Peter Hagoort

Publications

Displaying 1 - 100 of 426
  • Fitz, H., Hagoort, P., & Petersson, K. M. (in press). Neurobiological causal models of language processing. Neurobiology of Language.
  • Giglio, L., Ostarek, M., Sharoh, D., & Hagoort, P. (2024). Diverging neural dynamics for syntactic structure building in naturalistic speaking and listening. PNAS, 121(11): e2310766121. doi:10.1073/pnas.2310766121.

    Abstract

    The neural correlates of sentence production have been mostly studied with constraining task paradigms that introduce artificial task effects. In this study, we aimed to gain a better understanding of syntactic processing in spontaneous production vs. naturalistic comprehension. We extracted word-by-word metrics of phrase-structure building with top-down and bottom-up parsers that make different hypotheses about the timing of structure building. In comprehension, structure building proceeded in an integratory fashion and led to an increase in activity in posterior temporal and inferior frontal areas. In production, structure building was anticipatory and predicted an increase in activity in the inferior frontal gyrus. Newly developed production-specific parsers highlighted the anticipatory and incremental nature of structure building in production, which was confirmed by a converging analysis of the pausing patterns in speech. Overall, the results showed that the unfolding of syntactic processing diverges between speaking and listening.
  • Hagoort, P., & Özyürek, A. (2024). Extending the architecture of language from a multimodal perspective. Topics in Cognitive Science. Advance online publication. doi:10.1111/tops.12728.

    Abstract

    Language is inherently multimodal. In spoken languages, combined spoken and visual signals (e.g., co-speech gestures) are an integral part of linguistic structure and language representation. This requires an extension of the parallel architecture, which needs to include the visual signals concomitant to speech. We present the evidence for the multimodality of language. In addition, we propose that distributional semantics might provide a format for integrating speech and co-speech gestures in a common semantic representation.
  • Seijdel, N., Schoffelen, J.-M., Hagoort, P., & Drijvers, L. (2024). Attention drives visual processing and audiovisual integration during multimodal communication. The Journal of Neuroscience. Advance online publication. doi:10.1523/JNEUROSCI.0870-23.2023.

    Abstract

    During communication in real-life settings, our brain often needs to integrate auditory and visual information, and at the same time actively focus on the relevant sources of information, while ignoring interference from irrelevant events. The interaction between integration and attention processes remains poorly understood. Here, we use rapid invisible frequency tagging (RIFT) and magnetoencephalography (MEG) to investigate how attention affects auditory and visual information processing and integration, during multimodal communication. We presented human participants (male and female) with videos of an actress uttering action verbs (auditory; tagged at 58 Hz) accompanied by two movie clips of hand gestures on both sides of fixation (attended stimulus tagged at 65 Hz; unattended stimulus tagged at 63 Hz). Integration difficulty was manipulated by a lower-order auditory factor (clear/degraded speech) and a higher-order visual semantic factor (matching/mismatching gesture). We observed an enhanced neural response to the attended visual information during degraded speech compared to clear speech. For the unattended information, the neural response to mismatching gestures was enhanced compared to matching gestures. Furthermore, signal power at the intermodulation frequencies of the frequency tags, indexing non-linear signal interactions, was enhanced in left frontotemporal and frontal regions. Focusing on LIFG (Left Inferior Frontal Gyrus), this enhancement was specific for the attended information, for those trials that benefitted from integration with a matching gesture. Together, our results suggest that attention modulates audiovisual processing and interaction, depending on the congruence and quality of the sensory input.

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  • Terporten, R., Huizeling, E., Heidlmayr, K., Hagoort, P., & Kösem, A. (2024). The interaction of context constraints and predictive validity during sentence reading. Journal of Cognitive Neuroscience, 36(2), 225-238. doi:10.1162/jocn_a_02082.

    Abstract

    Words are not processed in isolation; instead, they are commonly embedded in phrases and sentences. The sentential context influences the perception and processing of a word. However, how this is achieved by brain processes and whether predictive mechanisms underlie this process remain a debated topic. Here, we employed an experimental paradigm in which we orthogonalized sentence context constraints and predictive validity, which was defined as the ratio of congruent to incongruent sentence endings within the experiment. While recording electroencephalography, participants read sentences with three levels of sentential context constraints (high, medium, and low). Participants were also separated into two groups that differed in their ratio of valid congruent to incongruent target words that could be predicted from the sentential context. For both groups, we investigated modulations of alpha power before, and N400 amplitude modulations after target word onset. The results reveal that the N400 amplitude gradually decreased with higher context constraints and cloze probability. In contrast, alpha power was not significantly affected by context constraint. Neither the N400 nor alpha power were significantly affected by changes in predictive validity.
  • Arana, S., Hagoort, P., Schoffelen, J.-M., & Rabovsky, M. (2023). Perceived similarity as a window into representations of integrated sentence meaning. Behavior Research Methods. Advance online publication. doi:10.3758/s13428-023-02129-x.

    Abstract

    When perceiving the world around us, we are constantly integrating pieces of information. The integrated experience consists of more than just the sum of its parts. For example, visual scenes are defined by a collection of objects as well as the spatial relations amongst them and sentence meaning is computed based on individual word semantic but also syntactic configuration. Having quantitative models of such integrated representations can help evaluate cognitive models of both language and scene perception. Here, we focus on language, and use a behavioral measure of perceived similarity as an approximation of integrated meaning representations. We collected similarity judgments of 200 subjects rating nouns or transitive sentences through an online multiple arrangement task. We find that perceived similarity between sentences is most strongly modulated by the semantic action category of the main verb. In addition, we show how non-negative matrix factorization of similarity judgment data can reveal multiple underlying dimensions reflecting both semantic as well as relational role information. Finally, we provide an example of how similarity judgments on sentence stimuli can serve as a point of comparison for artificial neural networks models (ANNs) by comparing our behavioral data against sentence similarity extracted from three state-of-the-art ANNs. Overall, our method combining the multiple arrangement task on sentence stimuli with matrix factorization can capture relational information emerging from integration of multiple words in a sentence even in the presence of strong focus on the verb.
  • Arana, S., Pesnot Lerousseau, J., & Hagoort, P. (2023). Deep learning models to study sentence comprehension in the human brain. Language, Cognition and Neuroscience. Advance online publication. doi:10.1080/23273798.2023.2198245.

    Abstract

    Recent artificial neural networks that process natural language achieve unprecedented performance in tasks requiring sentence-level understanding. As such, they could be interesting models of the integration of linguistic information in the human brain. We review works that compare these artificial language models with human brain activity and we assess the extent to which this approach has improved our understanding of the neural processes involved in natural language comprehension. Two main results emerge. First, the neural representation of word meaning aligns with the context-dependent, dense word vectors used by the artificial neural networks. Second, the processing hierarchy that emerges within artificial neural networks broadly matches the brain, but is surprisingly inconsistent across studies. We discuss current challenges in establishing artificial neural networks as process models of natural language comprehension. We suggest exploiting the highly structured representational geometry of artificial neural networks when mapping representations to brain data.

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    link to preprint
  • Hagoort, P. (2023). The language marker hypothesis. Cognition, 230: 105252. doi:10.1016/j.cognition.2022.105252.

    Abstract

    According to the language marker hypothesis language has provided homo sapiens with a rich symbolic system that plays a central role in interpreting signals delivered by our sensory apparatus, in shaping action goals, and in creating a powerful tool for reasoning and inferencing. This view provides an important correction on embodied accounts of language that reduce language to action, perception, emotion and mental simulation. The presence of a language system has, however, also important consequences for perception, action, emotion, and memory. Language stamps signals from perception, action, and emotional systems with rich cognitive markers that transform the role of these signals in the overall cognitive architecture of the human mind. This view does not deny that language is implemented by means of universal principles of neural organization. However, language creates the possibility to generate rich internal models of the world that are shaped and made accessible by the characteristics of a language system. This makes us less dependent on direct action-perception couplings and might even sometimes go at the expense of the veridicality of perception. In cognitive (neuro)science the pendulum has swung from language as the key to understand the organization of the human mind to the perspective that it is a byproduct of perception and action. It is time that it partly swings back again.
  • Hagoort, P. (2023). Zij zijn ons brein en andere beschouwingen. Nijmegen: Max Planck Instituut voor Psycholinguistiek.
  • Huizeling, E., Alday, P. M., Peeters, D., & Hagoort, P. (2023). Combining EEG and 3D-eye-tracking to study the prediction of upcoming speech in naturalistic virtual environments: A proof of principle. Neuropsychologia, 191: 108730. doi:10.1016/j.neuropsychologia.2023.108730.

    Abstract

    EEG and eye-tracking provide complementary information when investigating language comprehension. Evidence that speech processing may be facilitated by speech prediction comes from the observation that a listener's eye gaze moves towards a referent before it is mentioned if the remainder of the spoken sentence is predictable. However, changes to the trajectory of anticipatory fixations could result from a change in prediction or an attention shift. Conversely, N400 amplitudes and concurrent spectral power provide information about the ease of word processing the moment the word is perceived. In a proof-of-principle investigation, we combined EEG and eye-tracking to study linguistic prediction in naturalistic, virtual environments. We observed increased processing, reflected in theta band power, either during verb processing - when the verb was predictive of the noun - or during noun processing - when the verb was not predictive of the noun. Alpha power was higher in response to the predictive verb and unpredictable nouns. We replicated typical effects of noun congruence but not predictability on the N400 in response to the noun. Thus, the rich visual context that accompanied speech in virtual reality influenced language processing compared to previous reports, where the visual context may have facilitated processing of unpredictable nouns. Finally, anticipatory fixations were predictive of spectral power during noun processing and the length of time fixating the target could be predicted by spectral power at verb onset, conditional on the object having been fixated. Overall, we show that combining EEG and eye-tracking provides a promising new method to answer novel research questions about the prediction of upcoming linguistic input, for example, regarding the role of extralinguistic cues in prediction during language comprehension.
  • Kösem, A., Dai, B., McQueen, J. M., & Hagoort, P. (2023). Neural envelope tracking of speech does not unequivocally reflect intelligibility. NeuroImage, 272: 120040. doi:10.1016/j.neuroimage.2023.120040.

