Peter Hagoort

Publications

Displaying 1 - 70 of 70
  • Asaridou, S. S., Hagoort, P., & McQueen, J. M. (2015). Effects of early bilingual experience with a tone and a non-tone language on speech-music. PLoS One, 10(12): e0144225. doi:10.1371/journal.pone.0144225.

    Abstract

    We investigated music and language processing in a group of early bilinguals who spoke a tone language and a non-tone language (Cantonese and Dutch). We assessed online speech-music processing interactions, that is, interactions that occur when speech and music are processed simultaneously in songs, with a speeded classification task. In this task, participants judged sung pseudowords either musically (based on the direction of the musical interval) or phonologically (based on the identity of the sung vowel). We also assessed longer-term effects of linguistic experience on musical ability, that is, the influence of extensive prior experience with language when processing music. These effects were assessed with a task in which participants had to learn to identify musical intervals and with four pitch-perception tasks. Our hypothesis was that due to their experience in two different languages using lexical versus intonational tone, the early Cantonese-Dutch bilinguals would outperform the Dutch control participants. In online processing, the Cantonese-Dutch bilinguals processed speech and music more holistically than controls. This effect seems to be driven by experience with a tone language, in which integration of segmental and pitch information is fundamental. Regarding longer-term effects of linguistic experience, we found no evidence for a bilingual advantage in either the music-interval learning task or the pitch-perception tasks. Together, these results suggest that being a Cantonese-Dutch bilingual does not have any measurable longer-term effects on pitch and music processing, but does have consequences for how speech and music are processed jointly.

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  • Baggio, G., van Lambalgen, M., & Hagoort, P. (2015). Logic as Marr's computational level: Four case studies. Topics in Cognitive Science, 7, 287-298. doi:10.1111/tops.12125.

    Abstract

    We sketch four applications of Marr's levels-of-analysis methodology to the relations between logic and experimental data in the cognitive neuroscience of language and reasoning. The first part of the paper illustrates the explanatory power of computational level theories based on logic. We show that a Bayesian treatment of the suppression task in reasoning with conditionals is ruled out by EEG data, supporting instead an analysis based on defeasible logic. Further, we describe how results from an EEG study on temporal prepositions can be reanalyzed using formal semantics, addressing a potential confound. The second part of the article demonstrates the predictive power of logical theories drawing on EEG data on processing progressive constructions and on behavioral data on conditional reasoning in people with autism. Logical theories can constrain processing hypotheses all the way down to neurophysiology, and conversely neuroscience data can guide the selection of alternative computational level models of cognition.
  • Bašnákova, J., Van Berkum, J. J. A., Weber, K., & Hagoort, P. (2015). A job interview in the MRI scanner: How does indirectness affect addressees and overhearers? Neuropsychologia, 76, 79-91. doi:10.1016/j.neuropsychologia.2015.03.030.

    Abstract

    In using language, people not only exchange information, but also navigate their social world – for example, they can express themselves indirectly to avoid losing face. In this functional magnetic resonance imaging study, we investigated the neural correlates of interpreting face-saving indirect replies, in a situation where participants only overheard the replies as part of a conversation between two other people, as well as in a situation where the participants were directly addressed themselves. We created a fictional job interview context where indirect replies serve as a natural communicative strategy to attenuate one’s shortcomings, and asked fMRI participants to either pose scripted questions and receive answers from three putative job candidates (addressee condition) or to listen to someone else interview the same candidates (overhearer condition). In both cases, the need to evaluate the candidate ensured that participants had an active interest in comprehending the replies. Relative to direct replies, face-saving indirect replies increased activation in medial prefrontal cortex, bilateral temporo-parietal junction (TPJ), bilateral inferior frontal gyrus and bilateral middle temporal gyrus, in active overhearers and active addressees alike, with similar effect size, and comparable to findings obtained in an earlier passive listening study (Bašnáková et al., 2013). In contrast, indirectness effects in bilateral anterior insula and pregenual ACC, two regions implicated in emotional salience and empathy, were reliably stronger in addressees than in active overhearers. Our findings indicate that understanding face-saving indirect language requires additional cognitive perspective-taking and other discourse-relevant cognitive processing, to a comparable extent in active overhearers and addressees. Furthermore, they indicate that face-saving indirect language draws upon affective systems more in addressees than in overhearers, presumably because the addressee is the one being managed by a face-saving reply. In all, face-saving indirectness provides a window on the cognitive as well as affect-related neural systems involved in human communication.
  • Bastiaansen, M. C. M., & Hagoort, P. (2015). Frequency-based segregation of syntactic and semantic unification during online sentence level language comprehension. Journal of Cognitive Neuroscience, 27(11), 2095-2107. doi:10.1162/jocn_a_00829.

    Abstract

    During sentence level language comprehension, semantic and syntactic unification are functionally distinct operations. Nevertheless, both recruit roughly the same brain areas (spatially overlapping networks in the left frontotemporal cortex) and happen at the same time (in the first few hundred milliseconds after word onset). We tested the hypothesis that semantic and syntactic unification are segregated by means of neuronal synchronization of the functionally relevant networks in different frequency ranges: gamma (40 Hz and up) for semantic unification and lower beta (10–20 Hz) for syntactic unification. EEG power changes were quantified as participants read either correct sentences, syntactically correct though meaningless sentences (syntactic prose), or sentences that did not contain any syntactic structure (random word lists). Other sentences contained either a semantic anomaly or a syntactic violation at a critical word in the sentence. Larger EEG gamma-band power was observed for semantically coherent than for semantically anomalous sentences. Similarly, beta-band power was larger for syntactically correct sentences than for incorrect ones. These results confirm the existence of a functional dissociation in EEG oscillatory dynamics during sentence level language comprehension that is compatible with the notion of a frequency-based segregation of syntactic and semantic unification.
  • Francken, J. C., Meijs, E. L., Ridderinkhof, O. M., Hagoort, P., de Lange, F. P., & van Gaal, S. (2015). Manipulating word awareness dissociates feed-forward from feedback models of language-perception interactions. Neuroscience of consciousness, 1. doi:10.1093/nc/niv003.

    Abstract

    Previous studies suggest that linguistic material can modulate visual perception, but it is unclear at which level of processing these interactions occur. Here we aim to dissociate between two competing models of language–perception interactions: a feed-forward and a feedback model. We capitalized on the fact that the models make different predictions on the role of feedback. We presented unmasked (aware) or masked (unaware) words implying motion (e.g. “rise,” “fall”), directly preceding an upward or downward visual motion stimulus. Crucially, masking leaves intact feed-forward information processing from low- to high-level regions, whereas it abolishes subsequent feedback. Under this condition, participants remained faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. This suggests that language–perception interactions are driven by the feed-forward convergence of linguistic and perceptual information at higher-level conceptual and decision stages.
  • Francken, J. C., Meijs, E. L., Hagoort, P., van Gaal, S., & de Lange, F. P. (2015). Exploring the automaticity of language-perception interactions: Effects of attention and awareness. Scientific Reports, 5: 17725. doi:10.1038/srep17725.

    Abstract

    Previous studies have shown that language can modulate visual perception, by biasing and/
    or enhancing perceptual performance. However, it is still debated where in the brain visual and
    linguistic information are integrated, and whether the effects of language on perception are
    automatic and persist even in the absence of awareness of the linguistic material. Here, we aimed
    to explore the automaticity of language-perception interactions and the neural loci of these
    interactions in an fMRI study. Participants engaged in a visual motion discrimination task (upward
    or downward moving dots). Before each trial, a word prime was briefly presented that implied
    upward or downward motion (e.g., “rise”, “fall”). These word primes strongly influenced behavior:
    congruent motion words sped up reaction times and improved performance relative to incongruent
    motion words. Neural congruency effects were only observed in the left middle temporal gyrus,
    showing higher activity for congruent compared to incongruent conditions. This suggests that higherlevel
    conceptual areas rather than sensory areas are the locus of language-perception interactions.
    When motion words were rendered unaware by means of masking, they still affected visual motion
    perception, suggesting that language-perception interactions may rely on automatic feed-forward
    integration of perceptual and semantic material in language areas of the brain.
  • Francken, J. C., Kok, P., Hagoort, P., & De Lange, F. P. (2015). The behavioral and neural effects of language on motion perception. Journal of Cognitive Neuroscience, 27(1), 175-184. doi:10.1162/jocn_a_00682.

    Abstract

    Perception does not function as an isolated module but is tightly linked with other cognitive functions. Several studies have demonstrated an influence of language on motion perception, but it remains debated at which level of processing this modulation takes place. Some studies argue for an interaction in perceptual areas, but it is also possible that the interaction is mediated by "language areas" that integrate linguistic and visual information. Here, we investigated whether language-perception interactions were specific to the language-dominant left hemisphere by comparing the effects of language on visual material presented in the right (RVF) and left visual fields (LVF). Furthermore, we determined the neural locus of the interaction using fMRI. Participants performed a visual motion detection task. On each trial, the visual motion stimulus was presented in either the LVF or in the RVF, preceded by a centrally presented word (e.g., "rise"). The word could be congruent, incongruent, or neutral with regard to the direction of the visual motion stimulus that was presented subsequently. Participants were faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. Interestingly, the speed benefit was present only for motion stimuli that were presented in the RVF. We observed a neural counterpart of the behavioral facilitation effects in the left middle temporal gyrus, an area involved in semantic processing of verbal material. Together, our results suggest that semantic information about motion retrieved in language regions may automatically modulate perceptual decisions about motion.
  • Franken, M. K., McQueen, J. M., Hagoort, P., & Acheson, D. J. (2015). Assessing the link between speech perception and production through individual differences. In Proceedings of the 18th International Congress of Phonetic Sciences. Glasgow: the University of Glasgow.

