that the LMTG was more engaged in comprehension than production, while the LIFG was more engaged in production than comprehension. Finally, increasing constituent size was characterized by later BOLD peaks in comprehension but earlier peaks in production. These results show that syntactic encoding and parsing engage overlapping areas, but there are asymmetries in the engagement of the language network due to the specific requirements of production and comprehension.

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  • Hagoort, P. (2022). Reasoning and the brain. In M. Stokhof, & K. Stenning (Eds.), Rules, regularities, randomness. Festschrift for Michiel van Lambalgen (pp. 83-85). Amsterdam: Institute for Logic, Language and Computation.
  • Heilbron, M., Armeni, K., Schoffelen, J.-M., Hagoort, P., & De Lange, F. P. (2022). A hierarchy of linguistic predictions during natural language comprehension. Proceedings of the National Academy of Sciences of the United States of America, 119(32): e2201968119. doi:10.1073/pnas.2201968119.

    Abstract

    Understanding spoken language requires transforming ambiguous acoustic streams into a hierarchy of representations, from phonemes to meaning. It has been suggested that the brain uses prediction to guide the interpretation of incoming input. However, the role of prediction in language processing remains disputed, with disagreement about both the ubiquity and representational nature of predictions. Here, we address both issues by analyzing brain recordings of participants listening to audiobooks, and using a deep neural network (GPT-2) to precisely quantify contextual predictions. First, we establish that brain responses to words are modulated by ubiquitous predictions. Next, we disentangle model-based predictions into distinct dimensions, revealing dissociable neural signatures of predictions about syntactic category (parts of speech), phonemes, and semantics. Finally, we show that high-level (word) predictions inform low-level (phoneme) predictions, supporting hierarchical predictive processing. Together, these results underscore the ubiquity of prediction in language processing, showing that the brain spontaneously predicts upcoming language at multiple levels of abstraction.

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  • Hoeksema, N., Hagoort, P., & Vernes, S. C. (2022). Piecing together the building blocks of the vocal learning bat brain. 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. 294-296). Nijmegen: Joint Conference on Language Evolution (JCoLE).
  • Huizeling, E., Arana, S., Hagoort, P., & Schoffelen, J.-M. (2022). Lexical frequency and sentence context influence the brain’s response to single words. Neurobiology of Language, 3(1), 149-179. doi:10.1162/nol_a_00054.

    Abstract

    Typical adults read remarkably quickly. Such fast reading is facilitated by brain processes that are sensitive to both word frequency and contextual constraints. It is debated as to whether these attributes have additive or interactive effects on language processing in the brain. We investigated this issue by analysing existing magnetoencephalography data from 99 participants reading intact and scrambled sentences. Using a cross-validated model comparison scheme, we found that lexical frequency predicted the word-by-word elicited MEG signal in a widespread cortical network, irrespective of sentential context. In contrast, index (ordinal word position) was more strongly encoded in sentence words, in left front-temporal areas. This confirms that frequency influences word processing independently of predictability, and that contextual constraints affect word-by-word brain responses. With a conservative multiple comparisons correction, only the interaction between lexical frequency and surprisal survived, in anterior temporal and frontal cortex, and not between lexical frequency and entropy, nor between lexical frequency and index. However, interestingly, the uncorrected index*frequency interaction revealed an effect in left frontal and temporal cortex that reversed in time and space for intact compared to scrambled sentences. Finally, we provide evidence to suggest that, in sentences, lexical frequency and predictability may independently influence early (<150ms) and late stages of word processing, but interact during later stages of word processing (>150-250ms), thus helping to converge previous contradictory eye-tracking and electrophysiological literature. Current neuro-cognitive models of reading would benefit from accounting for these differing effects of lexical frequency and predictability on different stages of word processing.
  • Huizeling, E., Peeters, D., & Hagoort, P. (2022). Prediction of upcoming speech under fluent and disfluent conditions: Eye tracking evidence from immersive virtual reality. Language, Cognition and Neuroscience, 37(4), 481-508. doi:10.1080/23273798.2021.1994621.

    Abstract

    Traditional experiments indicate that prediction is important for efficient speech processing. In three virtual reality visual world paradigm experiments, we tested whether such findings hold in naturalistic settings (Experiment 1) and provided novel insights into whether disfluencies in speech (repairs/hesitations) inform one’s predictions in rich environments (Experiments 2–3). Experiment 1 supports that listeners predict upcoming speech in naturalistic environments, with higher proportions of anticipatory target fixations in predictable compared to unpredictable trials. In Experiments 2–3, disfluencies reduced anticipatory fixations towards predicted referents, compared to conjunction (Experiment 2) and fluent (Experiment 3) sentences. Unexpectedly, Experiment 2 provided no evidence that participants made new predictions from a repaired verb. Experiment 3 provided novel findings that fixations towards the speaker increase upon hearing a hesitation, supporting current theories of how hesitations influence sentence processing. Together, these findings unpack listeners’ use of visual (objects/speaker) and auditory (speech/disfluencies) information when predicting upcoming words.
  • Lai, V. T., Van Berkum, J. J. A., & Hagoort, P. (2022). Negative affect increases reanalysis of conflicts between discourse context and world knowledge. Frontiers in Communication, 7: 910482. doi:10.3389/fcomm.2022.910482.