    Abstract

    During listening, brain activity tracks the rhythmic structures of speech signals. Here, we directly dissociated the contribution of neural envelope tracking in the processing of speech acoustic cues from that related to linguistic processing. We examined the neural changes associated with the comprehension of Noise-Vocoded (NV) speech using magnetoencephalography (MEG). Participants listened to NV sentences in a 3-phase training paradigm: (1) pre-training, where NV stimuli were barely comprehended, (2) training with exposure of the original clear version of speech stimulus, and (3) post-training, where the same stimuli gained intelligibility from the training phase. Using this paradigm, we tested if the neural responses of a speech signal was modulated by its intelligibility without any change in its acoustic structure. To test the influence of spectral degradation on neural envelope tracking independently of training, participants listened to two types of NV sentences (4-band and 2-band NV speech), but were only trained to understand 4-band NV speech. Significant changes in neural tracking were observed in the delta range in relation to the acoustic degradation of speech. However, we failed to find a direct effect of intelligibility on the neural tracking of speech envelope in both theta and delta ranges, in both auditory regions-of-interest and whole-brain sensor-space analyses. This suggests that acoustics greatly influence the neural tracking response to speech envelope, and that caution needs to be taken when choosing the control signals for speech-brain tracking analyses, considering that a slight change in acoustic parameters can have strong effects on the neural tracking response.
  • Mishra, C., Verdonschot, R. G., Hagoort, P., & Skantze, G. (2023). Real-time emotion generation in human-robot dialogue using large language models. Frontiers in Robotics and AI, 10: 1271610. doi:10.3389/frobt.2023.1271610.

    Abstract

    Affective behaviors enable social robots to not only establish better connections with humans but also serve as a tool for the robots to express their internal states. It has been well established that emotions are important to signal understanding in Human-Robot Interaction (HRI). This work aims to harness the power of Large Language Models (LLM) and proposes an approach to control the affective behavior of robots. By interpreting emotion appraisal as an Emotion Recognition in Conversation (ERC) tasks, we used GPT-3.5 to predict the emotion of a robot’s turn in real-time, using the dialogue history of the ongoing conversation. The robot signaled the predicted emotion using facial expressions. The model was evaluated in a within-subjects user study (N = 47) where the model-driven emotion generation was compared against conditions where the robot did not display any emotions and where it displayed incongruent emotions. The participants interacted with the robot by playing a card sorting game that was specifically designed to evoke emotions. The results indicated that the emotions were reliably generated by the LLM and the participants were able to perceive the robot’s emotions. It was found that the robot expressing congruent model-driven facial emotion expressions were perceived to be significantly more human-like, emotionally appropriate, and elicit a more positive impression. Participants also scored significantly better in the card sorting game when the robot displayed congruent facial expressions. From a technical perspective, the study shows that LLMs can be used to control the affective behavior of robots reliably in real-time. Additionally, our results could be used in devising novel human-robot interactions, making robots more effective in roles where emotional interaction is important, such as therapy, companionship, or customer service.
  • Quaresima, A., Fitz, H., Duarte, R., Van den Broek, D., Hagoort, P., & Petersson, K. M. (2023). The Tripod neuron: A minimal structural reduction of the dendritic tree. The Journal of Physiology, 601(15), 3007-3437. doi:10.1113/JP283399.

    Abstract

    Neuron models with explicit dendritic dynamics have shed light on mechanisms for coincidence detection, pathway selection and temporal filtering. However, it is still unclear which morphological and physiological features are required to capture these phenomena. In this work, we introduce the Tripod neuron model and propose a minimal structural reduction of the dendritic tree that is able to reproduce these computations. The Tripod is a three-compartment model consisting of two segregated passive dendrites and a somatic compartment modelled as an adaptive, exponential integrate-and-fire neuron. It incorporates dendritic geometry, membrane physiology and receptor dynamics as measured in human pyramidal cells. We characterize the response of the Tripod to glutamatergic and GABAergic inputs and identify parameters that support supra-linear integration, coincidence-detection and pathway-specific gating through shunting inhibition. Following NMDA spikes, the Tripod neuron generates plateau potentials whose duration depends on the dendritic length and the strength of synaptic input. When fitted with distal compartments, the Tripod encodes previous activity into a dendritic depolarized state. This dendritic memory allows the neuron to perform temporal binding, and we show that it solves transition and sequence detection tasks on which a single-compartment model fails. Thus, the Tripod can account for dendritic computations previously explained only with more detailed neuron models or neural networks. Due to its simplicity, the Tripod neuron can be used efficiently in simulations of larger cortical circuits.
  • 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
  • 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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    Table 1.XLSX
  • 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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  • Coopmans, C. W., De Hoop, H., Kaushik, K., Hagoort, P., & Martin, A. E. (2021). Structure-(in)dependent interpretation of phrases in humans and LSTMs. In Proceedings of the Society for Computation in Linguistics (SCiL 2021) (pp. 459-463).

    Abstract

    In this study, we compared the performance of a long short-term memory (LSTM) neural network to the behavior of human participants on a language task that requires hierarchically structured knowledge. We show that humans interpret ambiguous noun phrases, such as second blue ball, in line with their hierarchical constituent structure. LSTMs, instead, only do
    so after unambiguous training, and they do not systematically generalize to novel items. Overall, the results of our simulations indicate that a model can behave hierarchically without relying on hierarchical constituent structure.
  • Healthy Brain Study Consortium, Aarts, E., Akkerman, A., Altgassen, M., Bartels, R., Beckers, D., Bevelander, K., Bijleveld, E., Blaney Davidson, E., Boleij, A., Bralten, J., Cillessen, T., Claassen, J., Cools, R., Cornelissen, I., Dresler, M., Eijsvogels, T., Faber, M., Fernández, G., Figner, B., Fritsche, M. and 67 moreHealthy Brain Study Consortium, Aarts, E., Akkerman, A., Altgassen, M., Bartels, R., Beckers, D., Bevelander, K., Bijleveld, E., Blaney Davidson, E., Boleij, A., Bralten, J., Cillessen, T., Claassen, J., Cools, R., Cornelissen, I., Dresler, M., Eijsvogels, T., Faber, M., Fernández, G., Figner, B., Fritsche, M., Füllbrunn, S., Gayet, S., Van Gelder, M. M. H. J., Van Gerven, M., Geurts, S., Greven, C. U., Groefsema, M., Haak, K., Hagoort, P., Hartman, Y., Van der Heijden, B., Hermans, E., Heuvelmans, V., Hintz, F., Den Hollander, J., Hulsman, A. M., Idesis, S., Jaeger, M., Janse, E., Janzing, J., Kessels, R. P. C., Karremans, J. C., De Kleijn, W., Klein, M., Klumpers, F., Kohn, N., Korzilius, H., Krahmer, B., De Lange, F., Van Leeuwen, J., Liu, H., Luijten, M., Manders, P., Manevska, K., Marques, J. P., Matthews, J., McQueen, J. M., Medendorp, P., Melis, R., Meyer, A. S., Oosterman, J., Overbeek, L., Peelen, M., Popma, J., Postma, G., Roelofs, K., Van Rossenberg, Y. G. T., Schaap, G., Scheepers, P., Selen, L., Starren, M., Swinkels, D. W., Tendolkar, I., Thijssen, D., Timmerman, H., Tutunji, R., Tuladhar, A., Veling, H., Verhagen, M., Verkroost, J., Vink, J., Vriezekolk, V., Vrijsen, J., Vyrastekova, J., Van der Wal, S., Willems, R. M., & Willemsen, A. (2021). Protocol of the Healthy Brain Study: An accessible resource for understanding the human brain and how it dynamically and individually operates in its bio-social context. PLoS One, 16(12): e0260952. doi:10.1371/journal.pone.0260952.

    Abstract

    The endeavor to understand the human brain has seen more progress in the last few decades than in the previous two millennia. Still, our understanding of how the human brain relates to behavior in the real world and how this link is modulated by biological, social, and environmental factors is limited. To address this, we designed the Healthy Brain Study (HBS), an interdisciplinary, longitudinal, cohort study based on multidimensional, dynamic assessments in both the laboratory and the real world. Here, we describe the rationale and design of the currently ongoing HBS. The HBS is examining a population-based sample of 1,000 healthy participants (age 30-39) who are thoroughly studied across an entire year. Data are collected through cognitive, affective, behavioral, and physiological testing, neuroimaging, bio-sampling, questionnaires, ecological momentary assessment, and real-world assessments using wearable devices. These data will become an accessible resource for the scientific community enabling the next step in understanding the human brain and how it dynamically and individually operates in its bio-social context. An access procedure to the collected data and bio-samples is in place and published on https://www.healthybrainstudy.nl/en/data-and-methods.

    https://www.trialregister.nl/trial/7955

    Additional information

    supplementary material
  • Heyselaar, E., Peeters, D., & Hagoort, P. (2021). Do we predict upcoming speech content in naturalistic environments? Language, Cognition and Neuroscience, 36(4), 440-461. doi:10.1080/23273798.2020.1859568.

    Abstract

    The ability to predict upcoming actions is a hallmark of cognition. It remains unclear, however, whether the predictive behaviour observed in controlled lab environments generalises to rich, everyday settings. In four virtual reality experiments, we tested whether a well-established marker of linguistic prediction (anticipatory eye movements) replicated when increasing the naturalness of the paradigm by means of immersing participants in naturalistic scenes (Experiment 1), increasing the number of distractor objects (Experiment 2), modifying the proportion of predictable noun-referents (Experiment 3), and manipulating the location of referents relative to the joint attentional space (Experiment 4). Robust anticipatory eye movements were observed for Experiments 1–3. The anticipatory effect disappeared, however, in Experiment 4. Our findings suggest that predictive processing occurs in everyday communication if the referents are situated in the joint attentional space. Methodologically, our study confirms that ecological validity and experimental control may go hand-in-hand in the study of human predictive behaviour.
  • Misersky, J., Slivac, K., Hagoort, P., & Flecken, M. (2021). The State of the Onion: Grammatical aspect modulates object representation during event comprehension. Cognition, 214: 104744. doi:10.1016/j.cognition.2021.104744.

    Abstract

    The present ERP study assessed whether grammatical aspect is used as a cue in online event comprehension, in particular when reading about events in which an object is visually changed. While perfective aspect cues holistic event representations, including an event's endpoint, progressive aspect highlights intermediate phases of an event. In a 2 × 3 design, participants read SVO sentences describing a change-of-state event (e.g., to chop an onion), with grammatical Aspect manipulated (perfective “chopped” vs progressive “was chopping”). Thereafter, they saw a Picture of an object either having undergone substantial state-change (SC; a chopped onion), no state-change (NSC; an onion in its original state) or an unrelated object (U; a cactus, acting as control condition). Their task was to decide whether the object in the Picture was mentioned in the sentence. We focused on N400 modulation, with ERPs time-locked to picture onset. U pictures elicited an N400 response as expected, suggesting detection of categorical mismatches in object type. For SC and NSC pictures, a whole-head follow-up analysis revealed a P300, implying people were engaged in detailed evaluation of pictures of matching objects. SC pictures received most positive responses overall. Crucially, there was an interaction of Aspect and Picture: SC pictures resulted in a higher amplitude P300 after sentences in the perfective compared to the progressive. Thus, while the perfective cued for a holistic event representation, including the resultant state of the affected object (i.e., the chopped onion) constraining object representations online, the progressive defocused event completion and object-state change. Grammatical aspect thus guided online event comprehension by cueing the visual representation(s) of an object's state.
  • Preisig, B., Riecke, L., Sjerps, M. J., Kösem, A., Kop, B. R., Bramson, B., Hagoort, P., & Hervais-Adelman, A. (2021). Selective modulation of interhemispheric connectivity by transcranial alternating current stimulation influences binaural integration. Proceedings of the National Academy of Sciences of the United States of America, 118(7): e2015488118. doi:10.1073/pnas.2015488118.

