Peter Hagoort

Publications

Displaying 1 - 39 of 39
  • Arana, S., Marquand, A., Hulten, A., Hagoort, P., & Schoffelen, J.-M. (2020). Sensory modality-independent activation of the brain network for language. The Journal of Neuroscience, 40(14), 2914-2924. doi:10.1523/JNEUROSCI.2271-19.2020.

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Additional information

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

    Abstract

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

    Additional information

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

    Abstract

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

    Additional information

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

    Abstract

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

    Abstract

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

    Abstract

    Catecholamine (CA) function has been widely implicated in cognitive functions that are tied to the prefrontal cortex and striatal areas. The present study investigated the effects of methylphenidate, which is a CA agonist, on the electroencephalogram (EEG) response related to semantic processing using a double-blind, placebo-controlled, randomized, crossover, within-subject design. Forty-eight healthy participants read semantically congruent or incongruent sentences after receiving 20-mg methylphenidate or a placebo while their brain activity was monitored with EEG. To probe whether the catecholaminergic modulation is task-dependent, in one condition participants had to focus on comprehending the sentences, while in the other condition, they only had to attend to the font size of the sentence. The results demonstrate that methylphenidate has a task-dependent effect on semantic processing. Compared to placebo, when semantic processing was task-irrelevant, methylphenidate enhanced the detection of semantic incongruence as indexed by a larger N400 amplitude in the incongruent sentences; when semantic processing was task-relevant, methylphenidate induced a larger N400 amplitude in the semantically congruent condition, which was followed by a larger late positive complex effect. These results suggest that CA-related neurotransmitters influence language processing, possibly through the projections between the prefrontal cortex and the striatum, which contain many CA receptors.
  • Baggio, G., & Hagoort, P. (2011). The balance between memory and unification in semantics: A dynamic account of the N400. Language and Cognitive Processes, 26, 1338-1367. doi:10.1080/01690965.2010.542671.

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

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

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    mmc1.pdf
  • Segaert, K., Menenti, L., Weber, K., & Hagoort, P. (2011). A paradox of syntactic priming: Why response tendencies show priming for passives, and response latencies show priming for actives. PLoS One, 6(10), e24209. doi:10.1371/journal.pone.0024209.

    Abstract

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

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

    Abstract

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

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

    A debated issue in the relationship between language and thought is how our linguistic abilities are involved in understanding the intentions of others (‘mentalizing’). The results of both theoretical and empirical work have been used to argue that linguistic, and more specifically, grammatical, abilities are crucial in representing the mental states of others. Here we contribute to this debate by investigating how damage to the language system influences the generation and understanding of intentional communicative behaviors. Four patients with pervasive language difficulties (severe global or agrammatic aphasia) engaged in an experimentally controlled non-verbal communication paradigm, which required signaling and understanding a communicative message. Despite their profound language problems they were able to engage in recipient design as well as intention recognition, showing similar indicators of mentalizing as have been observed in the neurologically healthy population. Our results show that aspects of the ability to communicate remain present even when core capacities of the language system are dysfunctional
  • Aleman, A., Formisano, E., Koppenhagen, H., Hagoort, P., De Haan, E. H. F., & Kahn, R. S. (2005). The functional neuroanatomy of metrical stress evaluation of perceived and imagined spoken words. Cerebral Cortex, 15(2), 221-228. doi:10.1093/cercor/bhh124.

    Abstract

    We hypothesized that areas in the temporal lobe that have been implicated in the phonological processing of spoken words would also be activated during the generation and phonological processing of imagined speech. We tested this hypothesis using functional magnetic resonance imaging during a behaviorally controlled task of metrical stress evaluation. Subjects were presented with bisyllabic words and had to determine the alternation of strong and weak syllables. Thus, they were required to discriminate between weak-initial words and strong-initial words. In one condition, the stimuli were presented auditorily to the subjects (by headphones). In the other condition the stimuli were presented visually on a screen and subjects were asked to imagine hearing the word. Results showed activation of the supplementary motor area, inferior frontal gyrus (Broca's area) and insula in both conditions. In the superior temporal gyrus (STG) and in the superior temporal sulcus (STS) strong activation was observed during the auditory (perceptual) condition. However, a region located in the posterior part of the STS/STG also responded during the imagery condition. No activation of this same region of the STS was observed during a control condition which also involved processing of visually presented words, but which required a semantic decision from the subject. We suggest that processing of metrical stress, with or without auditory input, relies in part on cortical interface systems located in the posterior part of STS/STG. These results corroborate behavioral evidence regarding phonological loop involvement in auditory–verbal imagery.
  • Bastiaansen, M. C. M., Van der Linden, M., Ter Keurs, M., Dijkstra, T., & Hagoort, P. (2005). Theta responses are involved in lexico-semantic retrieval during language processing. Journal of Cognitive Neuroscience, 17, 530-541. doi:10.1162/0898929053279469.