    Abstract

    Introduction: Mood is a constant in our daily life and can permeate all levels of cognition. We examined whether and how mood influences the processing of discourse content that is relatively neutral and not loaded with emotion. During discourse processing, readers have to constantly strike a balance between what they know in long term memory and what the current discourse is about. Our general hypothesis is that mood states would affect this balance. We hypothesized that readers in a positive mood would rely more on default world knowledge, whereas readers in a negative mood would be more inclined to analyze the details in the current discourse.

    Methods: Participants were put in a positive and a negative mood via film clips, one week apart. In each session, after mood manipulation, they were presented with sentences in discourse materials. We created sentences such as “With the lights on you can see...” that end with critical words (CWs) “more” or “less”, where general knowledge supports “more”, not “less”. We then embedded each of these sentences in a wider discourse that does/does not support the CWs (a story about driving in the night vs. stargazing). EEG was recorded throughout.

    Results: The results showed that first, mood manipulation was successful in that there was a significant mood difference between sessions. Second, mood did not modulate the N400 effects. Participants in both moods detected outright semantic violations and allowed world knowledge to be overridden by discourse context. Third, mood modulated the LPC (Late Positive Component) effects, distributed in the frontal region. In negative moods, the LPC was sensitive to one-level violation. That is, CWs that were supported by only world knowledge, only discourse, and neither, elicited larger frontal LPCs, in comparison to the condition where CWs were supported by both world knowledge and discourse.

    Discussion: These results suggest that mood does not influence all processes involved in discourse processing. Specifically, mood does not influence lexical-semantic retrieval (N400), but it does influence elaborative processes for sensemaking (P600) during discourse processing. These results advance our understanding of the impact and time course of mood on discourse.

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  • Murphy, E., Woolnough, O., Rollo, P. S., Roccaforte, Z., Segaert, K., Hagoort, P., & Tandon, N. (2022). Minimal phrase composition revealed by intracranial recordings. The Journal of Neuroscience, 42(15), 3216-3227. doi:10.1523/JNEUROSCI.1575-21.2022.

    Abstract

    The ability to comprehend phrases is an essential integrative property of the brain. Here we evaluate the neural processes that enable the transition from single word processing to a minimal compositional scheme. Previous research has reported conflicting timing effects of composition, and disagreement persists with respect to inferior frontal and posterior temporal contributions. To address these issues, 19 patients (10 male, 19 female) implanted with penetrating depth or surface subdural intracranial electrodes heard auditory recordings of adjective-noun, pseudoword-noun and adjective-pseudoword phrases and judged whether the phrase matched a picture. Stimulus-dependent alterations in broadband gamma activity, low frequency power and phase-locking values across the language-dominant left hemisphere were derived. This revealed a mosaic located on the lower bank of the posterior superior temporal sulcus (pSTS), in which closely neighboring cortical sites displayed exclusive sensitivity to either lexicality or phrase structure, but not both. Distinct timings were found for effects of phrase composition (210–300 ms) and pseudoword processing (approximately 300–700 ms), and these were localized to neighboring electrodes in pSTS. The pars triangularis and temporal pole encoded anticipation of composition in broadband low frequencies, and both regions exhibited greater functional connectivity with pSTS during phrase composition. Our results suggest that the pSTS is a highly specialized region comprised of sparsely interwoven heterogeneous constituents that encodes both lower and higher level linguistic features. This hub in pSTS for minimal phrase processing may form the neural basis for the human-specific computational capacity for forming hierarchically organized linguistic structures.
  • Udden, J., Hulten, A., Schoffelen, J.-M., Lam, N. H. L., Harbusch, K., Van den Bosch, A., Kempen, G., Petersson, K. M., & Hagoort, P. (2022). Supramodal sentence processing in the human brain: fMRI evidence for the influence of syntactic complexity in more than 200 participants. Neurobiology of Language, 3(4), 575-598. doi:10.1162/nol_a_00076.

    Abstract

    This study investigated two questions. One is: To what degree is sentence processing beyond single words independent of the input modality (speech vs. reading)? The second question is: Which parts of the network recruited by both modalities is sensitive to syntactic complexity? These questions were investigated by having more than 200 participants read or listen to well-formed sentences or series of unconnected words. A largely left-hemisphere frontotemporoparietal network was found to be supramodal in nature, i.e., independent of input modality. In addition, the left inferior frontal gyrus (LIFG) and the left posterior middle temporal gyrus (LpMTG) were most clearly associated with left-branching complexity. The left anterior temporal lobe (LaTL) showed the greatest sensitivity to sentences that differed in right-branching complexity. Moreover, activity in LIFG and LpMTG increased from sentence onset to end, in parallel with an increase of the left-branching complexity. While LIFG, bilateral anterior temporal lobe, posterior MTG, and left inferior parietal lobe (LIPL) all contribute to the supramodal unification processes, the results suggest that these regions differ in their respective contributions to syntactic complexity related processing. The consequences of these findings for neurobiological models of language processing are discussed.

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  • Vernes, S. C., Devanna, P., Hörpel, S. G., Alvarez van Tussenbroek, I., Firzlaff, U., Hagoort, P., Hiller, M., Hoeksema, N., Hughes, G. M., Lavrichenko, K., Mengede, J., Morales, A. E., & Wiesmann, M. (2022). The pale spear‐nosed bat: A neuromolecular and transgenic model for vocal learning. Annals of the New York Academy of Sciences, 1517, 125-142. doi:10.1111/nyas.14884.