    Abstract

    This study aims to test a prediction of recent
    theoretical frameworks in speech motor control: if speech production targets are specified in auditory
    terms, people with better auditory acuity should have more precise speech targets.
    To investigate this, we had participants perform speech perception and production tasks in a counterbalanced order. To assess speech perception acuity, we used an adaptive speech discrimination
    task. To assess variability in speech production, participants performed a pseudo-word reading task; formant values were measured for each recording.
    We predicted that speech production variability to correlate inversely with discrimination performance.
    The results suggest that people do vary in their production and perceptual abilities, and that better discriminators have more distinctive vowel production targets, confirming our prediction. This
    study highlights the importance of individual
    differences in the study of speech motor control, and sheds light on speech production-perception interaction.
  • Franken, M. K., Hagoort, P., & Acheson, D. J. (2015). Modulations of the auditory M100 in an Imitation Task. Brain and Language, 142, 18-23. doi:10.1016/j.bandl.2015.01.001.

    Abstract

    Models of speech production explain event-related suppression of the auditory cortical
    response as reflecting a comparison between auditory predictions and feedback. The present MEG
    study was designed to test two predictions from this framework: 1) whether the reduced auditory
    response varies as a function of the mismatch between prediction and feedback; 2) whether individual
    variation in this response is predictive of speech-motor adaptation.
    Participants alternated between online imitation and listening tasks. In the imitation task, participants
    began each trial producing the same vowel (/e/) and subsequently listened to and imitated auditorilypresented
    vowels varying in acoustic distance from /e/.
    Results replicated suppression, with a smaller M100 during speaking than listening. Although we did
    not find unequivocal support for the first prediction, participants with less M100 suppression were
    better at the imitation task. These results are consistent with the enhancement of M100 serving as an
    error signal to drive subsequent speech-motor adaptation.
  • Guadalupe, T., Zwiers, M. P., Wittfeld, K., Teumer, A., Vasquez, A. A., Hoogman, M., Hagoort, P., Fernandez, G., Buitelaar, J., van Bokhoven, H., Hegenscheid, K., Völzke, H., Franke, B., Fisher, S. E., Grabe, H. J., & Francks, C. (2015). Asymmetry within and around the human planum temporale is sexually dimorphic and influenced by genes involved in steroid hormone receptor activity. Cortex, 62, 41-55. doi:10.1016/j.cortex.2014.07.015.

    Abstract

    The genetic determinants of cerebral asymmetries are unknown. Sex differences in asymmetry of the planum temporale, that overlaps Wernicke’s classical language area, have been inconsistently reported. Meta-analysis of previous studies has suggested that publication bias established this sex difference in the literature. Using probabilistic definitions of cortical regions we screened over the cerebral cortex for sexual dimorphisms of asymmetry in 2337 healthy subjects, and found the planum temporale to show the strongest sex-linked asymmetry of all regions, which was supported by two further datasets, and also by analysis with the Freesurfer package that performs automated parcellation of cerebral cortical regions. We performed a genome-wide association scan meta-analysis of planum temporale asymmetry in a pooled sample of 3095 subjects, followed by a candidate-driven approach which measured a significant enrichment of association in genes of the ´steroid hormone receptor activity´ and 'steroid metabolic process' pathways. Variants in the genes and pathways identified may affect the role of the planum temporale in language cognition.
  • Hagoort, P. (2015). Het talige brein. In A. Aleman, & H. E. Hulshoff Pol (Eds.), Beeldvorming van het brein: Imaging voor psychiaters en psychologen (pp. 169-176). Utrecht: De Tijdstroom.
  • Hagoort, P. (2015). Spiegelneuronen. In J. Brockmann (Ed.), Wetenschappelijk onkruid: 179 hardnekkige ideeën die vooruitgang blokkeren (pp. 455-457). Amsterdam: Maven Publishing.
  • Holler, J., Kokal, I., Toni, I., Hagoort, P., Kelly, S. D., & Ozyurek, A. (2015). Eye’m talking to you: Speakers’ gaze direction modulates co-speech gesture processing in the right MTG. Social Cognitive & Affective Neuroscience, 10, 255-261. doi:10.1093/scan/nsu047.

    Abstract

    Recipients process information from speech and co-speech gestures, but it is currently unknown how this processing is influenced by the presence of other important social cues, especially gaze direction, a marker of communicative intent. Such cues may modulate neural activity in regions associated either with the processing of ostensive cues, such as eye gaze, or with the processing of semantic information, provided by speech and gesture.
    Participants were scanned (fMRI) while taking part in triadic communication involving two recipients and a speaker. The speaker uttered sentences that
    were and were not accompanied by complementary iconic gestures. Crucially, the speaker alternated her gaze direction, thus creating two recipient roles: addressed (direct gaze) vs unaddressed (averted gaze) recipient. The comprehension of Speech&Gesture relative to SpeechOnly utterances recruited middle occipital, middle temporal and inferior frontal gyri, bilaterally. The calcarine sulcus and posterior cingulate cortex were sensitive to differences between direct and averted gaze. Most importantly, Speech&Gesture utterances, but not SpeechOnly utterances, produced additional activity in the right middle temporal gyrus when participants were addressed. Marking communicative intent with gaze direction modulates the processing of speech–gesture utterances in cerebral areas typically associated with the semantic processing of multi-modal communicative acts.
  • Kunert, R., Willems, R. M., Casasanto, D., Patel, A. D., & Hagoort, P. (2015). Music and language syntax interact in Broca’s Area: An fMRI study. PLoS One, 10(11): e0141069. doi:10.1371/journal.pone.0141069.

    Abstract

    Instrumental music and language are both syntactic systems, employing complex, hierarchically-structured sequences built using implicit structural norms. This organization allows listeners to understand the role of individual words or tones in the context of an unfolding sentence or melody. Previous studies suggest that the brain mechanisms of syntactic processing may be partly shared between music and language. However, functional neuroimaging evidence for anatomical overlap of brain activity involved in linguistic and musical syntactic processing has been lacking. In the present study we used functional magnetic resonance imaging (fMRI) in conjunction with an interference paradigm based on sung sentences. We show that the processing demands of musical syntax (harmony) and language syntax interact in Broca’s area in the left inferior frontal gyrus (without leading to music and language main effects). A language main effect in Broca’s area only emerged in the complex music harmony condition, suggesting that (with our stimuli and tasks) a language effect only becomes visible under conditions of increased demands on shared neural resources. In contrast to previous studies, our design allows us to rule out that the observed neural interaction is due to: (1) general attention mechanisms, as a psychoacoustic auditory anomaly behaved unlike the harmonic manipulation, (2) error processing, as the language and the music stimuli contained no structural errors. The current results thus suggest that two different cognitive domains—music and language—might draw on the same high level syntactic integration resources in Broca’s area.
  • Lai, V. T., Willems, R. M., & Hagoort, P. (2015). Feel between the Lines: Implied emotion from combinatorial semantics. Journal of Cognitive Neuroscience, 27(8), 1528-1541. doi:10.1162/jocn_a_00798.

    Abstract

    This study investigated the brain regions for the comprehension of implied emotion in sentences. Participants read negative sentences without negative words, for example, “The boy fell asleep and never woke up again,” and their neutral counterparts “The boy stood up and grabbed his bag.” This kind of negative sentence allows us to examine implied emotion derived at the sentence level, without associative emotion coming from word retrieval. We found that implied emotion in sentences, relative to neutral sentences, led to activation in some emotion-related areas, including the medial prefrontal cortex, the amygdala, and the insula, as well as certain language-related areas, including the inferior frontal gyrus, which has been implicated in combinatorial processing. These results suggest that the emotional network involved in implied emotion is intricately related to the network for combinatorial processing in language, supporting the view that sentence meaning is more than simply concatenating the meanings of its lexical building blocks.
  • Peeters, D., Chu, M., Holler, J., Hagoort, P., & Ozyurek, A. (2015). Electrophysiological and kinematic correlates of communicative intent in the planning and production of pointing gestures and speech. Journal of Cognitive Neuroscience, 27(12), 2352-2368. doi:10.1162/jocn_a_00865.

    Abstract

    In everyday human communication, we often express our communicative intentions by manually pointing out referents in the material world around us to an addressee, often in tight synchronization with referential speech. This study investigated whether and how the kinematic form of index finger pointing gestures is shaped by the gesturer's communicative intentions and how this is modulated by the presence of concurrently produced speech. Furthermore, we explored the neural mechanisms underpinning the planning of communicative pointing gestures and speech. Two experiments were carried out in which participants pointed at referents for an addressee while the informativeness of their gestures and speech was varied. Kinematic and electrophysiological data were recorded online. It was found that participants prolonged the duration of the stroke and poststroke hold phase of their gesture to be more communicative, in particular when the gesture was carrying the main informational burden in their multimodal utterance. Frontal and P300 effects in the ERPs suggested the importance of intentional and modality-independent attentional mechanisms during the planning phase of informative pointing gestures. These findings contribute to a better understanding of the complex interplay between action, attention, intention, and language in the production of pointing gestures, a communicative act core to human interaction.
  • Peeters, D., Hagoort, P., & Ozyurek, A. (2015). Electrophysiological evidence for the role of shared space in online comprehension of spatial demonstratives. Cognition, 136, 64-84. doi:10.1016/j.cognition.2014.10.010.