    Abstract

    Brain connectivity plays a major role in the encoding, transfer, and
    integration of sensory information. Interregional synchronization
    of neural oscillations in the γ-frequency band has been suggested
    as a key mechanism underlying perceptual integration. In a recent
    study, we found evidence for this hypothesis showing that the
    modulation of interhemispheric oscillatory synchrony by means of
    bihemispheric high-density transcranial alternating current stimulation
    (HD-TACS) affects binaural integration of dichotic acoustic features.
    Here, we aimed to establish a direct link between oscillatory
    synchrony, effective brain connectivity, and binaural integration.
    We experimentally manipulated oscillatory synchrony (using bihemispheric
    γ-TACS with different interhemispheric phase lags) and
    assessed the effect on effective brain connectivity and binaural integration
    (as measured with functional MRI and a dichotic listening
    task, respectively). We found that TACS reduced intrahemispheric
    connectivity within the auditory cortices and antiphase (interhemispheric
    phase lag 180°) TACS modulated connectivity between the
    two auditory cortices. Importantly, the changes in intra- and interhemispheric
    connectivity induced by TACS were correlated with
    changes in perceptual integration. Our results indicate that γ-band
    synchronization between the two auditory cortices plays a functional
    role in binaural integration, supporting the proposed role
    of interregional oscillatory synchrony in perceptual integration.
  • Slivac, K., Hervais-Adelman, A., Hagoort, P., & Flecken, M. (2021). Linguistic labels cue biological motion perception and misperception. Scientific Reports, 11: 17239. doi:10.1038/s41598-021-96649-1.

    Abstract

    Linguistic labels exert a particularly strong top-down influence on perception. The potency of this influence has been ascribed to their ability to evoke category-diagnostic features of concepts. In doing this, they facilitate the formation of a perceptual template concordant with those features, effectively biasing perceptual activation towards the labelled category. In this study, we employ a cueing paradigm with moving, point-light stimuli across three experiments, in order to examine how the number of biological motion features (form and kinematics) encoded in lexical cues modulates the efficacy of lexical top-down influence on perception. We find that the magnitude of lexical influence on biological motion perception rises as a function of the number of biological motion-relevant features carried by both cue and target. When lexical cues encode multiple biological motion features, this influence is robust enough to mislead participants into reporting erroneous percepts, even when a masking level yielding high performance is used.
  • Arana, S., Marquand, A., Hulten, A., Hagoort, P., & Schoffelen, J.-M. (2020). Sensory modality-independent activation of the brain network for language. The Journal of Neuroscience, 40(14), 2914-2924. doi:10.1523/JNEUROSCI.2271-19.2020.

    Abstract

    The meaning of a sentence can be understood, whether presented in written or spoken form. Therefore it is highly probable that brain processes supporting language comprehension are at least partly independent of sensory modality. To identify where and when in the brain language processing is independent of sensory modality, we directly compared neuromagnetic brain signals of 200 human subjects (102 males) either reading or listening to sentences. We used multiset canonical correlation analysis to align individual subject data in a way that boosts those aspects of the signal that are common to all, allowing us to capture word-by-word signal variations, consistent across subjects and at a fine temporal scale. Quantifying this consistency in activation across both reading and listening tasks revealed a mostly left hemispheric cortical network. Areas showing consistent activity patterns include not only areas previously implicated in higher-level language processing, such as left prefrontal, superior & middle temporal areas and anterior temporal lobe, but also parts of the control-network as well as subcentral and more posterior temporal-parietal areas. Activity in this supramodal sentence processing network starts in temporal areas and rapidly spreads to the other regions involved. The findings do not only indicate the involvement of a large network of brain areas in supramodal language processing, but also indicate that the linguistic information contained in the unfolding sentences modulates brain activity in a word-specific manner across subjects.
  • Casasanto, D., Casasanto, L. S., Gijssels, T., & Hagoort, P. (2020). The Reverse Chameleon Effect: Negative social consequences of anatomical mimicry. Frontiers in Psychology, 11: 1876. doi:10.3389/fpsyg.2020.01876.

    Abstract

    Bodily mimicry often makes the mimickee have more positive feelings about the mimicker. Yet, little is known about the causes of mimicry’s social effects. When people mimic each other’s bodily movements face to face, they can either adopt a mirrorwise perspective (moving in the same absolute direction) or an anatomical perspective (moving in the same direction relative to their own bodies). Mirrorwise mimicry maximizes visuo-spatial similarity between the mimicker and mimickee, whereas anatomical mimicry maximizes the similarity in the states of their motor systems. To compare the social consequences of visuo-spatial and motoric similarity, we asked participants to converse with an embodied virtual agent (VIRTUO), who mimicked their head movements either mirrorwise, anatomically, or not at all. Compared to participants who were not mimicked, those who were mimicked mirrorwise tended to rate VIRTUO more positively, but those who were mimicked anatomically rated him more negatively. During face-to-face conversation, mirrorwise and anatomical mimicry have opposite social consequences. Results suggest that visuo-spatial similarity between mimicker and mimickee, not similarity in motor system activity, gives rise to the positive social effects of bodily mimicry.
  • Fitz, H., Uhlmann, M., Van den Broek, D., Duarte, R., Hagoort, P., & Petersson, K. M. (2020). Neuronal spike-rate adaptation supports working memory in language processing. Proceedings of the National Academy of Sciences of the United States of America, 117(34), 20881-20889. doi:10.1073/pnas.2000222117.

    Abstract

    Language processing involves the ability to store and integrate pieces of
    information in working memory over short periods of time. According to
    the dominant view, information is maintained through sustained, elevated
    neural activity. Other work has argued that short-term synaptic facilitation
    can serve as a substrate of memory. Here, we propose an account where
    memory is supported by intrinsic plasticity that downregulates neuronal
    firing rates. Single neuron responses are dependent on experience and we
    show through simulations that these adaptive changes in excitability pro-
    vide memory on timescales ranging from milliseconds to seconds. On this
    account, spiking activity writes information into coupled dynamic variables
    that control adaptation and move at slower timescales than the membrane
    potential. From these variables, information is continuously read back into
    the active membrane state for processing. This neuronal memory mech-
    anism does not rely on persistent activity, excitatory feedback, or synap-
    tic plasticity for storage. Instead, information is maintained in adaptive
    conductances that reduce firing rates and can be accessed directly with-
    out cued retrieval. Memory span is systematically related to both the time
    constant of adaptation and baseline levels of neuronal excitability. Inter-
    ference effects within memory arise when adaptation is long-lasting. We
    demonstrate that this mechanism is sensitive to context and serial order
    which makes it suitable for temporal integration in sequence processing
    within the language domain. We also show that it enables the binding of
    linguistic features over time within dynamic memory registers. This work
    provides a step towards a computational neurobiology of language.
  • Hagoort, P. (2020). Taal. In O. Van den Heuvel, Y. Van der Werf, B. Schmand, & B. Sabbe (Eds.), Leerboek neurowetenschappen voor de klinische psychiatrie (pp. 234-239). Amsterdam: Boom Uitgevers.
  • Heidlmayr, K., Weber, K., Takashima, A., & Hagoort, P. (2020). No title, no theme: The joined neural space between speakers and listeners during production and comprehension of multi-sentence discourse. Cortex, 130, 111-126. doi:10.1016/j.cortex.2020.04.035.

    Abstract

    Speakers and listeners usually interact in larger discourses than single words or even single sentences. The goal of the present study was to identify the neural bases reflecting how the mental representation of the situation denoted in a multi-sentence discourse (situation model) is constructed and shared between speakers and listeners. An fMRI study using a variant of the ambiguous text paradigm was designed. Speakers (n=15) produced ambiguous texts in the scanner and listeners (n=27) subsequently listened to these texts in different states of ambiguity: preceded by a highly informative, intermediately informative or no title at all. Conventional BOLD activation analyses in listeners, as well as inter-subject correlation analyses between the speakers’ and the listeners’ hemodynamic time courses were performed. Critically, only the processing of disambiguated, coherent discourse with an intelligible situation model representation involved (shared) activation in bilateral lateral parietal and medial prefrontal regions. This shared spatiotemporal pattern of brain activation between the speaker and the listener suggests that the process of memory retrieval in medial prefrontal regions and the binding of retrieved information in the lateral parietal cortex constitutes a core mechanism underlying the communication of complex conceptual representations.

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  • Heilbron, M., Richter, D., Ekman, M., Hagoort, P., & De Lange, F. P. (2020). Word contexts enhance the neural representation of individual letters in early visual cortex. Nature Communications, 11: 321. doi:10.1038/s41467-019-13996-4.

    Abstract

    Visual context facilitates perception, but how this is neurally implemented remains unclear. One example of contextual facilitation is found in reading, where letters are more easily identified when embedded in a word. Bottom-up models explain this word advantage as a post-perceptual decision bias, while top-down models propose that word contexts enhance perception itself. Here, we arbitrate between these accounts by presenting words and nonwords and probing the representational fidelity of individual letters using functional magnetic resonance imaging. In line with top-down models, we find that word contexts enhance letter representations in early visual cortex. Moreover, we observe increased coupling between letter information in visual cortex and brain activity in key areas of the reading network, suggesting these areas may be the source of the enhancement. Our results provide evidence for top-down representational enhancement in word recognition, demonstrating that word contexts can modulate perceptual processing already at the earliest visual regions.

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  • Hoeksema, N., Wiesmann, M., Kiliaan, A., Hagoort, P., & Vernes, S. C. (2020). Bats and the comparative neurobiology of vocal learning. In A. Ravignani, C. Barbieri, M. Flaherty, Y. Jadoul, E. Lattenkamp, H. Little, M. Martins, K. Mudd, & T. Verhoef (Eds.), The Evolution of Language: Proceedings of the 13th International Conference (Evolang13) (pp. 165-167). Nijmegen: The Evolution of Language Conferences.
  • Kösem, A., Bosker, H. R., Jensen, O., Hagoort, P., & Riecke, L. (2020). Biasing the perception of spoken words with transcranial alternating current stimulation. Journal of Cognitive Neuroscience, 32(8), 1428-1437. doi:10.1162/jocn_a_01579.