    Abstract

    Vocal learning, the ability to produce modified vocalizations via learning from acoustic signals, is a key trait in the evolution of speech. While extensively studied in songbirds, mammalian models for vocal learning are rare. Bats present a promising study system given their gregarious natures, small size, and the ability of some species to be maintained in captive colonies. We utilize the pale spear-nosed bat (Phyllostomus discolor) and report advances in establishing this species as a tractable model for understanding vocal learning. We have taken an interdisciplinary approach, aiming to provide an integrated understanding across genomics (Part I), neurobiology (Part II), and transgenics (Part III). In Part I, we generated new, high-quality genome annotations of coding genes and noncoding microRNAs to facilitate functional and evolutionary studies. In Part II, we traced connections between auditory-related brain regions and reported neuroimaging to explore the structure of the brain and gene expression patterns to highlight brain regions. In Part III, we created the first successful transgenic bats by manipulating the expression of FoxP2, a speech-related gene. These interdisciplinary approaches are facilitating a mechanistic and evolutionary understanding of mammalian vocal learning and can also contribute to other areas of investigation that utilize P. discolor or bats as study species.

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  • Eichert, N., Peeters, D., & Hagoort, P. (2018). Language-driven anticipatory eye movements in virtual reality. Behavior Research Methods, 50(3), 1102-1115. doi:10.3758/s13428-017-0929-z.

    Abstract

    Predictive language processing is often studied by measuring eye movements as participants look at objects on a computer screen while they listen to spoken sentences. The use of this variant of the visual world paradigm has shown that information encountered by a listener at a spoken verb can give rise to anticipatory eye movements to a target object, which is taken to indicate that people predict upcoming words. The ecological validity of such findings remains questionable, however, because these computer experiments used two-dimensional (2D) stimuli that are mere abstractions of real world objects. Here we present a visual world paradigm study in a three-dimensional (3D) immersive virtual reality environment. Despite significant changes in the stimulus material and the different mode of stimulus presentation, language-mediated anticipatory eye movements were observed. These findings thus indicate prediction of upcoming words in language comprehension in a more naturalistic setting where natural depth cues are preserved. Moreover, the results confirm the feasibility of using eye-tracking in rich and multimodal 3D virtual environments.

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    13428_2017_929_MOESM1_ESM.docx
  • Franken, M. K., Acheson, D. J., McQueen, J. M., Hagoort, P., & Eisner, F. (2018). Opposing and following responses in sensorimotor speech control: Why responses go both ways. Psychonomic Bulletin & Review, 25(4), 1458-1467. doi:10.3758/s13423-018-1494-x.

    Abstract

    When talking, speakers continuously monitor and use the auditory feedback of their own voice to control and inform speech production processes. When speakers are provided with auditory feedback that is perturbed in real time, most of them compensate for this by opposing the feedback perturbation. But some speakers follow the perturbation. In the current study, we investigated whether the state of the speech production system at perturbation onset may determine what type of response (opposing or following) is given. The results suggest that whether a perturbation-related response is opposing or following depends on ongoing fluctuations of the production system: It initially responds by doing the opposite of what it was doing. This effect and the non-trivial proportion of following responses suggest that current production models are inadequate: They need to account for why responses to unexpected sensory feedback depend on the production-system’s state at the time of perturbation.
  • Franken, M. K., Eisner, F., Acheson, D. J., McQueen, J. M., Hagoort, P., & Schoffelen, J.-M. (2018). Self-monitoring in the cerebral cortex: Neural responses to pitch-perturbed auditory feedback during speech production. NeuroImage, 179, 326-336. doi:10.1016/j.neuroimage.2018.06.061.

    Abstract

    Speaking is a complex motor skill which requires near instantaneous integration of sensory and motor-related information. Current theory hypothesizes a complex interplay between motor and auditory processes during speech production, involving the online comparison of the speech output with an internally generated forward model. To examine the neural correlates of this intricate interplay between sensory and motor processes, the current study uses altered auditory feedback (AAF) in combination with magnetoencephalography (MEG). Participants vocalized the vowel/e/and heard auditory feedback that was temporarily pitch-shifted by only 25 cents, while neural activity was recorded with MEG. As a control condition, participants also heard the recordings of the same auditory feedback that they heard in the first half of the experiment, now without vocalizing. The participants were not aware of any perturbation of the auditory feedback. We found auditory cortical areas responded more strongly to the pitch shifts during vocalization. In addition, auditory feedback perturbation resulted in spectral power increases in the θ and lower β bands, predominantly in sensorimotor areas. These results are in line with current models of speech production, suggesting auditory cortical areas are involved in an active comparison between a forward model's prediction and the actual sensory input. Subsequently, these areas interact with motor areas to generate a motor response. Furthermore, the results suggest that θ and β power increases support auditory-motor interaction, motor error detection and/or sensory prediction processing.
  • De Groot, A. M. B., & Hagoort, P. (Eds.). (2018). Research methods in psycholinguistics and the neurobiology of language: A practical guide. Oxford: Wiley.
  • Hagoort, P. (2018). Prerequisites for an evolutionary stance on the neurobiology of language. Current Opinion in Behavioral Sciences, 21, 191-194. doi:10.1016/j.cobeha.2018.05.012.
  • Heyselaar, E., Mazaheri, A., Hagoort, P., & Segaert, K. (2018). Changes in alpha activity reveal that social opinion modulates attention allocation during face processing. NeuroImage, 174, 432-440. doi:10.1016/j.neuroimage.2018.03.034.