    Abstract

    Oscillatory neuronal dynamics, observed in the human electroencephalogram (EEG) during language processing, have been related to the dynamic formation of functionally coherent networks that serve the role of integrating the different sources of information needed for understanding the linguistic input. To further explore the functional role of oscillatory synchrony during language processing, we quantified event-related EEG power changes induced by the presentation of open-class (OC) words and closed-class (CC) words in a wide range of frequencies (from 1 to 30 Hz), while subjects read a short story. Word presentation induced three oscillatory components: a theta power increase (4–7 Hz), an alpha power decrease (10–12 Hz), and a beta power decrease (16–21 Hz). Whereas the alpha and beta responses showed mainly quantitative differences between the two word classes, the theta responses showed qualitative differences between OC words and CC words: A theta power increase was found over left temporal areas for OC words, but not for CC words. The left temporal theta increase may index the activation of a network involved in retrieving the lexical–semantic properties of the OC items.
  • Hagoort, P. (2005). On Broca, brain, and binding: A new framework. Trends in Cognitive Sciences, 9(9), 416-423. doi:10.1016/j.tics.2005.07.004.

    Abstract

    In speaking and comprehending language, word information is retrieved from memory and combined into larger units (unification). Unification operations take place in parallel at the semantic, syntactic and phonological levels of processing. This article proposes a new framework that connects psycholinguistic models to a neurobiological account of language. According to this proposal the left inferior frontal gyrus (LIFG) plays an important role in unification. Research in other domains of cognition indicates that left prefrontal cortex has the necessary neurobiological characteristics for its involvement in the unification for language. I offer here a psycholinguistic perspective on the nature of language unification and the role of LIFG.
  • Hagoort, P. (2005). De talige aap. Linguaan, 26-35.
  • Hagoort, P. (2005). Breintaal. In S. Knols, & D. Redeker (Eds.), NWO-Spinozapremies 2005 (pp. 21-34). Den Haag: NWO.
  • Hagoort, P. (2005). Broca's complex as the unification space for language. In A. Cutler (Ed.), Twenty-first century psycholinguistics: Four cornerstones (pp. 157-173). Mahwah, NJ: Erlbaum.
  • Kooijman, V., Hagoort, P., & Cutler, A. (2005). Electrophysiological evidence for prelinguistic infants' word recognition in continuous speech. Cognitive Brain Research, 24(1), 109-116. doi:10.1016/j.cogbrainres.2004.12.009.

    Abstract

    Children begin to talk at about age one. The vocabulary they need to do so must be built on perceptual evidence and, indeed, infants begin to recognize spoken words long before they talk. Most of the utterances infants hear, however, are continuous, without pauses between words, so constructing a vocabulary requires them to decompose continuous speech in order to extract the individual words. Here, we present electrophysiological evidence that 10-month-old infants recognize two-syllable words they have previously heard only in isolation when these words are presented anew in continuous speech. Moreover, they only need roughly the first syllable of the word to begin doing this. Thus, prelinguistic infants command a highly efficient procedure for segmentation and recognition of spoken words in the absence of an existing vocabulary, allowing them to tackle effectively the problem of bootstrapping a lexicon out of the highly variable, continuous speech signals in their environment.
  • De Lange, F. P., Kalkman, J. S., Bleijenberg, G., Hagoort, P., Van der Meer, J. W. M., & Toni, I. (2005). Gray matter volume reduction in the chronic fatigue syndrome. NeuroImage, 26, 777-781. doi:10.1016/j.neuroimage.2005.02.037.