    Abstract

    Recent neuroimaging evidence suggests that the frequency of entrained oscillations in auditory cortices influences the perceived duration of speech segments, impacting word perception (Kösem et al. 2018). We further tested the causal influence of neural entrainment frequency during speech processing, by manipulating entrainment with continuous transcranial alternating
    current stimulation (tACS) at distinct oscillatory frequencies (3 Hz and 5.5 Hz) above the auditory cortices. Dutch participants listened to speech and were asked to report their percept of a target Dutch word, which contained a vowel with an ambiguous duration. Target words
    were presented either in isolation (first experiment) or at the end of spoken sentences (second experiment). We predicted that the tACS frequency would influence neural entrainment and
    therewith how speech is perceptually sampled, leading to a perceptual over- or underestimation of the vowel’s duration. Whereas results from Experiment 1 did not confirm this prediction, results from experiment 2 suggested a small effect of tACS frequency on target word
    perception: Faster tACS lead to more long-vowel word percepts, in line with the previous neuroimaging findings. Importantly, the difference in word perception induced by the different tACS frequencies was significantly larger in experiment 1 vs. experiment 2, suggesting that the
    impact of tACS is dependent on the sensory context. tACS may have a stronger effect on spoken word perception when the words are presented in continuous speech as compared to when they are isolated, potentially because prior (stimulus-induced) entrainment of brain oscillations
    might be a prerequisite for tACS to be effective.

    Additional information

    Data availability
  • Preisig, B., Sjerps, M. J., Hervais-Adelman, A., Kösem, A., Hagoort, P., & Riecke, L. (2020). Bilateral gamma/delta transcranial alternating current stimulation affects interhemispheric speech sound integration. Journal of Cognitive Neuroscience, 32(7), 1242-1250. doi:10.1162/jocn_a_01498.

    Abstract

    Perceiving speech requires the integration of different speech cues, that is, formants. When the speech signal is split so that different cues are presented to the right and left ear (dichotic listening), comprehension requires the integration of binaural information. Based on prior electrophysiological evidence, we hypothesized that the integration of dichotically presented speech cues is enabled by interhemispheric phase synchronization between primary and secondary auditory cortex in the gamma frequency band. We tested this hypothesis by applying transcranial alternating current stimulation (TACS) bilaterally above the superior temporal lobe to induce or disrupt interhemispheric gamma-phase coupling. In contrast to initial predictions, we found that gamma TACS applied in-phase above the two hemispheres (interhemispheric lag 0°) perturbs interhemispheric integration of speech cues, possibly because the applied stimulation perturbs an inherent phase lag between the left and right auditory cortex. We also observed this disruptive effect when applying antiphasic delta TACS (interhemispheric lag 180°). We conclude that interhemispheric phase coupling plays a functional role in interhemispheric speech integration. The direction of this effect may depend on the stimulation frequency.
  • Takashima, A., Konopka, A. E., Meyer, A. S., Hagoort, P., & Weber, K. (2020). Speaking in the brain: The interaction between words and syntax in sentence production. Journal of Cognitive Neuroscience, 32(8), 1466-1483. doi:10.1162/jocn_a_01563.

    Abstract

    This neuroimaging study investigated the neural infrastructure of sentence-level language production. We compared brain activation patterns, as measured with BOLD-fMRI, during production of sentences that differed in verb argument structures (intransitives, transitives, ditransitives) and the lexical status of the verb (known verbs or pseudoverbs). The experiment consisted of 30 mini-blocks of six sentences each. Each mini-block started with an example for the type of sentence to be produced in that block. On each trial in the mini-blocks, participants were first given the (pseudo-)verb followed by three geometric shapes to serve as verb arguments in the sentences. Production of sentences with known verbs yielded greater activation compared to sentences with pseudoverbs in the core language network of the left inferior frontal gyrus, the left posterior middle temporalgyrus, and a more posterior middle temporal region extending into the angular gyrus, analogous to effects observed in language comprehension. Increasing the number of verb arguments led to greater activation in an overlapping left posterior middle temporal gyrus/angular gyrus area, particularly for known verbs, as well as in the bilateral precuneus. Thus, producing sentences with more complex structures using existing verbs leads to increased activation in the language network, suggesting some reliance on memory retrieval of stored lexical–syntactic information during sentence production. This study thus provides evidence from sentence-level language production in line with functional models of the language network that have so far been mainly based on single-word production, comprehension, and language processing in aphasia.
  • Tan, Y., & Hagoort, P. (2020). Catecholaminergic modulation of semantic processing in sentence comprehension. Cerebral Cortex, 30(12), 6426-6443. doi:10.1093/cercor/bhaa204.

    Abstract

    Catecholamine (CA) function has been widely implicated in cognitive functions that are tied to the prefrontal cortex and striatal areas. The present study investigated the effects of methylphenidate, which is a CA agonist, on the electroencephalogram (EEG) response related to semantic processing using a double-blind, placebo-controlled, randomized, crossover, within-subject design. Forty-eight healthy participants read semantically congruent or incongruent sentences after receiving 20-mg methylphenidate or a placebo while their brain activity was monitored with EEG. To probe whether the catecholaminergic modulation is task-dependent, in one condition participants had to focus on comprehending the sentences, while in the other condition, they only had to attend to the font size of the sentence. The results demonstrate that methylphenidate has a task-dependent effect on semantic processing. Compared to placebo, when semantic processing was task-irrelevant, methylphenidate enhanced the detection of semantic incongruence as indexed by a larger N400 amplitude in the incongruent sentences; when semantic processing was task-relevant, methylphenidate induced a larger N400 amplitude in the semantically congruent condition, which was followed by a larger late positive complex effect. These results suggest that CA-related neurotransmitters influence language processing, possibly through the projections between the prefrontal cortex and the striatum, which contain many CA receptors.
  • Franken, M. K., Acheson, D. J., McQueen, J. M., Hagoort, P., & Eisner, F. (2019). Consistency influences altered auditory feedback processing. Quarterly Journal of Experimental Psychology, 72(10), 2371-2379. doi:10.1177/1747021819838939.

    Abstract

    Previous research on the effect of perturbed auditory feedback in speech production has focused on two types of responses. In the short term, speakers generate compensatory motor commands in response to unexpected perturbations. In the longer term, speakers adapt feedforward motor programmes in response to feedback perturbations, to avoid future errors. The current study investigated the relation between these two types of responses to altered auditory feedback. Specifically, it was hypothesised that consistency in previous feedback perturbations would influence whether speakers adapt their feedforward motor programmes. In an altered auditory feedback paradigm, formant perturbations were applied either across all trials (the consistent condition) or only to some trials, whereas the others remained unperturbed (the inconsistent condition). The results showed that speakers’ responses were affected by feedback consistency, with stronger speech changes in the consistent condition compared with the inconsistent condition. Current models of speech-motor control can explain this consistency effect. However, the data also suggest that compensation and adaptation are distinct processes, which are not in line with all current models.
  • Hagoort, P. (Ed.). (2019). Human language: From genes and brains to behavior. Cambridge, MA: MIT Press.
  • Hagoort, P., & Beckmann, C. F. (2019). Key issues and future directions: The neural architecture for language. In P. Hagoort (Ed.), Human language: From genes and brains to behavior (pp. 527-532). Cambridge, MA: MIT Press.
  • Hagoort, P. (2019). Introduction. In P. Hagoort (Ed.), Human language: From genes and brains to behavior (pp. 1-6). Cambridge, MA: MIT Press.
  • Hagoort, P. (2019). The meaning making mechanism(s) behind the eyes and between the ears. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 375: 20190301. doi:10.1098/rstb.2019.0301.

    Abstract

    In this contribution, the following four questions are discussed: (i) where is meaning?; (ii) what is meaning?; (iii) what is the meaning of mechanism?; (iv) what are the mechanisms of meaning? I will argue that meanings are in the head. Meanings have multiple facets, but minimally one needs to make a distinction between single word meanings (lexical meaning) and the meanings of multi-word utterances. The latter ones cannot be retrieved from memory, but need to be constructed on the fly. A mechanistic account of the meaning-making mind requires an analysis at both a functional and a neural level, the reason being that these levels are causally interdependent. I will show that an analysis exclusively focusing on patterns of brain activation lacks explanatory power. Finally, I shall present an initial sketch of how the dynamic interaction between temporo-parietal areas and inferior frontal cortex might instantiate the interpretation of linguistic utterances in the context of a multimodal setting and ongoing discourse information.
  • Hagoort, P. (2019). The neurobiology of language beyond single word processing. Science, 366(6461), 55-58. doi:10.1126/science.aax0289.

    Abstract

    In this Review, I propose a multiple-network view for the neurobiological basis of distinctly human language skills. A much more complex picture of interacting brain areas emerges than in the classical neurobiological model of language. This is because using language is more than single-word processing, and much goes on beyond the information given in the acoustic or orthographic tokens that enter primary sensory cortices. This requires the involvement of multiple networks with functionally nonoverlapping contributions

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  • Heilbron, M., Ehinger, B., Hagoort, P., & De Lange, F. P. (2019). Tracking naturalistic linguistic predictions with deep neural language models. In Proceedings of the 2019 Conference on Cognitive Computational Neuroscience (pp. 424-427). doi:10.32470/CCN.2019.1096-0.

    Abstract

    Prediction in language has traditionally been studied using
    simple designs in which neural responses to expected
    and unexpected words are compared in a categorical
    fashion. However, these designs have been contested
    as being ‘prediction encouraging’, potentially exaggerating
    the importance of prediction in language understanding.
    A few recent studies have begun to address
    these worries by using model-based approaches to probe
    the effects of linguistic predictability in naturalistic stimuli
    (e.g. continuous narrative). However, these studies
    so far only looked at very local forms of prediction, using
    models that take no more than the prior two words into
    account when computing a word’s predictability. Here,
    we extend this approach using a state-of-the-art neural
    language model that can take roughly 500 times longer
    linguistic contexts into account. Predictability estimates
    fromthe neural network offer amuch better fit to EEG data
    from subjects listening to naturalistic narrative than simpler
    models, and reveal strong surprise responses akin to
    the P200 and N400. These results show that predictability
    effects in language are not a side-effect of simple designs,
    and demonstrate the practical use of recent advances
    in AI for the cognitive neuroscience of language.
  • Hulten, A., Schoffelen, J.-M., Udden, J., Lam, N. H. L., & Hagoort, P. (2019). How the brain makes sense beyond the processing of single words – An MEG study. NeuroImage, 186, 586-594. doi:10.1016/j.neuroimage.2018.11.035.