    Abstract

    Participants’ performance differs when conducting a task in the presence of a secondary individual, moreover the opinion the participant has of this individual also plays a role. Using EEG, we investigated how previous interactions with, and evaluations of, an avatar in virtual reality subsequently influenced attentional allocation to the face of that avatar. We focused on changes in the alpha activity as an index of attentional allocation. We found that the onset of an avatar’s face whom the participant had developed a rapport with induced greater alpha suppression. This suggests greater attentional resources are allocated to the interacted-with avatars. The evaluative ratings of the avatar induced a U-shaped change in alpha suppression, such that participants paid most attention when the avatar was rated as average. These results suggest that attentional allocation is an important element of how behaviour is altered in the presence of a secondary individual and is modulated by our opinion of that individual.

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  • Kösem, A., Bosker, H. R., Takashima, A., Meyer, A. S., Jensen, O., & Hagoort, P. (2018). Neural entrainment determines the words we hear. Current Biology, 28, 2867-2875. doi:10.1016/j.cub.2018.07.023.

    Abstract

    Low-frequency neural entrainment to rhythmic input
    has been hypothesized as a canonical mechanism
    that shapes sensory perception in time. Neural
    entrainment is deemed particularly relevant for
    speech analysis, as it would contribute to the extraction
    of discrete linguistic elements from continuous
    acoustic signals. However, its causal influence in
    speech perception has been difficult to establish.
    Here, we provide evidence that oscillations build temporal
    predictions about the duration of speech tokens
    that affect perception. Using magnetoencephalography
    (MEG), we studied neural dynamics during
    listening to sentences that changed in speech rate.
    Weobserved neural entrainment to preceding speech
    rhythms persisting for several cycles after the change
    in rate. The sustained entrainment was associated
    with changes in the perceived duration of the last
    word’s vowel, resulting in the perception of words
    with different meanings. These findings support oscillatory
    models of speech processing, suggesting that
    neural oscillations actively shape speech perception.
  • Lam, N. H. L., Hulten, A., Hagoort, P., & Schoffelen, J.-M. (2018). Robust neuronal oscillatory entrainment to speech displays individual variation in lateralisation. Language, Cognition and Neuroscience, 33(8), 943-954. doi:10.1080/23273798.2018.1437456.

    Abstract

    Neural oscillations may be instrumental for the tracking and segmentation of continuous speech. Earlier work has suggested that delta, theta and gamma oscillations entrain to the speech rhythm. We used magnetoencephalography and a large sample of 102 participants to investigate oscillatory entrainment to speech, and observed robust entrainment of delta and theta activity, and weak group-level gamma entrainment. We show that the peak frequency and the hemispheric lateralisation of the entrainment are subject to considerable individual variability. The first finding may support the involvement of intrinsic oscillations in entrainment, and the second finding suggests that there is no systematic default right-hemispheric bias for processing acoustic signals on a slow time scale. Although low frequency entrainment to speech is a robust phenomenon, the characteristics of entrainment vary across individuals, and this variation is important for understanding the underlying neural mechanisms of entrainment, as well as its functional significance.
  • Segaert, K., Mazaheri, A., & Hagoort, P. (2018). Binding language: Structuring sentences through precisely timed oscillatory mechanisms. European Journal of Neuroscience, 48(7), 2651-2662. doi:10.1111/ejn.13816.

    Abstract

    Syntactic binding refers to combining words into larger structures. Using EEG, we investigated the neural processes involved in syntactic binding. Participants were auditorily presented two-word sentences (i.e. pronoun and pseudoverb such as ‘I grush’, ‘she grushes’, for which syntactic binding can take place) and wordlists (i.e. two pseudoverbs such as ‘pob grush’, ‘pob grushes’, for which no binding occurs). Comparing these two conditions, we targeted syntactic binding while minimizing contributions of semantic binding and of other cognitive processes such as working memory. We found a converging pattern of results using two distinct analysis approaches: one approach using frequency bands as defined in previous literature, and one data-driven approach in which we looked at the entire range of frequencies between 3-30 Hz without the constraints of pre-defined frequency bands. In the syntactic binding (relative to the wordlist) condition, a power increase was observed in the alpha and beta frequency range shortly preceding the presentation of the target word that requires binding, which was maximal over frontal-central electrodes. Our interpretation is that these signatures reflect that language comprehenders expect the need for binding to occur. Following the presentation of the target word in a syntactic binding context (relative to the wordlist condition), an increase in alpha power maximal over a left lateralized cluster of frontal-temporal electrodes was observed. We suggest that this alpha increase relates to syntactic binding taking place. Taken together, our findings suggest that increases in alpha and beta power are reflections of distinct the neural processes underlying syntactic binding.
  • Tromp, J., Peeters, D., Meyer, A. S., & Hagoort, P. (2018). The combined use of Virtual Reality and EEG to study language processing in naturalistic environments. Behavior Research Methods, 50(2), 862-869. doi:10.3758/s13428-017-0911-9.