    Abstract

    A fundamental property of language is that it can be used to refer to entities in the extra-linguistic physical context of a conversation in order to establish a joint focus of attention on a referent. Typological and psycholinguistic work across a wide range of languages has put forward at least two different theoretical views on demonstrative reference. Here we contrasted and tested these two accounts by investigating the electrophysiological brain activity underlying the construction of indexical meaning in comprehension. In two EEG experiments, participants watched pictures of a speaker who referred to one of two objects using speech and an index-finger pointing gesture. In contrast with separately collected native speakers’ linguistic intuitions, N400 effects showed a preference for a proximal demonstrative when speaker and addressee were in a face-to-face orientation and all possible referents were located in the shared space between them, irrespective of the physical proximity of the referent to the speaker. These findings reject egocentric proximity-based accounts of demonstrative reference, support a sociocentric approach to deixis, suggest that interlocutors construe a shared space during conversation, and imply that the psychological proximity of a referent may be more important than its physical proximity.
  • Peeters, D., Snijders, T. M., Hagoort, P., & Ozyurek, A. (2015). The role of left inferior frontal Gyrus in the integration of point- ing gestures and speech. In G. Ferré, & M. Tutton (Eds.), Proceedings of the4th GESPIN - Gesture & Speech in Interaction Conference. Nantes: Université de Nantes.

    Abstract

    Comprehension of pointing gestures is fundamental to human communication. However, the neural mechanisms
    that subserve the integration of pointing gestures and speech in visual contexts in comprehension
    are unclear. Here we present the results of an fMRI study in which participants watched images of an
    actor pointing at an object while they listened to her referential speech. The use of a mismatch paradigm
    revealed that the semantic unication of pointing gesture and speech in a triadic context recruits left
    inferior frontal gyrus. Complementing previous ndings, this suggests that left inferior frontal gyrus
    semantically integrates information across modalities and semiotic domains.
  • Samur, D., Lai, V. T., Hagoort, P., & Willems, R. M. (2015). Emotional context modulates embodied metaphor comprehension. Neuropsychologia, 78, 108-114. doi:10.1016/j.neuropsychologia.2015.10.003.

    Abstract

    Emotions are often expressed metaphorically, and both emotion and metaphor are ways through which abstract meaning can be grounded in language. Here we investigate specifically whether motion-related verbs when used metaphorically are differentially sensitive to a preceding emotional context, as compared to when they are used in a literal manner. Participants read stories that ended with ambiguous action/motion sentences (e.g., he got it), in which the action/motion could be interpreted metaphorically (he understood the idea) or literally (he caught the ball) depending on the preceding story. Orthogonal to the metaphorical manipulation, the stories were high or low in emotional content. The results showed that emotional context modulated the neural response in visual motion areas to the metaphorical interpretation of the sentences, but not to their literal interpretations. In addition, literal interpretations of the target sentences led to stronger activation in the visual motion areas as compared to metaphorical readings of the sentences. We interpret our results as suggesting that emotional context specifically modulates mental simulation during metaphor processing
  • Simanova, I., Van Gerven, M. A., Oostenveld, R., & Hagoort, P. (2015). Predicting the semantic category of internally generated words from neuromagnetic recordings. Journal of Cognitive Neuroscience, 27(1), 35-45. doi:10.1162/jocn_a_00690.

    Abstract

    In this study, we explore the possibility to predict the semantic category of words from brain signals in a free word generation task. Participants produced single words from different semantic categories in a modified semantic fluency task. A Bayesian logistic regression classifier was trained to predict the semantic category of words from single-trial MEG data. Significant classification accuracies were achieved using sensor-level MEG time series at the time interval of conceptual preparation. Semantic category prediction was also possible using source-reconstructed time series, based on minimum norm estimates of cortical activity. Brain regions that contributed most to classification on the source level were identified. These were the left inferior frontal gyrus, left middle frontal gyrus, and left posterior middle temporal gyrus. Additionally, the temporal dynamics of brain activity underlying the semantic preparation during word generation was explored. These results provide important insights about central aspects of language production
  • Xiang, H., Van Leeuwen, T. M., Dediu, D., Roberts, L., Norris, D. G., & Hagoort, P. (2015). L2-proficiency-dependent laterality shift in structural connectivity of brain language pathways. Brain Connectivity, 5(6), 349-361. doi:10.1089/brain.2013.0199.

    Abstract

    Diffusion tensor imaging (DTI) and a longitudinal language learning approach were applied to investigate the relationship between the achieved second language (L2) proficiency during L2 learning and the reorganization of structural connectivity between core language areas. Language proficiency tests and DTI scans were obtained from German students before and after they completed an intensive 6-week course of the Dutch language. In the initial learning stage, with increasing L2 proficiency, the hemispheric dominance of the BA6-temporal pathway (mainly along the arcuate fasciculus) shifted from the left to the right hemisphere. With further increased proficiency, however, lateralization dominance was again found in the left BA6-temporal pathway. This result is consistent with reports in the literature that imply a stronger involvement of the right hemisphere in L2-processing especially for less proficient L2-speakers. This is the first time that a L2-proficiency-dependent laterality shift in structural connectivity of language pathways during L2 acquisition has been observed to shift from left to right, and back to left hemisphere dominance with increasing L2-proficiency. We additionally find that changes in fractional anisotropy values after the course are related to the time elapsed between the two scans. The results suggest that structural connectivity in (at least part of) the perisylvian language network may be subject to fast dynamic changes following language learning
  • Brouwer, G. J., Tong, F., Hagoort, P., & Van Ee, R. (2009). Perceptual incongruence influences bistability and cortical activation. Plos One, 4(3): e5056. doi:10.1371/journal.pone.0005056.

    Abstract

    We employed a parametric psychophysical design in combination with functional imaging to examine the influence of metric changes in perceptual incongruence on perceptual alternation rates and cortical responses. Subjects viewed a bistable stimulus defined by incongruent depth cues; bistability resulted from incongruence between binocular disparity and monocular perspective cues that specify different slants (slant rivalry). Psychophysical results revealed that perceptual alternation rates were positively correlated with the degree of perceived incongruence. Functional imaging revealed systematic increases in activity that paralleled the psychophysical results within anterior intraparietal sulcus, prior to the onset of perceptual alternations. We suggest that this cortical activity predicts the frequency of subsequent alternations, implying a putative causal role for these areas in initiating bistable perception. In contrast, areas implicated in form and depth processing (LOC and V3A) were sensitive to the degree of slant, but failed to show increases in activity when these cues were in conflict.
  • Casasanto, D., Willems, R. M., & Hagoort, P. (2009). Body-specific representations of action verbs: Evidence from fMRI in right- and left-handers. In N. Taatgen, & H. Van Rijn (Eds.), Proceedings of the 31st Annual Meeting of the Cognitive Science Society (pp. 875-880). Austin: Cognitive Science Society.

    Abstract

    According to theories of embodied cognition, understanding a verb like throw involves unconsciously simulating the action throwing, using areas of the brain that support motor planning. If understanding action words involves mentally simulating our own actions, then the neurocognitive representation of word meanings should differ for people with different kinds of bodies, who perform actions in systematically different ways. In a test of the body-specificity hypothesis (Casasanto, 2009), we used fMRI to compare premotor activity correlated with action verb understanding in right- and left-handers. Right-handers preferentially activated left premotor cortex during lexical decision on manual action verbs (compared with non-manual action verbs), whereas left-handers preferentially activated right premotor areas. This finding helps refine theories of embodied semantics, suggesting that implicit mental simulation during language processing is body-specific: Right and left-handers, who perform actions differently, use correspondingly different areas of the brain for representing action verb meanings.
  • Fedor, A., Pléh, C., Brauer, J., Caplan, D., Friederici, A. D., Gulyás, B., Hagoort, P., Nazir, T., & Singer, W. (2009). What are the brain mechanisms underlying syntactic operations? In D. Bickerton, & E. Szathmáry (Eds.), Biological foundations and origin of syntax (pp. 299-324). Cambridge, MA: MIT Press.

    Abstract

    This chapter summarizes the extensive discussions that took place during the Forum as well as the subsequent months thereafter. It assesses current understanding of the neuronal mechanisms that underlie syntactic structure and processing.... It is posited that to understand the neurobiology of syntax, it might be worthwhile to shift the balance from comprehension to syntactic encoding in language production
  • Folia, V., Forkstam, C., Hagoort, P., & Petersson, K. M. (2009). Language comprehension: The interplay between form and content. In N. Taatgen, & H. van Rijn (Eds.), Proceedings of the 31th Annual Conference of the Cognitive Science Society (pp. 1686-1691). Austin, TX: Cognitive Science Society.

    Abstract

    In a 2x2 event-related FMRI study we find support for the idea that the inferior frontal cortex, centered on Broca’s region and its homologue, is involved in constructive unification operations during the structure-building process in parsing for comprehension. Tentatively, we provide evidence for a role of the dorsolateral prefrontal cortex centered on BA 9/46 in the control component of the language system. Finally, the left temporo-parietal cortex, in the vicinity of Wernicke’s region, supports the interaction between the syntax of gender agreement and sentence-level semantics.
  • Hagoort, P. (2009). The fractionation of spoken language understanding by measuring electrical and magnetic brain signals. In B. C. J. Moore, L. K. Tyler, & W. Marslen-Wilson (Eds.), The perception of speech: From sound to meaning (pp. 223-248). New York: Oxford University Press.
  • Hagoort, P. (2009). Reflections on the neurobiology of syntax. In D. Bickerton, & E. Szathmáry (Eds.), Biological foundations and origin of syntax (pp. 279-296). Cambridge, MA: MIT Press.