    Abstract

    Human language processing involves combinatorial operations that make human communication stand out in the animal kingdom. These operations rely on a dynamic interplay between the inferior frontal and the posterior temporal cortices. Using source reconstructed magnetoencephalography, we tracked language processing in the brain, in order to investigate how individual words are interpreted when part of sentence context. The large sample size in this study (n = 68) allowed us to assess how event-related activity is associated across distinct cortical areas, by means of inter-areal co-modulation within an individual. We showed that, within 500 ms of seeing a word, the word's lexical information has been retrieved and unified with the sentence context. This does not happen in a strictly feed-forward manner, but by means of co-modulation between the left posterior temporal cortex (LPTC) and left inferior frontal cortex (LIFC), for each individual word. The co-modulation of LIFC and LPTC occurs around 400 ms after the onset of each word, across the progression of a sentence. Moreover, these core language areas are supported early on by the attentional network. The results provide a detailed description of the temporal orchestration related to single word processing in the context of ongoing language.

    Additional information

    1-s2.0-S1053811918321165-mmc1.pdf
  • Mongelli, V., Meijs, E. L., Van Gaal, S., & Hagoort, P. (2019). No language unification without neural feedback: How awareness affects sentence processing. Neuroimage, 202: 116063. doi:10.1016/j.neuroimage.2019.116063.

    Abstract

    How does the human brain combine a finite number of words to form an infinite variety of sentences? According to the Memory, Unification and Control (MUC) model, sentence processing requires long-range feedback from the left inferior frontal cortex (LIFC) to left posterior temporal cortex (LPTC). Single word processing however may only require feedforward propagation of semantic information from sensory regions to LPTC. Here we tested the claim that long-range feedback is required for sentence processing by reducing visual awareness of words using a masking technique. Masking disrupts feedback processing while leaving feedforward processing relatively intact. Previous studies have shown that masked single words still elicit an N400 ERP effect, a neural signature of semantic incongruency. However, whether multiple words can be combined to form a sentence under reduced levels of awareness is controversial. To investigate this issue, we performed two experiments in which we measured electroencephalography (EEG) while 40 subjects performed a masked priming task. Words were presented either successively or simultaneously, thereby forming a short sentence that could be congruent or incongruent with a target picture. This sentence condition was compared with a typical single word condition. In the masked condition we only found an N400 effect for single words, whereas in the unmasked condition we observed an N400 effect for both unmasked sentences and single words. Our findings suggest that long-range feedback processing is required for sentence processing, but not for single word processing.
  • Schoffelen, J.-M., Oostenveld, R., Lam, N. H. L., Udden, J., Hulten, A., & Hagoort, P. (2019). A 204-subject multimodal neuroimaging dataset to study language processing. Scientific Data, 6(1): 17. doi:10.1038/s41597-019-0020-y.

    Abstract

    This dataset, colloquially known as the Mother Of Unification Studies (MOUS) dataset, contains multimodal neuroimaging data that has been acquired from 204 healthy human subjects. The neuroimaging protocol consisted of magnetic resonance imaging (MRI) to derive information at high spatial resolution about brain anatomy and structural connections, and functional data during task, and at rest. In addition, magnetoencephalography (MEG) was used to obtain high temporal resolution electrophysiological measurements during task, and at rest. All subjects performed a language task, during which they processed linguistic utterances that either consisted of normal or scrambled sentences. Half of the subjects were reading the stimuli, the other half listened to the stimuli. The resting state measurements consisted of 5 minutes eyes-open for the MEG and 7 minutes eyes-closed for fMRI. The neuroimaging data, as well as the information about the experimental events are shared according to the Brain Imaging Data Structure (BIDS) format. This unprecedented neuroimaging language data collection allows for the investigation of various aspects of the neurobiological correlates of language.
  • Schoot, L., Hagoort, P., & Segaert, K. (2019). Stronger syntactic alignment in the presence of an interlocutor. Frontiers in Psychology, 10: 685. doi:10.3389/fpsyg.2019.00685.

    Abstract

    Speakers are influenced by the linguistic context: hearing one syntactic alternative leads to an increased chance that the speaker will repeat this structure in the subsequent utterance (i.e., syntactic priming, or structural persistence). Top-down influences, such as whether a conversation partner (or, interlocutor) is present, may modulate the degree to which syntactic priming occurs. In the current study, we indeed show that the magnitude of syntactic alignment increases when speakers are interacting with an interlocutor as opposed to doing the experiment alone. The structural persistence effect for passive sentences is stronger in the presence of an interlocutor than when no interlocutor is present (i.e., when the participant is primed by a recording). We did not find evidence, however, that a speaker’s syntactic priming magnitude is influenced by the degree of their conversation partner’s priming magnitude. Together, these results support a mediated account of syntactic priming, in which syntactic choices are not only affected by preceding linguistic input, but also by top-down influences, such as the speakers’ communicative intent.
  • Sharoh, D., Van Mourik, T., Bains, L. J., Segaert, K., Weber, K., Hagoort, P., & Norris, D. (2019). Laminar specific fMRI reveals directed interactions in distributed networks during language processing. Proceedings of the National Academy of Sciences of the United States of America, 116(42), 21185-21190. doi:10.1073/pnas.1907858116.

    Abstract

    Interactions between top-down and bottom-up information streams are integral to brain function but challenging to measure noninvasively. Laminar resolution, functional MRI (lfMRI) is sensitive to depth-dependent properties of the blood oxygen level-dependent (BOLD) response, which can be potentially related to top-down and bottom-up signal contributions. In this work, we used lfMRI to dissociate the top-down and bottom-up signal contributions to the left occipitotemporal sulcus (LOTS) during word reading. We further demonstrate that laminar resolution measurements could be used to identify condition-specific distributed networks on the basis of whole-brain connectivity patterns specific to the depth-dependent BOLD signal. The networks corresponded to top-down and bottom-up signal pathways targeting the LOTS during word reading. We show that reading increased the top-down BOLD signal observed in the deep layers of the LOTS and that this signal uniquely related to the BOLD response in other language-critical regions. These results demonstrate that lfMRI can reveal important patterns of activation that are obscured at standard resolution. In addition to differences in activation strength as a function of depth, we also show meaningful differences in the interaction between signals originating from different depths both within a region and with the rest of the brain. We thus show that lfMRI allows the noninvasive measurement of directed interaction between brain regions and is capable of resolving different connectivity patterns at submillimeter resolution, something previously considered to be exclusively in the domain of invasive recordings.
  • Udden, J., Hulten, A., Bendt, K., Mineroff, Z., Kucera, K. S., Vino, A., Fedorenko, E., Hagoort, P., & Fisher, S. E. (2019). Towards robust functional neuroimaging genetics of cognition. Journal of Neuroscience, 39(44), 8778-8787. doi:10.1523/JNEUROSCI.0888-19.2019.

    Abstract

    A commonly held assumption in cognitive neuroscience is that, because measures of human brain function are closer to underlying biology than distal indices of behavior/cognition, they hold more promise for uncovering genetic pathways. Supporting this view is an influential fMRI-based study of sentence reading/listening by Pinel et al. (2012), who reported that common DNA variants in specific candidate genes were associated with altered neural activation in language-related regions of healthy individuals that carried them. In particular, different single-nucleotide polymorphisms (SNPs) of FOXP2 correlated with variation in task-based activation in left inferior frontal and precentral gyri, whereas a SNP at the KIAA0319/TTRAP/THEM2 locus was associated with variable functional asymmetry of the superior temporal sulcus. Here, we directly test each claim using a closely matched neuroimaging genetics approach in independent cohorts comprising 427 participants, four times larger than the original study of 94 participants. Despite demonstrating power to detect associations with substantially smaller effect sizes than those of the original report, we do not replicate any of the reported associations. Moreover, formal Bayesian analyses reveal substantial to strong evidence in support of the null hypothesis (no effect). We highlight key aspects of the original investigation, common to functional neuroimaging genetics studies, which could have yielded elevated false-positive rates. Genetic accounts of individual differences in cognitive functional neuroimaging are likely to be as complex as behavioral/cognitive tests, involving many common genetic variants, each of tiny effect. Reliable identification of true biological signals requires large sample sizes, power calculations, and validation in independent cohorts with equivalent paradigms.

    SIGNIFICANCE STATEMENT A pervasive idea in neuroscience is that neuroimaging-based measures of brain function, being closer to underlying neurobiology, are more amenable for uncovering links to genetics. This is a core assumption of prominent studies that associate common DNA variants with altered activations in task-based fMRI, despite using samples (10–100 people) that lack power for detecting the tiny effect sizes typical of genetically complex traits. Here, we test central findings from one of the most influential prior studies. Using matching paradigms and substantially larger samples, coupled to power calculations and formal Bayesian statistics, our data strongly refute the original findings. We demonstrate that neuroimaging genetics with task-based fMRI should be subject to the same rigorous standards as studies of other complex traits.
  • Weber, K., Christiansen, M., Indefrey, P., & Hagoort, P. (2019). Primed from the start: Syntactic priming during the first days of language learning. Language Learning, 69(1), 198-221. doi:10.1111/lang.12327.

    Abstract

    New linguistic information must be integrated into our existing language system. Using a novel experimental task that incorporates a syntactic priming paradigm into artificial language learning, we investigated how new grammatical regularities and words are learned. This innovation allowed us to control the language input the learner received, while the syntactic priming paradigm provided insight into the nature of the underlying syntactic processing machinery. The results of the present study pointed to facilitatory syntactic processing effects within the first days of learning: Syntactic and lexical priming effects revealed participants’ sensitivity to both novel words and word orders. This suggested that novel syntactic structures and their meaning (form–function mapping) can be acquired rapidly through incidental learning. More generally, our study indicated similar mechanisms for learning and processing in both artificial and natural languages, with implications for the relationship between first and second language learning.
  • Zhu, Z., Bastiaansen, M. C. M., Hakun, J. G., Petersson, K. M., Wang, S., & Hagoort, P. (2019). Semantic unification modulates N400 and BOLD signal change in the brain: A simultaneous EEG-fMRI study. Journal of Neurolinguistics, 52: 100855. doi:10.1016/j.jneuroling.2019.100855.