    Abstract

    When we comprehend language, we often do this in rich settings in which we can use many cues to understand what someone is saying. However, it has traditionally been difficult to design experiments with rich three-dimensional contexts that resemble our everyday environments, while maintaining control over the linguistic and non-linguistic information that is available. Here we test the validity of combining electroencephalography (EEG) and Virtual Reality (VR) to overcome this problem. We recorded electrophysiological brain activity during language processing in a well-controlled three-dimensional virtual audiovisual environment. Participants were immersed in a virtual restaurant, while wearing EEG equipment. In the restaurant participants encountered virtual restaurant guests. Each guest was seated at a separate table with an object on it (e.g. a plate with salmon). The restaurant guest would then produce a sentence (e.g. “I just ordered this salmon.”). The noun in the spoken sentence could either match (“salmon”) or mismatch (“pasta”) with the object on the table, creating a situation in which the auditory information was either appropriate or inappropriate in the visual context. We observed a reliable N400 effect as a consequence of the mismatch. This finding validates the combined use of VR and EEG as a tool to study the neurophysiological mechanisms of everyday language comprehension in rich, ecologically valid settings.
  • Vanlangendonck, F., Takashima, A., Willems, R. M., & Hagoort, P. (2018). Distinguishable memory retrieval networks for collaboratively and non-collaboratively learned information. Neuropsychologia, 111, 123-132. doi:10.1016/j.neuropsychologia.2017.12.008.

    Abstract

    Learning often occurs in communicative and collaborative settings, yet almost all research into the neural basis of memory relies on participants encoding and retrieving information on their own. We investigated whether learning linguistic labels in a collaborative context at least partly relies on cognitively and neurally distinct representations, as compared to learning in an individual context. Healthy human participants learned labels for sets of abstract shapes in three different tasks. They came up with labels with another person in a collaborative communication task (collaborative condition), by themselves (individual condition), or were given pre-determined unrelated labels to learn by themselves (arbitrary condition). Immediately after learning, participants retrieved and produced the labels aloud during a communicative task in the MRI scanner. The fMRI results show that the retrieval of collaboratively generated labels as compared to individually learned labels engages brain regions involved in understanding others (mentalizing or theory of mind) and autobiographical memory, including the medial prefrontal cortex, the right temporoparietal junction and the precuneus. This study is the first to show that collaboration during encoding affects the neural networks involved in retrieval.
  • Vanlangendonck, F., Willems, R. M., & Hagoort, P. (2018). Taking common ground into account: Specifying the role of the mentalizing network in communicative language production. PLoS One, 13(10): e0202943. doi:10.1371/journal.pone.0202943.
  • Wang, L., Hagoort, P., & Jensen, O. (2018). Language prediction is reflected by coupling between frontal gamma and posterior alpha oscillations. Journal of Cognitive Neuroscience, 30(3), 432-447. doi:10.1162/jocn_a_01190.

    Abstract

    Readers and listeners actively predict upcoming words during language processing. These predictions might serve to support the unification of incoming words into sentence context and thus rely on interactions between areas in the language network. In the current magnetoencephalography study, participants read sentences that varied in contextual constraints so that the predictability of the sentence-final words was either high or low. Before the sentence-final words, we observed stronger alpha power suppression for the highly compared with low constraining sentences in the left inferior frontal cortex, left posterior temporal region, and visual word form area. Importantly, the temporal and visual word form area alpha power correlated negatively with left frontal gamma power for the highly constraining sentences. We suggest that the correlation between alpha power decrease in temporal language areas and left prefrontal gamma power reflects the initiation of an anticipatory unification process in the language network.
  • Wang, L., Hagoort, P., & Jensen, O. (2018). Gamma oscillatory activity related to language prediction. Journal of Cognitive Neuroscience, 30(8), 1075-1085. doi:10.1162/jocn_a_01275.

    Abstract

    Using magnetoencephalography, the current study examined gamma activity associated with language prediction. Participants read high- and low-constraining sentences in which the final word of the sentence was either expected or unexpected. Although no consistent gamma power difference induced by the sentence-final words was found between the expected and unexpected conditions, the correlation of gamma power during the prediction and activation intervals of the sentence-final words was larger when the presented words matched with the prediction compared with when the prediction was violated or when no prediction was available. This suggests that gamma magnitude relates to the match between predicted and perceived words. Moreover, the expected words induced activity with a slower gamma frequency compared with that induced by unexpected words. Overall, the current study establishes that prediction is related to gamma power correlations and a slowing of the gamma frequency.
  • Baggio, G., & Hagoort, P. (2011). The balance between memory and unification in semantics: A dynamic account of the N400. Language and Cognitive Processes, 26, 1338-1367. doi:10.1080/01690965.2010.542671.

    Abstract

    At least three cognitive brain components are necessary in order for us to be able to produce and comprehend language: a Memory repository for the lexicon, a Unification buffer where lexical information is combined into novel structures, and a Control apparatus presiding over executive function in language. Here we describe the brain networks that support Memory and Unification in semantics. A dynamic account of their interactions is presented, in which a balance between the two components is sought at each word-processing step. We use the theory to provide an explanation of the N400 effect.
  • Davids, N., Segers, E., Van den Brink, D., Mitterer, H., van Balkom, H., Hagoort, P., & Verhoeven, L. (2011). The nature of auditory discrimination problems in children with specific language impairment: An MMN study. Neuropsychologia, 49, 19-28. doi:10.1016/j.neuropsychologia.2010.11.001.