    Abstract

    This contribution focuses on the neural infrastructure for parsing and syntactic encoding. From an anatomical point of view, it is argued that Broca's area is an ill-conceived notion. Functionally, Broca's area and adjacent cortex (together Broca's complex) are relevant for language, but not exclusively for this domain of cognition. Its role can be characterized as providing the necessary infrastructure for unification (syntactic and semantic). A general proposal, but with required level of computational detail, is discussed to account for the distribution of labor between different components of the language network in the brain.Arguments are provided for the immediacy principle, which denies a privileged status for syntax in sentence processing. The temporal profile of event-related brain potential (ERP) is suggested to require predictive processing. Finally, since, next to speed, diversity is a hallmark of human languages, the language readiness of the brain might not depend on a universal, dedicated neural machinery for syntax, but rather on a shaping of the neural infrastructure of more general cognitive systems (e.g., memory, unification) in a direction that made it optimally suited for the purpose of communication through language.
  • Hagoort, P., Baggio, G., & Willems, R. M. (2009). Semantic unification. In M. S. Gazzaniga (Ed.), The cognitive neurosciences, 4th ed. (pp. 819-836). Cambridge, MA: MIT Press.

    Abstract

    Language and communication are about the exchange of meaning. A key feature of understanding and producing language is the construction of complex meaning from more elementary semantic building blocks. The functional characteristics of this semantic unification process are revealed by studies using event related brain potentials. These studies have found that word meaning is assembled into compound meaning in not more than 500 ms. World knowledge, information about the speaker, co-occurring visual input and discourse all have an immediate impact on semantic unification, and trigger similar electrophysiological responses as sentence-internal semantic information. Neuroimaging studies show that a network of brain areas, including the left inferior frontal gyrus, the left superior/middle temporal cortex, the left inferior parietal cortex and, to a lesser extent their right hemisphere homologues are recruited to perform semantic unification.
  • Hagoort, P. (2009). Taalontwikkeling: Meer dan woorden alleen. In M. Evenblij (Ed.), Brein in beeld: Beeldvorming bij heersenonderzoek (pp. 53-57). Den Haag: Stichting Bio-Wetenschappen en Maatschappij.
  • Hagoort, P., & Levelt, W. J. M. (2009). The speaking brain. Science, 326(5951), 372-373. doi:10.1126/science.1181675.

    Abstract

    How does intention to speak become the action of speaking? It involves the generation of a preverbal message that is tailored to the requirements of a particular language, and through a series of steps, the message is transformed into a linear sequence of speech sounds (1, 2). These steps include retrieving different kinds of information from memory (semantic, syntactic, and phonological), and combining them into larger structures, a process called unification. Despite general agreement about the steps that connect intention to articulation, there is no consensus about their temporal profile or the role of feedback from later steps (3, 4). In addition, since the discovery by the French physician Pierre Paul Broca (in 1865) of the role of the left inferior frontal cortex in speaking, relatively little progress has been made in understanding the neural infrastructure that supports speech production (5). One reason is that the characteristics of natural language are uniquely human, and thus the neurobiology of language lacks an adequate animal model. But on page 445 of this issue, Sahin et al. (6) demonstrate, by recording neuronal activity in the human brain, that different kinds of linguistic information are indeed sequentially processed within Broca's area.
  • Kooijman, V., Hagoort, P., & Cutler, A. (2009). Prosodic structure in early word segmentation: ERP evidence from Dutch ten-month-olds. Infancy, 14, 591 -612. doi:10.1080/15250000903263957.

    Abstract

    Recognizing word boundaries in continuous speech requires detailed knowledge of the native language. In the first year of life, infants acquire considerable word segmentation abilities. Infants at this early stage in word segmentation rely to a large extent on the metrical pattern of their native language, at least in stress-based languages. In Dutch and English (both languages with a preferred trochaic stress pattern), segmentation of strong-weak words develops rapidly between 7 and 10 months of age. Nevertheless, trochaic languages contain not only strong-weak words but also words with a weak-strong stress pattern. In this article, we present electrophysiological evidence of the beginnings of weak-strong word segmentation in Dutch 10-month-olds. At this age, the ability to combine different cues for efficient word segmentation does not yet seem to be completely developed. We provide evidence that Dutch infants still largely rely on strong syllables, even for the segmentation of weak-strong words.
  • Koten Jr., J. W., Wood, G., Hagoort, P., Goebel, R., Propping, P., Willmes, K., & Boomsma, D. I. (2009). Genetic contribution to variation in cognitive function: An fMRI study in twins. Science, 323(5922), 1737-1740. doi:10.1126/science.1167371.

    Abstract

    Little is known about the genetic contribution to individual differences in neural networks subserving cognition function. In this functional magnetic resonance imaging (fMRI) twin study, we found a significant genetic influence on brain activation in neural networks supporting digit working memory tasks. Participants activating frontal-parietal networks responded faster than individuals relying more on language-related brain networks.There were genetic influences on brain activation in language-relevant brain circuits that were atypical for numerical working memory tasks as such. This suggests that differences in cognition might be related to brain activation patterns that differ qualitatively among individuals.
  • De Lange, F. P., Koers, A., Kalkman, J. S., Bleijenberg, G., Hagoort, P., Van der Meer, J. W. M., & Toni, I. (2009). Reply to: "Can CBT substantially change grey matter volume in chronic fatigue syndrome" [Letter to the editor]. Brain, 132(6), e111. doi:10.1093/brain/awn208.
  • De Lange, F., Bleijenberg, G., Van der Meer, J. W. M., Hagoort, P., & Toni, I. (2009). Reply: Change in grey matter volume cannot be assumed to be due to cognitive behavioural therapy [Letter to the editor]. Brain, 132(7), e120. doi:10.1093/brain/awn359.
  • De Lange, F. P., Knoop, H., Bleijenberg, G., Van der Meer, J. W. M., Hagoort, P., & Toni, I. (2009). The experience of fatigue in the brain [Letter to the editor]. Psychological Medicine, 39, 523-524. doi:10.1017/S0033291708004844.
  • Menenti, L., Petersson, K. M., Scheeringa, R., & Hagoort, P. (2009). When elephants fly: Differential sensitivity of right and left inferior frontal gyri to discourse and world knowledge. Journal of Cognitive Neuroscience, 21, 2358-2368. doi:10.1162/jocn.2008.21163.

    Abstract

    Both local discourse and world knowledge are known to influence sentence processing. We investigated how these two sources of information conspire in language comprehension. Two types of critical sentences, correct and world knowledge anomalies, were preceded by either a neutral or a local context. The latter made the world knowledge anomalies more acceptable or plausible. We predicted that the effect of world knowledge anomalies would be weaker for the local context. World knowledge effects have previously been observed in the left inferior frontal region (Brodmann's area 45/47). In the current study, an effect of world knowledge was present in this region in the neutral context. We also observed an effect in the right inferior frontal gyrus, which was more sensitive to the discourse manipulation than the left inferior frontal gyrus. In addition, the left angular gyrus reacted strongly to the degree of discourse coherence between the context and critical sentence. Overall, both world knowledge and the discourse context affect the process of meaning unification, but do so by recruiting partly different sets of brain areas.
  • Newman-Norlund, S. E., Noordzij, M. L., Newman-Norlund, R. D., Volman, I. A., De Ruiter, J. P., Hagoort, P., & Toni, I. (2009). Recipient design in tacit communication. Cognition, 111, 46-54. doi:10.1016/j.cognition.2008.12.004.

    Abstract

    The ability to design tailored messages for specific listeners is an important aspect of
    human communication. The present study investigates whether a mere belief about an
    addressee’s identity influences the generation and production of a communicative message in
    a novel, non-verbal communication task. Participants were made to believe they were playing a game with a child or an adult partner, while a confederate acted as both child
    and adult partners with matched performance and response times. The participants’ belief
    influenced their behavior, spending longer when interacting with the presumed child
    addressee, but only during communicative portions of the game, i.e. using time as a tool
    to place emphasis on target information. This communicative adaptation attenuated with
    experience, and it was related to personality traits, namely Empathy and Need for Cognition
    measures. Overall, these findings indicate that novel nonverbal communicative interactions
    are selected according to a socio-centric perspective, and they are strongly
    influenced by participants’ traits.
  • Noordzij, M., Newman-Norlund, S. E., De Ruiter, J. P., Hagoort, P., Levinson, S. C., & Toni, I. (2009). Brain mechanisms underlying human communication. Frontiers in Human Neuroscience, 3:14. doi:10.3389/neuro.09.014.2009.

    Abstract

    Human communication has been described as involving the coding-decoding of a conventional symbol system, which could be supported by parts of the human motor system (i.e. the “mirror neurons system”). However, this view does not explain how these conventions could develop in the first place. Here we target the neglected but crucial issue of how people organize their non-verbal behavior to communicate a given intention without pre-established conventions. We have measured behavioral and brain responses in pairs of subjects during communicative exchanges occurring in a real, interactive, on-line social context. In two fMRI studies, we found robust evidence that planning new communicative actions (by a sender) and recognizing the communicative intention of the same actions (by a receiver) relied on spatially overlapping portions of their brains (the right posterior superior temporal sulcus). The response of this region was lateralized to the right hemisphere, modulated by the ambiguity in meaning of the communicative acts, but not by their sensorimotor complexity. These results indicate that the sender of a communicative signal uses his own intention recognition system to make a prediction of the intention recognition performed by the receiver. This finding supports the notion that our communicative abilities are distinct from both sensorimotor processes and language abilities.
  • Pijnacker, J., Geurts, B., Van Lambalgen, M., Kan, C. C., Buitelaar, J. K., & Hagoort, P. (2009). Defeasible reasoning in high-functioning adults with autism: Evidence for impaired exception-handling. Neuropsychologia, 47, 644-651. doi:10.1016/j.neuropsychologia.2008.11.011.