    Abstract

    Semantic unification during sentence comprehension has been associated with amplitude change of the N400 in event-related potential (ERP) studies, and activation in the left inferior frontal gyrus (IFG) in functional magnetic resonance imaging (fMRI) studies. However, the specificity of this activation to semantic unification remains unknown. To more closely examine the brain processes involved in semantic unification, we employed simultaneous EEG-fMRI to time-lock the semantic unification related N400 change, and integrated trial-by-trial variation in both N400 and BOLD change beyond the condition-level BOLD change difference measured in traditional fMRI analyses. Participants read sentences in which semantic unification load was parametrically manipulated by varying cloze probability. Separately, ERP and fMRI results replicated previous findings, in that semantic unification load parametrically modulated the amplitude of N400 and cortical activation. Integrated EEG-fMRI analyses revealed a different pattern in which functional activity in the left IFG and bilateral supramarginal gyrus (SMG) was associated with N400 amplitude, with the left IFG activation and bilateral SMG activation being selective to the condition-level and trial-level of semantic unification load, respectively. By employing the EEG-fMRI integrated analyses, this study among the first sheds light on how to integrate trial-level variation in language comprehension.
  • 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.

    Additional information

    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.

    Additional information

    mmc1.docx
  • 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.
  • Dai, B., McQueen, J. M., Hagoort, P., & Kösem, A. (2017). Pure linguistic interference during comprehension of competing speech signals. The Journal of the Acoustical Society of America, 141, EL249-EL254. doi:10.1121/1.4977590.

    Abstract

    Speech-in-speech perception can be challenging because the processing of competing acoustic and linguistic information leads to informational masking. Here, a method is proposed to isolate the linguistic component of informational masking while keeping the distractor's acoustic information unchanged. Participants performed a dichotic listening cocktail-party task before and after training on 4-band noise-vocoded sentences that became intelligible through the training. Distracting noise-vocoded speech interfered more with target speech comprehension after training (i.e., when intelligible) than before training (i.e., when unintelligible) at −3 dB SNR. These findings confirm that linguistic and acoustic information have distinct masking effects during speech-in‐speech comprehension
  • Franken, M. K., Eisner, F., Schoffelen, J.-M., Acheson, D. J., Hagoort, P., & McQueen, J. M. (2017). Audiovisual recalibration of vowel categories. In Proceedings of Interspeech 2017 (pp. 655-658). doi:10.21437/Interspeech.2017-122.

    Abstract

    One of the most daunting tasks of a listener is to map a
    continuous auditory stream onto known speech sound
    categories and lexical items. A major issue with this mapping
    problem is the variability in the acoustic realizations of sound
    categories, both within and across speakers. Past research has
    suggested listeners may use visual information (e.g., lipreading)
    to calibrate these speech categories to the current
    speaker. Previous studies have focused on audiovisual
    recalibration of consonant categories. The present study
    explores whether vowel categorization, which is known to show
    less sharply defined category boundaries, also benefit from
    visual cues.
    Participants were exposed to videos of a speaker
    pronouncing one out of two vowels, paired with audio that was
    ambiguous between the two vowels. After exposure, it was
    found that participants had recalibrated their vowel categories.
    In addition, individual variability in audiovisual recalibration is
    discussed. It is suggested that listeners’ category sharpness may
    be related to the weight they assign to visual information in
    audiovisual speech perception. Specifically, listeners with less
    sharp categories assign more weight to visual information
    during audiovisual speech recognition.
  • Franken, M. K., Acheson, D. J., McQueen, J. M., Eisner, F., & Hagoort, P. (2017). Individual variability as a window on production-perception interactions in speech motor control. The Journal of the Acoustical Society of America, 142(4), 2007-2018. doi:10.1121/1.5006899.

    Abstract

    An important part of understanding speech motor control consists of capturing the
    interaction between speech production and speech perception. This study tests a
    prediction of theoretical frameworks that have tried to account for these interactions: if
    speech production targets are specified in auditory terms, individuals with better
    auditory acuity should have more precise speech targets, evidenced by decreased
    within-phoneme variability and increased between-phoneme distance. A study was
    carried out consisting of perception and production tasks in counterbalanced order.
    Auditory acuity was assessed using an adaptive speech discrimination task, while
    production variability was determined using a pseudo-word reading task. Analyses of
    the production data were carried out to quantify average within-phoneme variability as
    well as average between-phoneme contrasts. Results show that individuals not only
    vary in their production and perceptual abilities, but that better discriminators have
    more distinctive vowel production targets (that is, targets with less within-phoneme
    variability and greater between-phoneme distances), confirming the initial hypothesis.
    This association between speech production and perception did not depend on local
    phoneme density in vowel space. This study suggests that better auditory acuity leads
    to more precise speech production targets, which may be a consequence of auditory
    feedback affecting speech production over time.
  • Hagoort, P. (2017). It is the facts, stupid. In J. Brockman, F. Van der Wa, & H. Corver (Eds.), Wetenschappelijke parels: het belangrijkste wetenschappelijke nieuws volgens 193 'briljante geesten'. Amsterdam: Maven Press.
  • Hagoort, P. (2017). Don't forget neurobiology: An experimental approach to linguistic representation. Commentary on Branigan and Pickering "An experimental approach to linguistic representation". Behavioral and Brain Sciences, 40: e292. doi:10.1017/S0140525X17000401.

    Abstract

    Acceptability judgments are no longer acceptable as the holy grail for testing the nature of linguistic representations. Experimental and quantitative methods should be used to test theoretical claims in psycholinguistics. These methods should include not only behavior, but also the more recent possibilities to probe the neural codes for language-relevant representation
  • Hagoort, P. (2017). The core and beyond in the language-ready brain. Neuroscience and Biobehavioral Reviews, 81, 194-204. doi:10.1016/j.neubiorev.2017.01.048.

    Abstract

    In this paper a general cognitive architecture of spoken language processing is specified. This is followed by an account of how this cognitive architecture is instantiated in the human brain. Both the spatial aspects of the networks for language are discussed, as well as the temporal dynamics and the underlying neurophysiology. A distinction is proposed between networks for coding/decoding linguistic information and additional networks for getting from coded meaning to speaker meaning, i.e. for making the inferences that enable the listener to understand the intentions of the speaker
  • Hagoort, P. (2017). The neural basis for primary and acquired language skills. In E. Segers, & P. Van den Broek (Eds.), Developmental Perspectives in Written Language and Literacy: In honor of Ludo Verhoeven (pp. 17-28). Amsterdam: Benjamins. doi:10.1075/z.206.02hag.

    Abstract

    Reading is a cultural invention that needs to recruit cortical infrastructure that was not designed for it (cultural recycling of cortical maps). In the case of reading both visual cortex and networks for speech processing are recruited. Here I discuss current views on the neurobiological underpinnings of spoken language that deviate in a number of ways from the classical Wernicke-Lichtheim-Geschwind model. More areas than Broca’s and Wernicke’s region are involved in language. Moreover, a division along the axis of language production and language comprehension does not seem to be warranted. Instead, for central aspects of language processing neural infrastructure is shared between production and comprehension. Arguments are presented in favor of a dynamic network view, in which the functionality of a region is co-determined by the network of regions in which it is embedded at particular moments in time. Finally, core regions of language processing need to interact with other networks (e.g. the attentional networks and the ToM network) to establish full functionality of language and communication. The consequences of this architecture for reading are discussed.
  • Hartung, F., Hagoort, P., & Willems, R. M. (2017). Readers select a comprehension mode independent of pronoun: Evidence from fMRI during narrative comprehension. Brain and Language, 170, 29-38. doi:10.1016/j.bandl.2017.03.007.

    Abstract

    Perspective is a crucial feature for communicating about events. Yet it is unclear how linguistically encoded perspective relates to cognitive perspective taking. Here, we tested the effect of perspective taking with short literary stories. Participants listened to stories with 1st or 3rd person pronouns referring to the protagonist, while undergoing fMRI. When comparing action events with 1st and 3rd person pronouns, we found no evidence for a neural dissociation depending on the pronoun. A split sample approach based on the self-reported experience of perspective taking revealed 3 comprehension preferences. One group showed a strong 1st person preference, another a strong 3rd person preference, while a third group engaged in 1st and 3rd person perspective taking simultaneously. Comparing brain activations of the groups revealed different neural networks. Our results suggest that comprehension is perspective dependent, but not on the perspective suggested by the text, but on the reader’s (situational) preference
  • Hartung, F., Withers, P., Hagoort, P., & Willems, R. M. (2017). When fiction is just as real as fact: No differences in reading behavior between stories believed to be based on true or fictional events. Frontiers in Psychology, 8: 1618. doi:10.3389/fpsyg.2017.01618.

    Abstract

    Experiments have shown that compared to fictional texts, readers read factual texts faster and have better memory for described situations. Reading fictional texts on the other hand seems to improve memory for exact wordings and expressions. Most of these studies used a ‘newspaper’ versus ‘literature’ comparison. In the present study, we investigated the effect of reader’s expectation to whether information is true or fictional with a subtler manipulation by labelling short stories as either based on true or fictional events. In addition, we tested whether narrative perspective or individual preference in perspective taking affects reading true or fictional stories differently. In an online experiment, participants (final N=1742) read one story which was introduced as based on true events or as fictional (factor fictionality). The story could be narrated in either 1st or 3rd person perspective (factor perspective). We measured immersion in and appreciation of the story, perspective taking, as well as memory for events. We found no evidence that knowing a story is fictional or based on true events influences reading behavior or experiential aspects of reading. We suggest that it is not whether a story is true or fictional, but rather expectations towards certain reading situations (e.g. reading newspaper or literature) which affect behavior by activating appropriate reading goals. Results further confirm that narrative perspective partially influences perspective taking and experiential aspects of reading
  • Heyselaar, E., Hagoort, P., & Segaert, K. (2017). How social opinion influences syntactic processing – An investigation using virtual reality. PLoS One, 12(4): e0174405. doi:10.1371/journal.pone.0174405.
  • Heyselaar, E., Hagoort, P., & Segaert, K. (2017). In dialogue with an avatar, language behavior is identical to dialogue with a human partner. Behavior Research Methods, 49(1), 46-60. doi:10.3758/s13428-015-0688-7.