    Abstract

    Many children with Specific Language Impairment (SLI) show impairments in discriminating auditorily presented stimuli. The present study investigates whether these discrimination problems are speech specific or of a general auditory nature. This was studied by using a linguistic and nonlinguistic contrast that were matched for acoustic complexity in an active behavioral task and a passive ERP paradigm, known to elicit the mismatch negativity (MMN). In addition, attention skills and a variety of language skills were measured. Participants were 25 five-year-old Dutch children with SLI having receptive as well as productive language problems and 25 control children with typical speechand language development. At the behavioral level, the SLI group was impaired in discriminating the linguistic contrast as compared to the control group, while both groups were unable to distinguish the non-linguistic contrast. Moreover, the SLI group tended to have impaired attention skills which correlated with performance on most of the language tests. At the neural level, the SLI group, in contrast to the control group, did not show an MMN in response to either the linguistic or nonlinguistic contrast. The MMN data are consistent with an account that relates the symptoms in children with SLI to non-speech processing difficulties.
  • Folia, V., Forkstam, C., Ingvar, M., Hagoort, P., & Petersson, K. M. (2011). Implicit artificial syntax processing: Genes, preference, and bounded recursion. Biolinguistics, 5(1/2), 105-132.

    Abstract

    The first objective of this study was to compare the brain network engaged by preference classification and the standard grammaticality classification after implicit artificial syntax acquisition by re-analyzing previously reported event-related fMRI data. The results show that preference and grammaticality classification engage virtually identical brain networks, including Broca’s region, consistent with previous behavioral findings. Moreover, the results showed that the effects related to artificial syntax in Broca’s region were essentially the same when masked with variability related to natural syntax processing in the same participants. The second objective was to explore CNTNAP2-related effects in implicit artificial syntax learning by analyzing behavioral and event-related fMRI data from a subsample. The CNTNAP2 gene has been linked to specific language impairment and is controlled by the FOXP2 transcription factor. CNTNAP2 is expressed in language related brain networks in the developing human brain and the FOXP2–CNTNAP2 pathway provides a mechanistic link between clinically distinct syndromes involving disrupted language. Finally, we discuss the implication of taking natural language to be a neurobiological system in terms of bounded recursion and suggest that the left inferior frontal region is a generic on-line sequence processor that unifies information from various sources in an incremental and recursive manner.
  • Habets, B., Kita, S., Shao, Z., Ozyurek, A., & Hagoort, P. (2011). The role of synchrony and ambiguity in speech–gesture integration during comprehension. Journal of Cognitive Neuroscience, 23, 1845-1854. doi:10.1162/jocn.2010.21462.

    Abstract

    During face-to-face communication, one does not only hear speech but also see a speaker's communicative hand movements. It has been shown that such hand gestures play an important role in communication where the two modalities influence each other's interpretation. A gesture typically temporally overlaps with coexpressive speech, but the gesture is often initiated before (but not after) the coexpressive speech. The present ERP study investigated what degree of asynchrony in the speech and gesture onsets are optimal for semantic integration of the concurrent gesture and speech. Videos of a person gesturing were combined with speech segments that were either semantically congruent or incongruent with the gesture. Although gesture and speech always overlapped in time, gesture and speech were presented with three different degrees of asynchrony. In the SOA 0 condition, the gesture onset and the speech onset were simultaneous. In the SOA 160 and 360 conditions, speech was delayed by 160 and 360 msec, respectively. ERPs time locked to speech onset showed a significant difference between semantically congruent versus incongruent gesture–speech combinations on the N400 for the SOA 0 and 160 conditions. No significant difference was found for the SOA 360 condition. These results imply that speech and gesture are integrated most efficiently when the differences in onsets do not exceed a certain time span because of the fact that iconic gestures need speech to be disambiguated in a way relevant to the speech context.
  • Hagoort, P. (2011). The binding problem for language, and its consequences for the neurocognition of comprehension. In E. A. Gibson, & N. J. Pearlmutter (Eds.), The processing and acquisition of reference (pp. 403-436). Cambridge, MA: MIT Press.
  • Hagoort, P. (2011). The neuronal infrastructure for unification at multiple levels. In G. Gaskell, & P. Zwitserlood (Eds.), Lexical representation: A multidisciplinary approach (pp. 231-242). Berlin: De Gruyter Mouton.
  • Lai, V. T., Hagoort, P., & Casasanto, D. (2011). Affective and non-affective meaning in words and pictures. In L. Carlson, C. Holscher, & T. Shipley (Eds.), Proceedings of the 33rd Annual Meeting of the Cognitive Science Society (pp. 390-395). Austin, TX: Cognitive Science Society.
  • Menenti, L., Gierhan, S., Segaert, K., & Hagoort, P. (2011). Shared language: Overlap and segregation of the neuronal infrastructure for speaking and listening revealed by functional MRI. Psychological Science, 22, 1173-1182. doi:10.1177/0956797611418347.