    Abstract

    While autism is one of the most intensively researched psychiatric disorders, little is known about reasoning skills of people with autism. The focus of this study was on defeasible inferences, that is inferences that can be revised in the light of new information. We used a behavioral task to investigate (a) conditional reasoning and (b) the suppression of conditional inferences in high-functioning adults with autism. In the suppression task a possible exception was made salient which could prevent a conclusion from being drawn. We predicted that the autism group would have difficulties dealing with such exceptions because they require mental flexibility to adjust to the context, which is often impaired in autism. The findings confirm our hypothesis that high-functioning adults with autism have a specific difficulty with exception-handling during reasoning. It is suggested that defeasible reasoning is also involved in other cognitive domains. Implications for neural underpinnings of reasoning and autism are discussed.
  • Pijnacker, J., Hagoort, P., Buitelaar, J., Teunisse, J.-P., & Geurts, B. (2009). Pragmatic inferences in high-functioning adults with autism and Asperger syndrome. Journal of Autism and Developmental Disorders, 39(4), 607-618. doi:10.1007/s10803-008-0661-8.

    Abstract

    Although people with autism spectrum disorders (ASD) often have severe problems with pragmatic aspects of language, little is known about their pragmatic reasoning. We carried out a behavioral study on highfunctioning adults with autistic disorder (n = 11) and Asperger syndrome (n = 17) and matched controls (n = 28) to investigate whether they are capable of deriving scalar implicatures, which are generally considered to be pragmatic inferences. Participants were presented with underinformative sentences like ‘‘Some sparrows are birds’’. This sentence is logically true, but pragmatically inappropriate if the scalar implicature ‘‘Not all sparrows are birds’’ is derived. The present findings indicate that the combined ASD group was just as likely as controls to derive scalar implicatures, yet there was a difference between participants with autistic disorder and Asperger syndrome, suggesting a potential differentiation between these disorders in pragmatic reasoning. Moreover, our results suggest that verbal intelligence is a constraint for task performance in autistic disorder but not in Asperger syndrome.
  • Scheeringa, R., Petersson, K. M., Oostenveld, R., Norris, D. G., Hagoort, P., & Bastiaansen, M. C. M. (2009). Trial-by-trial coupling between EEG and BOLD identifies networks related to alpha and theta EEG power increases during working memory maintenance. Neuroimage, 44, 1224-1238. doi:10.1016/j.neuroimage.2008.08.041.

    Abstract

    PET and fMRI experiments have previously shown that several brain regions in the frontal and parietal lobe are involved in working memory maintenance. MEG and EEG experiments have shown parametric increases with load for oscillatory activity in posterior alpha and frontal theta power. In the current study we investigated whether the areas found with fMRI can be associated with these alpha and theta effects by measuring simultaneous EEG and fMRI during a modified Sternberg task This allowed us to correlate EEG at the single trial level with the fMRI BOLD signal by forming a regressor based on single trial alpha and theta
    power estimates. We observed a right posterior, parametric alpha power increase, which was functionally related to decreases in BOLD in the primary visual cortex and in the posterior part of the right middle temporal gyrus. We relate this finding to the inhibition of neuronal activity that may interfere with WM maintenance. An observed parametric increase in frontal theta power was correlated to a decrease in BOLD in
    regions that together form the default mode network. We did not observe correlations between oscillatory EEG phenomena and BOLD in the traditional WM areas. In conclusion, the study shows that simultaneous EEG fMRI recordings can be successfully used to identify the emergence of functional networks in the brain during the execution of a cognitive task.
  • Snijders, T. M., Vosse, T., Kempen, G., Van Berkum, J. J. A., Petersson, K. M., & Hagoort, P. (2009). Retrieval and unification of syntactic structure in sentence comprehension: An fMRI study using word-category ambiguity. Cerebral Cortex, 19, 1493-1503. doi:10.1093/cercor/bhn187.

    Abstract

    Sentence comprehension requires the retrieval of single word information from long-term memory, and the integration of this information into multiword representations. The current functional magnetic resonance imaging study explored the hypothesis that the left posterior temporal gyrus supports the retrieval of lexical-syntactic information, whereas left inferior frontal gyrus (LIFG) contributes to syntactic unification. Twenty-eight subjects read sentences and word sequences containing word-category (noun–verb) ambiguous words at critical positions. Regions contributing to the syntactic unification process should show enhanced activation for sentences compared to words, and only within sentences display a larger signal for ambiguous than unambiguous conditions. The posterior LIFG showed exactly this predicted pattern, confirming our hypothesis that LIFG contributes to syntactic unification. The left posterior middle temporal gyrus was activated more for ambiguous than unambiguous conditions (main effect over both sentences and word sequences), as predicted for regions subserving the retrieval of lexical-syntactic information from memory. We conclude that understanding language involves the dynamic interplay between left inferior frontal and left posterior temporal regions.

    Additional information

    suppl1.pdf suppl2_dutch_stimulus.pdf
  • Tesink, C. M. J. Y., Buitelaar, J. K., Petersson, K. M., Van der Gaag, R. J., Kan, C. C., Tendolkar, I., & Hagoort, P. (2009). Neural correlates of pragmatic language comprehension in autism disorders. Brain, 132, 1941-1952. doi:10.1093/brain/awp103.

    Abstract

    Difficulties with pragmatic aspects of communication are universal across individuals with autism spectrum disorders (ASDs). Here we focused on an aspect of pragmatic language comprehension that is relevant to social interaction in daily life: the integration of speaker characteristics inferred from the voice with the content of a message. Using functional magnetic resonance imaging (fMRI), we examined the neural correlates of the integration of voice-based inferences about the speaker’s age, gender or social background, and sentence content in adults with ASD and matched control participants. Relative to the control group, the ASD group showed increased activation in right inferior frontal gyrus (RIFG; Brodmann area 47) for speakerincongruent sentences compared to speaker-congruent sentences. Given that both groups performed behaviourally at a similar level on a debriefing interview outside the scanner, the increased activation in RIFG for the ASD group was interpreted as being compensatory in nature. It presumably reflects spill-over processing from the language dominant left hemisphere due to higher task demands faced by the participants with ASD when integrating speaker characteristics and the content of a spoken sentence. Furthermore, only the control group showed decreased activation for speaker-incongruent relative to speaker-congruent sentences in right ventral medial prefrontal cortex (vMPFC; Brodmann area 10), including right anterior cingulate cortex (ACC; Brodmann area 24/32). Since vMPFC is involved in self-referential processing related to judgments and inferences about self and others, the absence of such a modulation in vMPFC activation in the ASD group possibly points to atypical default self-referential mental activity in ASD. Our results show that in ASD compensatory mechanisms are necessary in implicit, low-level inferential processes in spoken language understanding. This indicates that pragmatic language problems in ASD are not restricted to high-level inferential processes, but encompass the most basic aspects of pragmatic language processing.
  • Tesink, C. M. J. Y., Petersson, K. M., Van Berkum, J. J. A., Van den Brink, D., Buitelaar, J. K., & Hagoort, P. (2009). Unification of speaker and meaning in language comprehension: An fMRI study. Journal of Cognitive Neuroscience, 21, 2085-2099. doi:10.1162/jocn.2008.21161.

    Abstract

    When interpreting a message, a listener takes into account several sources of linguistic and extralinguistic information. Here we focused on one particular form of extralinguistic information, certain speaker characteristics as conveyed by the voice. Using functional magnetic resonance imaging, we examined the neural structures involved in the unification of sentence meaning and voice-based inferences about the speaker's age, sex, or social background. We found enhanced activation in the inferior frontal gyrus bilaterally (BA 45/47) during listening to sentences whose meaning was incongruent with inferred speaker characteristics. Furthermore, our results showed an overlap in brain regions involved in unification of speaker-related information and those used for the unification of semantic and world knowledge information [inferior frontal gyrus bilaterally (BA 45/47) and left middle temporal gyrus (BA 21)]. These findings provide evidence for a shared neural unification system for linguistic and extralinguistic sources of information and extend the existing knowledge about the role of inferior frontal cortex as a crucial component for unification during language comprehension.
  • Uddén, J., Araújo, S., Forkstam, C., Ingvar, M., Hagoort, P., & Petersson, K. M. (2009). A matter of time: Implicit acquisition of recursive sequence structures. In N. Taatgen, & H. Van Rijn (Eds.), Proceedings of the Thirty-First Annual Conference of the Cognitive Science Society (pp. 2444-2449).