    Abstract

    The use of virtual reality (VR) as a methodological tool is becoming increasingly popular in behavioral research as its flexibility allows for a wide range of applications. This new method has not been as widely accepted in the field of psycholinguistics, however, possibly due to the assumption that language processing during human-computer interactions does not accurately reflect human-human interactions. Yet at the same time there is a growing need to study human-human language interactions in a tightly controlled context, which has not been possible using existing methods. VR, however, offers experimental control over parameters that cannot be (as finely) controlled in the real world. As such, in this study we aim to show that human-computer language interaction is comparable to human-human language interaction in virtual reality. In the current study we compare participants’ language behavior in a syntactic priming task with human versus computer partners: we used a human partner, a human-like avatar with human-like facial expressions and verbal behavior, and a computer-like avatar which had this humanness removed. As predicted, our study shows comparable priming effects between the human and human-like avatar suggesting that participants attributed human-like agency to the human-like avatar. Indeed, when interacting with the computer-like avatar, the priming effect was significantly decreased. This suggests that when interacting with a human-like avatar, sentence processing is comparable to interacting with a human partner. Our study therefore shows that VR is a valid platform for conducting language research and studying dialogue interactions in an ecologically valid manner.
  • Heyselaar, E., Segaert, K., Walvoort, S. J., Kessels, R. P., & Hagoort, P. (2017). The role of nondeclarative memory in the skill for language: Evidence from syntactic priming in patients with amnesia. Neuropsychologia, 101, 97-105. doi:10.1016/j.neuropsychologia.2017.04.033.

    Abstract

    Syntactic priming, the phenomenon in which participants adopt the linguistic behaviour of their partner, is widely used in psycholinguistics to investigate syntactic operations. Although the phenomenon of syntactic priming is well documented, the memory system that supports the retention of this syntactic information long enough to influence future utterances, is not as widely investigated. We aim to shed light on this issue by assessing patients with Korsakoff's amnesia on an active-passive syntactic priming task and compare their performance to controls matched in age, education, and premorbid intelligence. Patients with Korsakoff's syndrome display deficits in all subdomains of declarative memory, yet their nondeclarative memory remains intact, making them an ideal patient group to determine which memory system supports syntactic priming. In line with the hypothesis that syntactic priming relies on nondeclarative memory, the patient group shows strong priming tendencies (12.6% passive structure repetition). Our healthy control group did not show a priming tendency, presumably due to cognitive interference between declarative and nondeclarative memory. We discuss the results in relation to amnesia, aging, and compensatory mechanisms.
  • Peeters, D., Snijders, T. M., Hagoort, P., & Ozyurek, A. (2017). Linking language to the visual world: Neural correlates of comprehending verbal reference to objects through pointing and visual cues. Neuropsychologia, 95, 21-29. doi:10.1016/j.neuropsychologia.2016.12.004.

    Abstract

    In everyday communication speakers often refer in speech and/or gesture to objects in their immediate environment, thereby shifting their addressee's attention to an intended referent. The neurobiological infrastructure involved in the comprehension of such basic multimodal communicative acts remains unclear. In an event-related fMRI study, we presented participants with pictures of a speaker and two objects while they concurrently listened to her speech. In each picture, one of the objects was singled out, either through the speaker's index-finger pointing gesture or through a visual cue that made the object perceptually more salient in the absence of gesture. A mismatch (compared to a match) between speech and the object singled out by the speaker's pointing gesture led to enhanced activation in left IFG and bilateral pMTG, showing the importance of these areas in conceptual matching between speech and referent. Moreover, a match (compared to a mismatch) between speech and the object made salient through a visual cue led to enhanced activation in the mentalizing system, arguably reflecting an attempt to converge on a jointly attended referent in the absence of pointing. These findings shed new light on the neurobiological underpinnings of the core communicative process of comprehending a speaker's multimodal referential act and stress the power of pointing as an important natural device to link speech to objects.
  • Schoffelen, J.-M., Hulten, A., Lam, N. H. L., Marquand, A. F., Udden, J., & Hagoort, P. (2017). Frequency-specific directed interactions in the human brain network for language. Proceedings of the National Academy of Sciences of the United States of America, 114(30), 8083-8088. doi:10.1073/pnas.1703155114.

    Abstract

    The brain’s remarkable capacity for language requires bidirectional interactions between functionally specialized brain regions. We used magnetoencephalography to investigate interregional interactions in the brain network for language while 102 participants were reading sentences. Using Granger causality analysis, we identified inferior frontal cortex and anterior temporal regions to receive widespread input and middle temporal regions to send widespread output. This fits well with the notion that these regions play a central role in language processing. Characterization of the functional topology of this network, using data-driven matrix factorization, which allowed for partitioning into a set of subnetworks, revealed directed connections at distinct frequencies of interaction. Connections originating from temporal regions peaked at alpha frequency, whereas connections originating from frontal and parietal regions peaked at beta frequency. These findings indicate that the information flow between language-relevant brain areas, which is required for linguistic processing, may depend on the contributions of distinct brain rhythms

    Additional information

    pnas.201703155SI.pdf
  • Silva, S., Folia, V., Hagoort, P., & Petersson, K. M. (2017). The P600 in Implicit Artificial Grammar Learning. Cognitive Science, 41(1), 137-157. doi:10.1111/cogs.12343.

    Abstract

    The suitability of the Artificial Grammar Learning (AGL) paradigm to capture relevant aspects of the acquisition of linguistic structures has been empirically tested in a number of EEG studies. Some have shown a syntax-related P600 component, but it has not been ruled out that the AGL P600 effect is a response to surface features (e.g., subsequence familiarity) rather than the underlying syntax structure. Therefore, in this study, we controlled for the surface characteristics of the test sequences (associative chunk strength) and recorded the EEG before (baseline preference classification) and
    after (preference and grammaticality classification) exposure to a grammar. A typical, centroparietal P600 effect was elicited by grammatical violations after exposure, suggesting that the AGL P600 effect signals a response to structural irregularities. Moreover, preference and grammaticality classification showed a qualitatively similar ERP profile, strengthening the idea that the implicit structural mere
    exposure paradigm in combination with preference classification is a suitable alternative to the traditional grammaticality classification test.
  • Ye, Z., Stolk, A., Toni, I., & Hagoort, P. (2017). Oxytocin modulates semantic integration in speech comprehension. Journal of Cognitive Neuroscience, 29, 267-276. doi:10.1162/jocn_a_01044.

    Abstract

    Listeners interpret utterances by integrating information from multiple sources including word level semantics and world knowledge. When the semantics of an expression is inconsistent with his or her knowledge about the world, the listener may have to search through the conceptual space for alternative possible world scenarios that can make the expression more acceptable. Such cognitive exploration requires considerable computational resources and might depend on motivational factors. This study explores whether and how oxytocin, a neuropeptide known to influence socialmotivation by reducing social anxiety and enhancing affiliative tendencies, can modulate the integration of world knowledge and sentence meanings. The study used a betweenparticipant double-blind randomized placebo-controlled design. Semantic integration, indexed with magnetoencephalography through the N400m marker, was quantified while 45 healthymale participants listened to sentences that were either congruent or incongruent with facts of the world, after receiving intranasally delivered oxytocin or placebo. Compared with congruent sentences, world knowledge incongruent sentences elicited a stronger N400m signal from the left inferior frontal and anterior temporal regions and medial pFC (the N400m effect) in the placebo group. Oxytocin administration significantly attenuated the N400meffect at both sensor and cortical source levels throughout the experiment, in a state-like manner. Additional electrophysiological markers suggest that the absence of the N400m effect in the oxytocin group is unlikely due to the lack of early sensory or semantic processing or a general downregulation of attention. These findings suggest that oxytocin drives listeners to resolve challenges of semantic integration, possibly by promoting the cognitive exploration of alternative possible world scenarios.
  • Tsuji, S., Fikkert, P., Minagawa, Y., Dupoux, E., Filippin, L., Versteegh, M., Hagoort, P., & Cristia, A. (2017). The more, the better? Behavioral and neural correlates of frequent and infrequent vowel exposure. Developmental Psychobiology, 59, 603-612. doi:10.1002/dev.21534.

    Abstract

    A central assumption in the perceptual attunement literature holds that exposure to a speech sound contrast leads to improvement in native speech sound processing. However, whether the amount of exposure matters for this process has not been put to a direct test. We elucidated indicators of frequency-dependent perceptual attunement by comparing 5–8-month-old Dutch infants’ discrimination of tokens containing a highly frequent [hɪt-he:t] and a highly infrequent [hʏt-hø:t] native vowel contrast as well as a non-native [hɛt-hæt] vowel contrast in a behavioral visual habituation paradigm (Experiment 1). Infants discriminated both native contrasts similarly well, but did not discriminate the non-native contrast. We sought further evidence for subtle differences in the processing of the two native contrasts using near-infrared spectroscopy and a within-participant design (Experiment 2). The neuroimaging data did not provide additional evidence that responses to native contrasts are modulated by frequency of exposure. These results suggest that even large differences in exposure to a native contrast may not directly translate to behavioral and neural indicators of perceptual attunement, raising the possibility that frequency of exposure does not influence improvements in discriminating native contrasts.

    Additional information

    dev21534-sup-0001-SuppInfo-S1.docx
  • Udden, J., Ingvar, M., Hagoort, P., & Petersson, K. M. (2017). Broca’s region: A causal role in implicit processing of grammars with crossed non-adjacent dependencies. Cognition, 164, 188-198. doi:10.1016/j.cognition.2017.03.010.

    Abstract

    Non-adjacent dependencies are challenging for the language learning machinery and are acquired later than adjacent dependencies. In this transcranial magnetic stimulation (TMS) study, we show that participants successfully discriminated between grammatical and non-grammatical sequences after having implicitly acquired an artificial language with crossed non-adjacent dependencies. Subsequent to transcranial magnetic stimulation of Broca’s region, discrimination was impaired compared to when a language-irrelevant control region (vertex) was stimulated. These results support the view that Broca’s region is engaged in structured sequence processing and extend previous functional neuroimaging results on artificial grammar learning (AGL) in two directions: first, the results establish that Broca’s region is a causal component in the processing of non-adjacent dependencies, and second, they show that implicit processing of non-adjacent dependencies engages Broca’s region. Since patients with lesions in Broca’s region do not always show grammatical processing difficulties, the result that Broca’s region is causally linked to processing of non-adjacent dependencies is a step towards clarification of the exact nature of syntactic deficits caused by lesions or perturbation to Broca’s region. Our findings are consistent with previous results and support a role for Broca’s region in general structured sequence processing, rather than a specific role for the processing of hierarchically organized sentence structure.
  • Udden, J., Snijders, T. M., Fisher, S. E., & Hagoort, P. (2017). A common variant of the CNTNAP2 gene is associated with structural variation in the left superior occipital gyrus. Brain and Language, 172, 16-21. doi:10.1016/j.bandl.2016.02.003.