    Abstract

    Whether the brain’s speech-production system is also involved in speech comprehension is a topic of much debate. Research has focused on whether motor areas are involved in listening, but overlap between speaking and listening might occur not only at primary sensory and motor levels, but also at linguistic levels (where semantic, lexical, and syntactic processes occur). Using functional MRI adaptation during speech comprehension and production, we found that the brain areas involved in semantic, lexical, and syntactic processing are mostly the same for speaking and for listening. Effects of primary processing load (indicative of sensory and motor processes) overlapped in auditory cortex and left inferior frontal cortex, but not in motor cortex, where processing load affected activity only in speaking. These results indicate that the linguistic parts of the language system are used for both speaking and listening, but that the motor system does not seem to provide a crucial contribution to listening.
  • Pijnacker, J., Geurts, B., Van Lambalgen, M., Buitelaar, J., & Hagoort, P. (2011). Reasoning with exceptions: An event-related brain potentials study. Journal of Cognitive Neuroscience, 23, 471-480. doi:10.1162/jocn.2009.21360.

    Abstract

    Defeasible inferences are inferences that can be revised in the light of new information. Although defeasible inferences are pervasive in everyday communication, little is known about how and when they are processed by the brain. This study examined the electrophysiological signature of defeasible reasoning using a modified version of the suppression task. Participants were presented with conditional inferences (of the type “if p, then q; p, therefore q”) that were preceded by a congruent or a disabling context. The disabling context contained a possible exception or precondition that prevented people from drawing the conclusion. Acceptability of the conclusion was indeed lower in the disabling condition compared to the congruent condition. Further, we found a large sustained negativity at the conclusion of the disabling condition relative to the congruent condition, which started around 250 msec and was persistent throughout the entire epoch. Possible accounts for the observed effect are discussed.
  • Scheeringa, R., Fries, P., Petersson, K. M., Oostenveld, R., Grothe, I., Norris, D. G., Hagoort, P., & Bastiaansen, M. C. M. (2011). Neuronal dynamics underlying high- and low- frequency EEG oscillations contribute independently to the human BOLD signal. Neuron, 69, 572-583. doi:10.1016/j.neuron.2010.11.044.

    Abstract

    Work on animals indicates that BOLD is preferentially sensitive to local field potentials, and that it correlates most strongly with gamma band neuronal synchronization. Here we investigate how the BOLD signal in humans performing a cognitive task is related to neuronal synchronization across different frequency bands. We simultaneously recorded EEG and BOLD while subjects engaged in a visual attention task known to induce sustained changes in neuronal synchronization across a wide range of frequencies. Trial-by-trial BOLD luctuations correlated positively with trial-by-trial fluctuations in high-EEG gamma power (60–80 Hz) and negatively with alpha and beta power. Gamma power on the one hand, and alpha and beta power on the other hand, independently contributed to explaining BOLD variance. These results indicate that the BOLD-gamma coupling observed in animals can be extrapolated to humans performing a task and that neuronal dynamics underlying high- and low-frequency synchronization contribute independently to the BOLD signal.

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  • Segaert, K., Menenti, L., Weber, K., & Hagoort, P. (2011). A paradox of syntactic priming: Why response tendencies show priming for passives, and response latencies show priming for actives. PLoS One, 6(10), e24209. doi:10.1371/journal.pone.0024209.

    Abstract

    Speakers tend to repeat syntactic structures across sentences, a phenomenon called syntactic priming. Although it has been suggested that repeating syntactic structures should result in speeded responses, previous research has focused on effects in response tendencies. We investigated syntactic priming effects simultaneously in response tendencies and response latencies for active and passive transitive sentences in a picture description task. In Experiment 1, there were priming effects in response tendencies for passives and in response latencies for actives. However, when participants' pre-existing preference for actives was altered in Experiment 2, syntactic priming occurred for both actives and passives in response tendencies as well as in response latencies. This is the first investigation of the effects of structure frequency on both response tendencies and latencies in syntactic priming. We discuss the implications of these data for current theories of syntactic processing.

    Additional information

    Segaert_2011_Supporting_Info.doc
  • Small, S. L., Hickok, G., Nusbaum, H. C., Blumstein, S., Coslett, H. B., Dell, G., Hagoort, P., Kutas, M., Marantz, A., Pylkkanen, L., Thompson-Schill, S., Watkins, K., & Wise, R. J. (2011). The neurobiology of language: Two years later [Editorial]. Brain and Language, 116(3), 103-104. doi:10.1016/j.bandl.2011.02.004.
  • Tesink, C. M. J. Y., Buitelaar, J. K., Petersson, K. M., Van der Gaag, R. J., Teunisse, J.-P., & Hagoort, P. (2011). Neural correlates of language comprehension in autism spectrum disorders: When language conflicts with world knowledge. Neuropsychologia, 49, 1095-1104. doi:10.1016/j.neuropsychologia.2011.01.018.

    Abstract

    In individuals with ASD, difficulties with language comprehension are most evident when higher-level semantic-pragmatic language processing is required, for instance when context has to be used to interpret the meaning of an utterance. Until now, it is unclear at what level of processing and for what type of context these difficulties in language comprehension occur. Therefore, in the current fMRI study, we investigated the neural correlates of the integration of contextual information during auditory language comprehension in 24 adults with ASD and 24 matched control participants. Different levels of context processing were manipulated by using spoken sentences that were correct or contained either a semantic or world knowledge anomaly. Our findings demonstrated significant differences between the groups in inferior frontal cortex that were only present for sentences with a world knowledge anomaly. Relative to the ASD group, the control group showed significantly increased activation in left inferior frontal gyrus (LIFG) for sentences with a world knowledge anomaly compared to correct sentences. This effect possibly indicates reduced integrative capacities of the ASD group. Furthermore, world knowledge anomalies elicited significantly stronger activation in right inferior frontal gyrus (RIFG) in the control group compared to the ASD group. This additional RIFG activation probably reflects revision of the situation model after new, conflicting information. The lack of recruitment of RIFG is possibly related to difficulties with exception handling in the ASD group.