    Abstract

    A dominant hypothesis in empirical research on the evolution of language is the following: the fundamental difference between animal and human communication systems is captured by the distinction between regular and more complex non-regular grammars. Studies reporting successful artificial grammar learning of nested recursive structures and imaging studies of the same have methodological shortcomings since they typically allow explicit problem solving strategies and this has been shown to account for the learning effect in subsequent behavioral studies. The present study overcomes these shortcomings by using subtle violations of agreement structure in a preference classification task. In contrast to the studies conducted so far, we use an implicit learning paradigm, allowing the time needed for both abstraction processes and consolidation to take place. Our results demonstrate robust implicit learning of recursively embedded structures (context-free grammar) and recursive structures with cross-dependencies (context-sensitive grammar) in an artificial grammar learning task spanning 9 days. Keywords: Implicit artificial grammar learning; centre embedded; cross-dependency; implicit learning; context-sensitive grammar; context-free grammar; regular grammar; non-regular grammar
  • Wang, L., Hagoort, P., & Yang, Y. (2009). Semantic illusion depends on information structure: ERP evidence. Brain Research, 1282, 50-56. doi:10.1016/j.brainres.2009.05.069.

    Abstract

    Next to propositional content, speakers distribute information in their utterances in such a way that listeners can make a distinction between new (focused) and given (non-focused) information. This is referred to as information structure. We measured event-related potentials (ERPs) to explore the role of information structure in semantic processing. Following different questions in wh-question-answer pairs (e.g. What kind of vegetable did Ming buy for cooking today? /Who bought the vegetables for cooking today?), the answer sentences (e.g., Ming bought eggplant/beef to cook today.) contained a critical word, which was either semantically appropriate (eggplant) or inappropriate (beef), and either focus or non-focus. The results showed a full N400 effect only when the critical words were in focus position. In non-focus position a strongly reduced N400 effect was observed, in line with the well-known semantic illusion effect. The results suggest that information structure facilitates semantic processing by devoting more resources to focused information.
  • Willems, R. M., Toni, I., Hagoort, P., & Casasanto, D. (2009). Body-specific motor imagery of hand actions: Neural evidence from right- and left-handers. Frontiers in Human Neuroscience, 3: 39, pp. 39. doi:10.3389/neuro.09.039.2009.

    Abstract

    If motor imagery uses neural structures involved in action execution, then the neural correlates of imagining an action should differ between individuals who tend to execute the action differently. Here we report fMRI data showing that motor imagery is influenced by the way people habitually perform motor actions with their particular bodies; that is, motor imagery is ‘body-specific’ (Casasanto, 2009). During mental imagery for complex hand actions, activation of cortical areas involved in motor planning and execution was left-lateralized in right-handers but right-lateralized in left-handers. We conclude that motor imagery involves the generation of an action plan that is grounded in the participant’s motor habits, not just an abstract representation at the level of the action’s goal. People with different patterns of motor experience form correspondingly different neurocognitive representations of imagined actions.
  • Willems, R. M., & Hagoort, P. (2009). Broca's region: Battles are not won by ignoring half of the facts. Trends in Cognitive Sciences, 13(3), 101. doi:10.1016/j.tics.2008.12.001.
  • Willems, R. M., Ozyurek, A., & Hagoort, P. (2009). Differential roles for left inferior frontal and superior temporal cortex in multimodal integration of action and language. Neuroimage, 47, 1992-2004. doi:10.1016/j.neuroimage.2009.05.066.

    Abstract

    Several studies indicate that both posterior superior temporal sulcus/middle temporal gyrus (pSTS/MTG) and left inferior frontal gyrus (LIFG) are involved in integrating information from different modalities. Here we investigated the respective roles of these two areas in integration of action and language information. We exploited the fact that the semantic relationship between language and different forms of action (i.e. co-speech gestures and pantomimes) is radically different. Speech and co-speech gestures are always produced together, and gestures are not unambiguously understood without speech. On the contrary, pantomimes are not necessarily produced together with speech and can be easily understood without speech. We presented speech together with these two types of communicative hand actions in matching or mismatching combinations to manipulate semantic integration load. Left and right pSTS/MTG were only involved in semantic integration of speech and pantomimes. Left IFG on the other hand was involved in integration of speech and co-speech gestures as well as of speech and pantomimes. Effective connectivity analyses showed that depending upon the semantic relationship between language and action, LIFG modulates activation levels in left pSTS.

    This suggests that integration in pSTS/MTG involves the matching of two input streams for which there is a relatively stable common object representation, whereas integration in LIFG is better characterized as the on-line construction of a new and unified representation of the input streams. In conclusion, pSTS/MTG and LIFG are differentially involved in multimodal integration, crucially depending upon the semantic relationship between the input streams.

    Additional information

    Supplementary table S1
  • Willems, R. M., & Hagoort, P. (2009). Hand preference influences neural correlates of action observation. Brain Research, 1269, 90-104. doi:10.1016/j.brainres.2009.02.057.

    Abstract

    It has been argued that we map observed actions onto our own motor system. Here we added to this issue by investigating whether hand preference influences the neural correlates of action observation of simple, essentially meaningless hand actions. Such an influence would argue for an intricate neural coupling between action production and action observation, which goes beyond effects of motor repertoire or explicit motor training, as has been suggested before. Indeed, parts of the human motor system exhibited a close coupling between action production and action observation. Ventral premotor and inferior and superior parietal cortices showed differential activation for left- and right-handers that was similar during action production as well as during action observation. This suggests that mapping observed actions onto the observer's own motor system is a core feature of action observation - at least for actions that do not have a clear goal or meaning. Basic differences in the way we act upon the world are not only reflected in neural correlates of action production, but can also influence the brain basis of action observation.
  • Baggio, G., Van Lambalgen, M., & Hagoort, P. (2008). Computing and recomputing discourse models: An ERP study. Journal of Memory and Language, 59, 36-53. doi:10.1016/j.jml.2008.02.005.

    Abstract

    While syntactic reanalysis has been extensively investigated in psycholinguistics, comparatively little is known about reanalysis in the semantic domain. We used event-related brain potentials (ERPs) to keep track of semantic processes involved in understanding short narratives such as ‘The girl was writing a letter when her friend spilled coffee on the paper’. We hypothesize that these sentences are interpreted in two steps: (1) when the progressive clause is processed, a discourse model is computed in which the goal state (a complete letter) is predicted to hold; (2) when the subordinate clause is processed, the initial representation is recomputed to the effect that, in the final discourse structure, the goal state is not satisfied. Critical sentences evoked larger sustained anterior negativities (SANs) compared to controls, starting around 400 ms following the onset of the sentence-final word, and lasting for about 400 ms. The amplitude of the SAN was correlated with the frequency with which participants, in an offline probe-selection task, responded that the goal state was not attained. Our results raise the possibility that the brain supports some form of non-monotonic recomputation to integrate information which invalidates previously held assumptions.
  • Bastiaansen, M. C. M., Oostenveld, R., Jensen, O., & Hagoort, P. (2008). I see what you mean: Theta power increases are involved in the retrieval of lexical semantic information. Brain and Language, 106(1), 15-28. doi:10.1016/j.bandl.2007.10.006.

    Abstract

    An influential hypothesis regarding the neural basis of the mental lexicon is that semantic representations are neurally implemented as distributed networks carrying sensory, motor and/or more abstract functional information. This work investigates whether the semantic properties of words partly determine the topography of such networks. Subjects performed a visual lexical decision task while their EEG was recorded. We compared the EEG responses to nouns with either visual semantic properties (VIS, referring to colors and shapes) or with auditory semantic properties (AUD, referring to sounds). A time–frequency analysis of the EEG revealed power increases in the theta (4–7 Hz) and lower-beta (13–18 Hz) frequency bands, and an early power increase and subsequent decrease for the alpha (8–12 Hz) band. In the theta band we observed a double dissociation: temporal electrodes showed larger theta power increases in the AUD condition, while occipital leads showed larger theta responses in the VIS condition. The results support the notion that semantic representations are stored in functional networks with a topography that reflects the semantic properties of the stored items, and provide further evidence that oscillatory brain dynamics in the theta frequency range are functionally related to the retrieval of lexical semantic information.
  • Folia, V., Uddén, J., Forkstam, C., Ingvar, M., Hagoort, P., & Petersson, K. M. (2008). Implicit learning and dyslexia. Annals of the New York Academy of Sciences, 1145, 132-150. doi:10.1196/annals.1416.012.

    Abstract

    Several studies have reported an association between dyslexia and implicit learning deficits. It has been suggested that the weakness in implicit learning observed in dyslexic individuals may be related to sequential processing and implicit sequence learning. In the present article, we review the current literature on implicit learning and dyslexia. We describe a novel, forced-choice structural "mere exposure" artificial grammar learning paradigm and characterize this paradigm in normal readers in relation to the standard grammaticality classification paradigm. We argue that preference classification is a more optimal measure of the outcome of implicit acquisition since in the preference version participants are kept completely unaware of the underlying generative mechanism, while in the grammaticality version, the subjects have, at least in principle, been informed about the existence of an underlying complex set of rules at the point of classification (but not during acquisition). On the basis of the "mere exposure effect," we tested the prediction that the development of preference will correlate with the grammaticality status of the classification items. In addition, we examined the effects of grammaticality (grammatical/nongrammatical) and associative chunk strength (ACS; high/low) on the classification tasks (preference/grammaticality). Using a balanced ACS design in which the factors of grammaticality (grammatical/nongrammatical) and ACS (high/low) were independently controlled in a 2 × 2 factorial design, we confirmed our predictions. We discuss the suitability of this task for further investigation of the implicit learning characteristics in dyslexia.
  • Hagoort, P. (2008). Should psychology ignore the language of the brain? Current Directions in Psychological Science, 17(2), 96-101. doi:10.1111/j.1467-8721.2008.00556.x.