    Abstract

    The chronic fatigue syndrome (CFS) is a disabling disorder of unknown etiology. The symptomatology of CFS (central fatigue, impaired concentration, attention and memory) suggests that this disorder could be related to alterations at the level of the central nervous system. In this study, we have used an automated and unbiased morphometric technique to test whether CFS patients display structural cerebral abnormalities. We mapped structural cerebral morphology and volume in two cohorts of CFS patients (in total 28 patients) and healthy controls (in total 28 controls) from high-resolution structural magnetic resonance images, using voxel-based morphometry. Additionally, we recorded physical activity levels to explore the relation between severity of CFS symptoms and cerebral abnormalities. We observed significant reductions in global gray matter volume in both cohorts of CFS patients, as compared to matched control participants. Moreover, the decline in gray matter volume was linked to the reduction in physical activity, a core aspect of CFS. These findings suggest that the central nervous system plays a key role in the pathophysiology of CFS and point to a new objective and quantitative tool for clinical diagnosis of this disabling disorder.
  • De Lange, F. P., Hagoort, P., & Toni, I. (2005). Neural topography and content of movement representations. Journal of Cognitive Neuroscience, 17(1), 97-112. doi:10.1162/0898929052880039.

    Abstract

    We have used implicit motor imagery to investigate the neural correlates of motor planning independently from actual movements. Subjects were presented with drawings of left or right hands and asked to judge the hand laterality, regardless of the stimulus rotation from its upright orientation. We paired this task with a visual imagery control task, in which subjects were presented with typographical characters and asked to report whether they saw a canonical letter or its mirror image, regardless of its rotation. We measured neurovascular activity with fast event-related fMRI, distinguishing responses parametrically related to motor imagery from responses evoked by visual imagery and other task-related phenomena. By quantifying behavioral and neurovascular correlates of imagery on a trial-by-trial basis, we could discriminate between stimulusrelated, mental rotation-related, and response-related neural activity. We found that specific portions of the posterior parietal and precentral cortex increased their activity as a function of mental rotation only during the motor imagery task. Within these regions, the parietal cortex was visually responsive, whereas the dorsal precentral cortex was not. Response- but not rotation-related activity was found around the left central sulcus (putative primary motor cortex) during both imagery tasks. Our study provides novel evidence on the topography and content of movement representations in the human brain. During intended action, the posterior parietal cortex combines somatosensory and visuomotor information, whereas the dorsal premotor cortex generates the actual motor plan, and the primary motor cortex deals with movement execution. We discuss the relevance of these results in the context of current models of action planning.
  • Van Berkum, J. J. A., Brown, C. M., Zwitserlood, P., Kooijman, V., & Hagoort, P. (2005). Anticipating upcoming words in discourse: Evidence from ERPs and reading times. Journal of Experimental Psychology: Learning, Memory, and Cognition, 31(3), 443-467. doi:10.1037/0278-7393.31.3.443.

    Abstract

    The authors examined whether people can use their knowledge of the wider discourse rapidly enough to anticipate specific upcoming words as a sentence is unfolding. In an event-related brain potential (ERP) experiment, subjects heard Dutch stories that supported the prediction of a specific noun. To probe whether this noun was anticipated at a preceding indefinite article, stories were continued with a gender-marked adjective whose suffix mismatched the upcoming noun's syntactic gender. Prediction-inconsistent adjectives elicited a differential ERP effect, which disappeared in a no-discourse control experiment. Furthermore, in self-paced reading, prediction-inconsistent adjectives slowed readers down before the noun. These findings suggest that people can indeed predict upcoming words in fluent discourse and, moreover, that these predicted words can immediately begin to participate in incremental parsing operations.
  • Wassenaar, M., & Hagoort, P. (2005). Word-category violations in patients with Broca's aphasia: An ERP study. Brain and Language, 92, 117-137. doi:10.1016/j.bandl.2004.05.011.

    Abstract

    An event-related brain potential experiment was carried out to investigate on-line syntactic processing in patients with Broca’s aphasia. Subjects were visually presented with sentences that were either syntactically correct or contained violations of word-category. Three groups of subjects were tested: Broca patients (N=11), non-aphasic patients with a right hemisphere (RH) lesion (N=9), and healthy aged-matched controls (N=15). Both control groups appeared sensitive to the violations of word-category as shown by clear P600/SPS effects. The Broca patients displayed only a very reduced and delayed P600/SPS effect. The results are discussed in the context of a lexicalist parsing model. It is concluded that Broca patients are hindered to detect on-line violations of word-category, if word class information is incomplete or delayed available.

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