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  • Van Leeuwen, T. M., Den Ouden, H. E. M., & Hagoort, P. (2011). Effective connectivity determines the nature of subjective experience in grapheme-color synesthesia. Journal of Neuroscience, 31, 9879-9884. doi:10.1523/JNEUROSCI.0569-11.2011.

    Abstract

    Synesthesia provides an elegant model to investigate neural mechanisms underlying individual differences in subjective experience in humans. In grapheme–color synesthesia, written letters induce color sensations, accompanied by activation of color area V4. Competing hypotheses suggest that enhanced V4 activity during synesthesia is either induced by direct bottom-up cross-activation from grapheme processing areas within the fusiform gyrus, or indirectly via higher-order parietal areas. Synesthetes differ in the way synesthetic color is perceived: “projector” synesthetes experience color externally colocalized with a presented grapheme, whereas “associators” report an internally evoked association. Using dynamic causal modeling for fMRI, we show that V4 cross-activation during synesthesia was induced via a bottom-up pathway (within fusiform gyrus) in projector synesthetes, but via a top-down pathway (via parietal lobe) in associators. These findings show how altered coupling within the same network of active regions leads to differences in subjective experience. Our findings reconcile the two most influential cross-activation accounts of synesthesia.
  • Wang, L., Bastiaansen, M. C. M., Yang, Y., & Hagoort, P. (2011). The influence of information structure on the depth of semantic processing: How focus and pitch accent determine the size of the N400 effect. Neuropsychologia, 49, 813-820. doi:10.1016/j.neuropsychologia.2010.12.035.

    Abstract

    To highlight relevant information in dialogues, both wh-question context and pitch accent in answers can be used, such that focused information gains more attention and is processed more elaborately. To evaluate the relative influence of context and pitch accent on the depth of semantic processing, we measured Event-Related Potentials (ERPs) to auditorily presented wh-question-answer pairs. A semantically incongruent word in the answer occurred either in focus or non-focus position as determined by the context, and this word was either accented or unaccented. Semantic incongruency elicited different N400 effects in different conditions. The largest N400 effect was found when the question-marked focus was accented, while the other three conditions elicited smaller N400 effects. The results suggest that context and accentuation interact. Thus accented focused words were processed more deeply compared to conditions where focus and accentuation mismatched, or when the new information had no marking. In addition, there seems to be sex differences in the depth of semantic processing. For males, a significant N400 effect was observed only when the question-marked focus was accented, reduced N400 effects were found in the other dialogues. In contrast, females produced similar N400 effects in all the conditions. These results suggest that regardless of external cues, females tend to engage in more elaborate semantic processing compared to males.
  • Willems, R. M., Clevis, K., & Hagoort, P. (2011). Add a picture for suspense: Neural correlates of the interaction between language and visual information in the perception of fear. Social, Cognitive and Affective Neuroscience, 6, 404-416. doi:10.1093/scan/nsq050.

    Abstract

    We investigated how visual and linguistic information interact in the perception of emotion. We borrowed a phenomenon from film theory which states that presentation of an as such neutral visual scene intensifies the percept of fear or suspense induced by a different channel of information, such as language. Our main aim was to investigate how neutral visual scenes can enhance responses to fearful language content in parts of the brain involved in the perception of emotion. Healthy participants’ brain activity was measured (using functional magnetic resonance imaging) while they read fearful and less fearful sentences presented with or without a neutral visual scene. The main idea is that the visual scenes intensify the fearful content of the language by subtly implying and concretizing what is described in the sentence. Activation levels in the right anterior temporal pole were selectively increased when a neutral visual scene was paired with a fearful sentence, compared to reading the sentence alone, as well as to reading of non-fearful sentences presented with the same neutral scene. We conclude that the right anterior temporal pole serves a binding function of emotional information across domains such as visual and linguistic information.
  • Willems, R. M., Benn, Y., Hagoort, P., Tonia, I., & Varley, R. (2011). Communicating without a functioning language system: Implications for the role of language in mentalizing. Neuropsychologia, 49, 3130-3135. doi:10.1016/j.neuropsychologia.2011.07.023.

    Abstract

    A debated issue in the relationship between language and thought is how our linguistic abilities are involved in understanding the intentions of others (‘mentalizing’). The results of both theoretical and empirical work have been used to argue that linguistic, and more specifically, grammatical, abilities are crucial in representing the mental states of others. Here we contribute to this debate by investigating how damage to the language system influences the generation and understanding of intentional communicative behaviors. Four patients with pervasive language difficulties (severe global or agrammatic aphasia) engaged in an experimentally controlled non-verbal communication paradigm, which required signaling and understanding a communicative message. Despite their profound language problems they were able to engage in recipient design as well as intention recognition, showing similar indicators of mentalizing as have been observed in the neurologically healthy population. Our results show that aspects of the ability to communicate remain present even when core capacities of the language system are dysfunctional

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