    Abstract

    Claims that neuroscientific data do not contribute to our understanding of psychological functions have been made recently. Here I argue that these criticisms are solely based on an analysis of functional magnetic resonance imaging (fMRI) studies. However, fMRI is only one of the methods in the toolkit of cognitive neuroscience. I provide examples from research on event-related brain potentials (ERPs) that have contributed to our understanding of the cognitive architecture of human language functions. In addition, I provide evidence of (possible) contributions from fMRI measurements to our understanding of the functional architecture of language processing. Finally, I argue that a neurobiology of human language that integrates information about the necessary genetic and neural infrastructures will allow us to answer certain questions that are not answerable if all we have is evidence from behavior.
  • Hagoort, P. (2008). The fractionation of spoken language understanding by measuring electrical and magnetic brain signals. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 363, 1055-1069. doi:10.1098/rstb.2007.2159.

    Abstract

    This paper focuses on what electrical and magnetic recordings of human brain activity reveal about spoken language understanding. Based on the high temporal resolution of these recordings, a fine-grained temporal profile of different aspects of spoken language comprehension can be obtained. Crucial aspects of speech comprehension are lexical access, selection and semantic integration. Results show that for words spoken in context, there is no ‘magic moment’ when lexical selection ends and semantic integration begins. Irrespective of whether words have early or late recognition points, semantic integration processing is initiated before words can be identified on the basis of the acoustic information alone. Moreover, for one particular event-related brain potential (ERP) component (the N400), equivalent impact of sentence- and discourse-semantic contexts is observed. This indicates that in comprehension, a spoken word is immediately evaluated relative to the widest interpretive domain available. In addition, this happens very quickly. Findings are discussed that show that often an unfolding word can be mapped onto discourse-level representations well before the end of the word. Overall, the time course of the ERP effects is compatible with the view that the different information types (lexical, syntactic, phonological, pragmatic) are processed in parallel and influence the interpretation process incrementally, that is as soon as the relevant pieces of information are available. This is referred to as the immediacy principle.
  • Li, X., Hagoort, P., & Yang, Y. (2008). Event-related potential evidence on the influence of accentuation in spoken discourse comprehension in Chinese. Journal of Cognitive Neuroscience, 20(5), 906-915. doi:10.1162/jocn.2008.20512.

    Abstract

    In an event-related potential experiment with Chinese discourses as material, we investigated how and when accentuation influences spoken discourse comprehension in relation to the different information states of the critical words. These words could either provide new or old information. It was shown that variation of accentuation influenced the amplitude of the N400, with a larger amplitude for accented than deaccented words. In addition, there was an interaction between accentuation and information state. The N400 amplitude difference between accented and deaccented new information was smaller than that between accented and deaccented old information. The results demonstrate that, during spoken discourse comprehension, listeners rapidly extract the semantic consequences of accentuation in relation to the previous discourse context. Moreover, our results show that the N400 amplitude can be larger for correct (new,accented words) than incorrect (new, deaccented words) information. This, we argue, proves that the N400 does not react to semantic anomaly per se, but rather to semantic integration load, which is higher for new information.
  • Hagoort, P., Ramsey, N. F., & Jensen, O. (2008). De gereedschapskist van de cognitieve neurowetenschap. In F. Wijnen, & F. Verstraten (Eds.), Het brein te kijk: Verkenning van de cognitieve neurowetenschap (pp. 41-75). Amsterdam: Harcourt Assessment.
  • Hagoort, P. (2008). Mijn omweg naar de filosofie. Algemeen Nederlands Tijdschrift voor Wijsbegeerte, 100(4), 303-310.
  • Hagoort, P. (2008). Über Broca, Gehirn und Bindung. In Jahrbuch 2008: Tätigkeitsberichte der Institute. München: Generalverwaltung der Max-Planck-Gesellschaft. Retrieved from http://www.mpg.de/306524/forschungsSchwerpunkt1?c=166434.

    Abstract

    Beim Sprechen und beim Sprachverstehen findet man die Wortbedeutung im Gedächtnis auf und kombiniert sie zu größeren Einheiten (Unifikation). Solche Unifikations-Operationen laufen auf unterschiedlichen Ebenen der Sprachverarbeitung ab. In diesem Beitrag wird ein Rahmen vorgeschlagen, in dem psycholinguistische Modelle mit neurobiologischer Sprachbetrachtung in Verbindung gebracht werden. Diesem Vorschlag zufolge spielt der linke inferiore frontale Gyrus (LIFG) eine bedeutende Rolle bei der Unifi kation
  • Kho, K. H., Indefrey, P., Hagoort, P., Van Veelen, C. W. M., Van Rijen, P. C., & Ramsey, N. F. (2008). Unimpaired sentence comprehension after anterior temporal cortex resection. Neuropsychologia, 46(4), 1170-1178. doi:10.1016/j.neuropsychologia.2007.10.014.

    Abstract

    Functional imaging studies have demonstrated involvement of the anterior temporal cortex in sentence comprehension. It is unclear, however, whether the anterior temporal cortex is essential for this function.We studied two aspects of sentence comprehension, namely syntactic and prosodic comprehension in temporal lobe epilepsy patients who were candidates for resection of the anterior temporal lobe. Methods: Temporal lobe epilepsy patients (n = 32) with normal (left) language dominance were tested on syntactic and prosodic comprehension before and after removal of the anterior temporal cortex. The prosodic comprehension test was also compared with performance of healthy control subjects (n = 47) before surgery. Results: Overall, temporal lobe epilepsy patients did not differ from healthy controls in syntactic and prosodic comprehension before surgery. They did perform less well on an affective prosody task. Post-operative testing revealed that syntactic and prosodic comprehension did not change after removal of the anterior temporal cortex. Discussion: The unchanged performance on syntactic and prosodic comprehension after removal of the anterior temporal cortex suggests that this area is not indispensable for sentence comprehension functions in temporal epilepsy patients. Potential implications for the postulated role of the anterior temporal lobe in the healthy brain are discussed.
  • De Lange, F. P., Koers, A., Kalkman, J. S., Bleijenberg, G., Hagoort, P., Van der Meer, J. W. M., & Toni, I. (2008). Increase in prefrontal cortical volume following cognitive behavioural therapy in patients with chronic fatigue syndrome. Brain, 131, 2172-2180. doi:10.1093/brain/awn140.

    Abstract

    Chronic fatigue syndrome (CFS) is a disabling disorder, characterized by persistent or relapsing fatigue. Recent studies have detected a decrease in cortical grey matter volume in patients with CFS, but it is unclear whether this cerebral atrophy constitutes a cause or a consequence of the disease. Cognitive behavioural therapy (CBT) is an effective behavioural intervention for CFS, which combines a rehabilitative approach of a graded increase in physical activity with a psychological approach that addresses thoughts and beliefs about CFS which may impair recovery. Here, we test the hypothesis that cerebral atrophy may be a reversible state that can ameliorate with successful CBT. We have quantified cerebral structural changes in 22 CFS patients that underwent CBT and 22 healthy control participants. At baseline, CFS patients had significantly lower grey matter volume than healthy control participants. CBT intervention led to a significant improvement in health status, physical activity and cognitive performance. Crucially, CFS patients showed a significant increase in grey matter volume, localized in the lateral prefrontal cortex. This change in cerebral volume was related to improvements in cognitive speed in the CFS patients. Our findings indicate that the cerebral atrophy associated with CFS is partially reversed after effective CBT. This result provides an example of macroscopic cortical plasticity in the adult human brain, demonstrating a surprisingly dynamic relation between behavioural state and cerebral anatomy. Furthermore, our results reveal a possible neurobiological substrate of psychotherapeutic treatment.
  • Patel, A. D., Iversen, J. R., Wassenaar, M., & Hagoort, P. (2008). Musical syntactic processing in agrammatic Broca's aphasia. Aphasiology, 22(7/8), 776-789. doi:10.1080/02687030701803804.

    Abstract

    Background: Growing evidence for overlap in the syntactic processing of language and music in non-brain-damaged individuals leads to the question of whether aphasic individuals with grammatical comprehension problems in language also have problems processing structural relations in music.

    Aims: The current study sought to test musical syntactic processing in individuals with Broca's aphasia and grammatical comprehension deficits, using both explicit and implicit tasks.

    Methods & Procedures: Two experiments were conducted. In the first experiment 12 individuals with Broca's aphasia (and 14 matched controls) were tested for their sensitivity to grammatical and semantic relations in sentences, and for their sensitivity to musical syntactic (harmonic) relations in chord sequences. An explicit task (acceptability judgement of novel sequences) was used. The second experiment, with 9 individuals with Broca's aphasia (and 12 matched controls), probed musical syntactic processing using an implicit task (harmonic priming).

    Outcomes & Results: In both experiments the aphasic group showed impaired processing of musical syntactic relations. Control experiments indicated that this could not be attributed to low-level problems with the perception of pitch patterns or with auditory short-term memory for tones.

    Conclusions: The results suggest that musical syntactic processing in agrammatic aphasia deserves systematic investigation, and that such studies could help probe the nature of the processing deficits underlying linguistic agrammatism. Methodological suggestions are offered for future work in this little-explored area.
  • Scheeringa, R., Bastiaansen, M. C. M., Petersson, K. M., Oostenveld, R., Norris, D. G., & Hagoort, P. (2008). Frontal theta EEG activity correlates negatively with the default mode network in resting state. International Journal of Psychophysiology, 67, 242-251. doi:10.1016/j.ijpsycho.2007.05.017.