    Abstract

    The CNTNAP2 gene encodes a cell-adhesion molecule that influences the properties of neural networks and the morphology and density of neurons and glial cells. Previous studies have shown association of CNTNAP2 variants with language-related phenotypes in health and disease. Here, we report associations of a common CNTNAP2 polymorphism (rs7794745) with variation in grey matter in a region in the dorsal visual stream. We tried to replicate an earlier study on 314 subjects by Tan and colleagues (2010), but now in a substantially larger group of more than 1700 subjects. Carriers of the T allele showed reduced grey matter volume in left superior occipital gyrus, while we did not replicate associations with grey matter volume in other regions identified by Tan et al (2010). Our work illustrates the importance of independent replication in neuroimaging genetic studies of language-related candidate genes.
  • Asaridou, S. S., Takashima, A., Dediu, D., Hagoort, P., & McQueen, J. M. (2016). Repetition suppression in the left inferior frontal gyrus predicts tone learning performance. Cerebral Cortex, 26(6), 2728-2742. doi:10.1093/cercor/bhv126.

    Abstract

    Do individuals differ in how efficiently they process non-native sounds? To what extent do these differences relate to individual variability in sound-learning aptitude? We addressed these questions by assessing the sound-learning abilities of Dutch native speakers as they were trained on non-native tone contrasts. We used fMRI repetition suppression to the non-native tones to measure participants' neuronal processing efficiency before and after training. Although all participants improved in tone identification with training, there was large individual variability in learning performance. A repetition suppression effect to tone was found in the bilateral inferior frontal gyri (IFGs) before training. No whole-brain effect was found after training; a region-of-interest analysis, however, showed that, after training, repetition suppression to tone in the left IFG correlated positively with learning. That is, individuals who were better in learning the non-native tones showed larger repetition suppression in this area. Crucially, this was true even before training. These findings add to existing evidence that the left IFG plays an important role in sound learning and indicate that individual differences in learning aptitude stem from differences in the neuronal efficiency with which non-native sounds are processed.
  • Dimitrova, D. V., Chu, M., Wang, L., Ozyurek, A., & Hagoort, P. (2016). Beat that word: How listeners integrate beat gesture and focus in multimodal speech discourse. Journal of Cognitive Neuroscience, 28(9), 1255-1269. doi:10.1162/jocn_a_00963.

    Abstract

    Communication is facilitated when listeners allocate their attention to important information (focus) in the message, a process called "information structure." Linguistic cues like the preceding context and pitch accent help listeners to identify focused information. In multimodal communication, relevant information can be emphasized by nonverbal cues like beat gestures, which represent rhythmic nonmeaningful hand movements. Recent studies have found that linguistic and nonverbal attention cues are integrated independently in single sentences. However, it is possible that these two cues interact when information is embedded in context, because context allows listeners to predict what information is important. In an ERP study, we tested this hypothesis and asked listeners to view videos capturing a dialogue. In the critical sentence, focused and nonfocused words were accompanied by beat gestures, grooming hand movements, or no gestures. ERP results showed that focused words are processed more attentively than nonfocused words as reflected in an N1 and P300 component. Hand movements also captured attention and elicited a P300 component. Importantly, beat gesture and focus interacted in a late time window of 600-900 msec relative to target word onset, giving rise to a late positivity when nonfocused words were accompanied by beat gestures. Our results show that listeners integrate beat gesture with the focus of the message and that integration costs arise when beat gesture falls on nonfocused information. This suggests that beat gestures fulfill a unique focusing function in multimodal discourse processing and that they have to be integrated with the information structure of the message.
  • Gijssels, T., Staum Casasanto, L., Jasmin, K., Hagoort, P., & Casasanto, D. (2016). Speech accommodation without priming: The case of pitch. Discourse Processes, 53(4), 233-251. doi:10.1080/0163853X.2015.1023965.

    Abstract

    People often accommodate to each other's speech by aligning their linguistic production with their partner's. According to an influential theory, the Interactive Alignment Model (Pickering & Garrod, 2004), alignment is the result of priming. When people perceive an utterance, the corresponding linguistic representations are primed, and become easier to produce. Here we tested this theory by investigating whether pitch (F0) alignment shows two characteristic signatures of priming: dose dependence and persistence. In a virtual reality experiment, we manipulated the pitch of a virtual interlocutor's speech to find out (a.) whether participants accommodated to the agent's F0, (b.) whether the amount of accommodation increased with increasing exposure to the agent's speech, and (c.) whether changes to participants' F0 persisted beyond the conversation. Participants accommodated to the virtual interlocutor, but accommodation did not increase in strength over the conversation, and it disappeared immediately after the conversation ended. Results argue against a priming-based account of F0 accommodation, and indicate that an alternative mechanism is needed to explain alignment along continuous dimensions of language such as speech rate and pitch.
  • Hagoort, P. (2016). MUC (Memory, Unification, Control): A Model on the Neurobiology of Language Beyond Single Word Processing. In G. Hickok, & S. Small (Eds.), Neurobiology of language (pp. 339-347). Amsterdam: Elsever. doi:10.1016/B978-0-12-407794-2.00028-6.

    Abstract

    A neurobiological model of language is discussed that overcomes the shortcomings of the classical Wernicke-Lichtheim-Geschwind model. It is based on a subdivision of language processing into three components: Memory, Unification, and Control. The functional components as well as the neurobiological underpinnings of the model are discussed. In addition, the need for extension beyond the classical core regions for language is shown. Attentional networks as well as networks for inferential processing are crucial to realize language comprehension beyond single word processing and beyond decoding propositional content.
  • Hagoort, P. (2016). Zij zijn ons brein. In J. Brockman (Ed.), Machines die denken: Invloedrijke denkers over de komst van kunstmatige intelligentie (pp. 184-186). Amsterdam: Maven Publishing.
  • Hartung, F., Burke, M., Hagoort, P., & Willems, R. M. (2016). Taking perspective: Personal pronouns affect experiential aspects of literary reading. PLoS One, 11(5): e0154732. doi:10.1371/journal.pone.0154732.

    Abstract

    Personal pronouns have been shown to influence cognitive perspective taking during comprehension. Studies using single sentences found that 3rd person pronouns facilitate the construction of a mental model from an observer’s perspective, whereas 2nd person pronouns support an actor’s perspective. The direction of the effect for 1st person pronouns seems to depend on the situational context. In the present study, we investigated how personal pronouns influence discourse comprehension when people read fiction stories and if this has consequences for affective components like emotion during reading or appreciation of the story. We wanted to find out if personal pronouns affect immersion and arousal, as well as appreciation of fiction. In a natural reading paradigm, we measured electrodermal activity and story immersion, while participants read literary stories with 1st and 3rd person pronouns referring to the protagonist. In addition, participants rated and ranked the stories for appreciation. Our results show that stories with 1st person pronouns lead to higher immersion. Two factors—transportation into the story world and mental imagery during reading—in particular showed higher scores for 1st person as compared to 3rd person pronoun stories. In contrast, arousal as measured by electrodermal activity seemed tentatively higher for 3rd person pronoun stories. The two measures of appreciation were not affected by the pronoun manipulation. Our findings underscore the importance of perspective for language processing, and additionally show which aspects of the narrative experience are influenced by a change in perspective.
  • Kunert, R., Willems, R. M., & Hagoort, P. (2016). An independent psychometric evaluation of the PROMS measure of music perception skills. PLoS One, 11(7): e0159103. doi:10.1371/journal.pone.0159103.

    Abstract

    The Profile of Music Perception Skills (PROMS) is a recently developed measure of perceptual music skills which has been shown to have promising psychometric properties. In this paper we extend the evaluation of its brief version to three kinds of validity using an individual difference approach. The brief PROMS displays good discriminant validity with working memory, given that it does not correlate with backward digit span (r = .04). Moreover, it shows promising criterion validity (association with musical training (r = .45), musicianship status (r = .48), and self-rated musical talent (r = .51)). Finally, its convergent validity, i.e. relation to an unrelated measure of music perception skills, was assessed by correlating the brief PROMS to harmonic closure judgment accuracy. Two independent samples point to good convergent validity of the brief PROMS (r = .36; r = .40). The same association is still significant in one of the samples when including self-reported music skill in a partial correlation (rpartial = .30; rpartial = .17). Overall, the results show that the brief version of the PROMS displays a very good pattern of construct validity. Especially its tuning subtest stands out as a valuable part for music skill evaluations in Western samples. We conclude by briefly discussing the choice faced by music cognition researchers between different musical aptitude measures of which the brief PROMS is a well evaluated example.
  • Kunert, R., Willems, R. M., & Hagoort, P. (2016). Language influences music harmony perception: effects of shared syntactic integration resources beyond attention. Royal Society Open Science, 3(2): 150685. doi:10.1098/rsos.150685.

    Abstract

    Many studies have revealed shared music–language processing resources by finding an influence of music harmony manipulations on concurrent language processing. However, the nature of the shared resources has remained ambiguous. They have been argued to be syntax specific and thus due to shared syntactic integration resources. An alternative view regards them as related to general attention and, thus, not specific to syntax. The present experiments evaluated these accounts by investigating the influence of language on music. Participants were asked to provide closure judgements on harmonic sequences in order to assess the appropriateness of sequence endings. At the same time participants read syntactic garden-path sentences. Closure judgements revealed a change in harmonic processing as the result of reading a syntactically challenging word. We found no influence of an arithmetic control manipulation (experiment 1) or semantic garden-path sentences (experiment 2). Our results provide behavioural evidence for a specific influence of linguistic syntax processing on musical harmony judgements. A closer look reveals that the shared resources appear to be needed to hold a harmonic key online in some form of syntactic working memory or unification workspace related to the integration of chords and words. Overall, our results support the syntax specificity of shared music–language processing resources.
  • Lam, N. H. L., Schoffelen, J.-M., Udden, J., Hulten, A., & Hagoort, P. (2016). Neural activity during sentence processing as reflected in theta, alpha, beta and gamma oscillations. NeuroImage, 142(15), 43-54. doi:10.1016/j.neuroimage.2016.03.007.

    Abstract

    We used magnetoencephalography (MEG) to explore the spatio-temporal dynamics of neural oscillations associated with sentence processing, in 102 participants. We quantified changes in oscillatory power as the sentence unfolded, and in response to individual words in the sentence. For words early in a sentence compared to those late in the same sentence, we observed differences in left temporal and frontal areas, and bilateral frontal and right parietal regions for the theta, alpha, and beta frequency bands. The neural response to words in a sentence differed from the response to words in scrambled sentences in left-lateralized theta, alpha, beta, and gamma. The theta band effects suggest that a sentential context facilitates lexical retrieval, and that this facilitation is stronger for words late in the sentence. Effects in the alpha and beta band may reflect the unification of semantic and syntactic information, and are suggestive of easier unification late in a sentence. The gamma oscillations are indicative of predicting the upcoming word during sentence processing. In conclusion, changes in oscillatory neuronal activity capture aspects of sentence processing. Our results support earlier claims that language (sentence) processing recruits areas distributed across both hemispheres, and extends beyond the classical language regions

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