    Abstract

    We used simultaneously recorded EEG and fMRI to investigate in which areas the BOLD signal correlates with frontal theta power changes, while subjects were quietly lying resting in the scanner with their eyes open. To obtain a reliable estimate of frontal theta power we applied ICA on band-pass filtered (2–9 Hz) EEG data. For each subject we selected the component that best matched the mid-frontal scalp topography associated with the frontal theta rhythm. We applied a time-frequency analysis on this component and used the time course of the frequency bin with the highest overall power to form a regressor that modeled spontaneous fluctuations in frontal theta power. No significant positive BOLD correlations with this regressor were observed. Extensive negative correlations were observed in the areas that together form the default mode network. We conclude that frontal theta activity can be seen as an EEG index of default mode network activity.
  • Toni, I., De Lange, F. P., Noordzij, M. L., & Hagoort, P. (2008). Language beyond action. Journal of Physiology, 102, 71-79. doi:10.1016/j.jphysparis.2008.03.005.

    Abstract

    The discovery of mirror neurons in macaques and of a similar system in humans has provided a new and fertile neurobiological ground for rooting a variety of cognitive faculties. Automatic sensorimotor resonance has been invoked as the key elementary process accounting for disparate (dys)functions, like imitation, ideomotor apraxia, autism, and schizophrenia. In this paper, we provide a critical appraisal of three of these claims that deal with the relationship between language and the motor system. Does language comprehension require the motor system? Was there an evolutionary switch from manual gestures to speech as the primary mode of language? Is human communication explained by automatic sensorimotor resonances? A positive answer to these questions would open the tantalizing possibility of bringing language and human communication within the fold of the motor system. We argue that the available empirical evidence does not appear to support these claims, and their theoretical scope fails to account for some crucial features of the phenomena they are supposed to explain. Without denying the enormous importance of the discovery of mirror neurons, we highlight the limits of their explanatory power for understanding language and communication.
  • Uddén, J., Folia, V., Forkstam, C., Ingvar, M., Fernández, G., Overeem, S., Van Elswijk, G., Hagoort, P., & Petersson, K. M. (2008). The inferior frontal cortex in artificial syntax processing: An rTMS study. Brain Research, 1224, 69-78. doi:10.1016/j.brainres.2008.05.070.

    Abstract

    The human capacity to implicitly acquire knowledge of structured sequences has recently been investigated in artificial grammar learning using functional magnetic resonance imaging. It was found that the left inferior frontal cortex (IFC; Brodmann's area (BA) 44/45) was related to classification performance. The objective of this study was to investigate whether the IFC (BA 44/45) is causally related to classification of artificial syntactic structures by means of an off-line repetitive transcranial magnetic stimulation (rTMS) paradigm. We manipulated the stimulus material in a 2 × 2 factorial design with grammaticality status and local substring familiarity as factors. The participants showed a reliable effect of grammaticality on classification of novel items after 5days of exposure to grammatical exemplars without performance feedback in an implicit acquisition task. The results show that rTMS of BA 44/45 improves syntactic classification performance by increasing the rejection rate of non-grammatical items and by shortening reaction times of correct rejections specifically after left-sided stimulation. A similar pattern of results is observed in FMRI experiments on artificial syntactic classification. These results suggest that activity in the inferior frontal region is causally related to artificial syntax processing.
  • Van Berkum, J. J. A., Van den Brink, D., Tesink, C. M. J. Y., Kos, M., & Hagoort, P. (2008). The neural integration of speaker and message. Journal of Cognitive Neuroscience, 20(4), 580-591. doi:10.1162/jocn.2008.20054.

    Abstract

    When do listeners take into account who the speaker is? We asked people to listen to utterances whose content sometimes did not match inferences based on the identity of the speaker (e.g., “If only I looked like Britney Spears” in a male voice, or “I have a large tattoo on my back” spoken with an upper-class accent). Event-related brain responses revealed that the speaker's identity is taken into account as early as 200–300 msec after the beginning of a spoken word, and is processed by the same early interpretation mechanism that constructs sentence meaning based on just the words. This finding is difficult to reconcile with standard “Gricean” models of sentence interpretation in which comprehenders initially compute a local, context-independent meaning for the sentence (“semantics”) before working out what it really means given the wider communicative context and the particular speaker (“pragmatics”). Because the observed brain response hinges on voice-based and usually stereotype-dependent inferences about the speaker, it also shows that listeners rapidly classify speakers on the basis of their voices and bring the associated social stereotypes to bear on what is being said. According to our event-related potential results, language comprehension takes very rapid account of the social context, and the construction of meaning based on language alone cannot be separated from the social aspects of language use. The linguistic brain relates the message to the speaker immediately.
  • Van Heuven, W. J. B., Schriefers, H., Dijkstra, T., & Hagoort, P. (2008). Language conflict in the bilingual brain. Cerebral Cortex, 18(11), 2706-2716. doi:10.1093/cercor/bhn030.

    Abstract

    The large majority of humankind is more or less fluent in 2 or even more languages. This raises the fundamental question how the language network in the brain is organized such that the correct target language is selected at a particular occasion. Here we present behavioral and functional magnetic resonance imaging data showing that bilingual processing leads to language conflict in the bilingual brain even when the bilinguals’ task only required target language knowledge. This finding demonstrates that the bilingual brain cannot avoid language conflict, because words from the target and nontarget languages become automatically activated during reading. Importantly, stimulus-based language conflict was found in brain regions in the LIPC associated with phonological and semantic processing, whereas response-based language conflict was only found in the pre-supplementary motor area/anterior cingulate cortex when language conflict leads to response conflicts.
  • Willems, R. M., Ozyurek, A., & Hagoort, P. (2008). Seeing and hearing meaning: ERP and fMRI evidence of word versus picture integration into a sentence context. Journal of Cognitive Neuroscience, 20, 1235-1249. doi:10.1162/jocn.2008.20085.

    Abstract

    Understanding language always occurs within a situational context and, therefore, often implies combining streams of information from different domains and modalities. One such combination is that of spoken language and visual information, which are perceived together in a variety of ways during everyday communication. Here we investigate whether and how words and pictures differ in terms of their neural correlates when they are integrated into a previously built-up sentence context. This is assessed in two experiments looking at the time course (measuring event-related potentials, ERPs) and the locus (using functional magnetic resonance imaging, fMRI) of this integration process. We manipulated the ease of semantic integration of word and/or picture to a previous sentence context to increase the semantic load of processing. In the ERP study, an increased semantic load led to an N400 effect which was similar for pictures and words in terms of latency and amplitude. In the fMRI study, we found overlapping activations to both picture and word integration in the left inferior frontal cortex. Specific activations for the integration of a word were observed in the left superior temporal cortex. We conclude that despite obvious differences in representational format, semantic information coming from pictures and words is integrated into a sentence context in similar ways in the brain. This study adds to the growing insight that the language system incorporates (semantic) information coming from linguistic and extralinguistic domains with the same neural time course and by recruitment of overlapping brain areas.
  • Willems, R. M., Oostenveld, R., & Hagoort, P. (2008). Early decreases in alpha and gamma band power distinguish linguistic from visual information during spoken sentence comprehension. Brain Research, 1219, 78-90. doi:10.1016/j.brainres.2008.04.065.

    Abstract

    Language is often perceived together with visual information. This raises the question on how the brain integrates information conveyed in visual and/or linguistic format during spoken language comprehension. In this study we investigated the dynamics of semantic integration of visual and linguistic information by means of time-frequency analysis of the EEG signal. A modified version of the N400 paradigm with either a word or a picture of an object being semantically incongruous with respect to the preceding sentence context was employed. Event-Related Potential (ERP) analysis showed qualitatively similar N400 effects for integration of either word or picture. Time-frequency analysis revealed early specific decreases in alpha and gamma band power for linguistic and visual information respectively. We argue that these reflect a rapid context-based analysis of acoustic (word) or visual (picture) form information. We conclude that although full semantic integration of linguistic and visual information occurs through a common mechanism, early differences in oscillations in specific frequency bands reflect the format of the incoming information and, importantly, an early context-based detection of its congruity with respect to the preceding language context
  • Li, X., Yang, Y., & Hagoort, P. (2008). Pitch accent and lexical tone processing in Chinese discourse comprehension: An ERP study. Brain Research, 1222, 192-200. doi:10.1016/j.brainres.2008.05.031.

    Abstract

    In the present study, event-related brain potentials (ERP) were recorded to investigate the role of pitch accent and lexical tone in spoken discourse comprehension. Chinese was used as material to explore the potential difference in the nature and time course of brain responses to sentence meaning as indicated by pitch accent and to lexical meaning as indicated by tone. In both cases, the pitch contour of critical words was varied. The results showed that both inconsistent pitch accent and inconsistent lexical tone yielded N400 effects, and there was no interaction between them. The negativity evoked by inconsistent pitch accent had the some topography as that evoked by inconsistent lexical tone violation, with a maximum over central–parietal electrodes. Furthermore, the effect for the combined violations was the sum of effects for pure pitch accent and pure lexical tone violation. However, the effect for the lexical tone violation appeared approximately 90 ms earlier than the effect of the pitch accent violation. It is suggested that there might be a correspondence between the neural mechanism underlying pitch accent and lexical meaning processing in context. They both reflect the integration of the current information into a discourse context, independent of whether the current information was sentence meaning indicated by accentuation, or lexical meaning indicated by tone. In addition, lexical meaning was processed earlier than sentence meaning conveyed by pitch accent during spoken language processing.

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