Peter Hagoort

Publications

Displaying 201 - 300 of 431
  • Stein, J. L., Medland, S. E., Vasquez, A. A., Hibar, D. P., Senstad, R. E., Winkler, A. M., Toro, R., Appel, K., Bartecek, R., Bergmann, Ø., Bernard, M., Brown, A. A., Cannon, D. M., Chakravarty, M. M., Christoforou, A., Domin, M., Grimm, O., Hollinshead, M., Holmes, A. J., Homuth, G. and 184 moreStein, J. L., Medland, S. E., Vasquez, A. A., Hibar, D. P., Senstad, R. E., Winkler, A. M., Toro, R., Appel, K., Bartecek, R., Bergmann, Ø., Bernard, M., Brown, A. A., Cannon, D. M., Chakravarty, M. M., Christoforou, A., Domin, M., Grimm, O., Hollinshead, M., Holmes, A. J., Homuth, G., Hottenga, J.-J., Langan, C., Lopez, L. M., Hansell, N. K., Hwang, K. S., Kim, S., Laje, G., Lee, P. H., Liu, X., Loth, E., Lourdusamy, A., Mattingsdal, M., Mohnke, S., Maniega, S. M., Nho, K., Nugent, A. C., O'Brien, C., Papmeyer, M., Pütz, B., Ramasamy, A., Rasmussen, J., Rijpkema, M., Risacher, S. L., Roddey, J. C., Rose, E. J., Ryten, M., Shen, L., Sprooten, E., Strengman, E., Teumer, A., Trabzuni, D., Turner, J., van Eijk, K., van Erp, T. G. M., van Tol, M.-J., Wittfeld, K., Wolf, C., Woudstra, S., Aleman, A., Alhusaini, S., Almasy, L., Binder, E. B., Brohawn, D. G., Cantor, R. M., Carless, M. A., Corvin, A., Czisch, M., Curran, J. E., Davies, G., de Almeida, M. A. A., Delanty, N., Depondt, C., Duggirala, R., Dyer, T. D., Erk, S., Fagerness, J., Fox, P. T., Freimer, N. B., Gill, M., Göring, H. H. H., Hagler, D. J., Hoehn, D., Holsboer, F., Hoogman, M., Hosten, N., Jahanshad, N., Johnson, M. P., Kasperaviciute, D., Kent, J. W. J., Kochunov, P., Lancaster, J. L., Lawrie, S. M., Liewald, D. C., Mandl, R., Matarin, M., Mattheisen, M., Meisenzahl, E., Melle, I., Moses, E. K., Mühleisen, T. W., Nauck, M., Nöthen, M. M., Olvera, R. L., Pandolfo, M., Pike, G. B., Puls, R., Reinvang, I., Rentería, M. E., Rietschel, M., Roffman, J. L., Royle, N. A., Rujescu, D., Savitz, J., Schnack, H. G., Schnell, K., Seiferth, N., Smith, C., Hernández, M. C. V., Steen, V. M., den Heuvel, M. V., van der Wee, N. J., Haren, N. E. M. V., Veltman, J. A., Völzke, H., Walker, R., Westlye, L. T., Whelan, C. D., Agartz, I., Boomsma, D. I., Cavalleri, G. L., Dale, A. M., Djurovic, S., Drevets, W. C., Hagoort, P., Hall, J., Heinz, A., Clifford, R. J., Foroud, T. M., Le Hellard, S., Macciardi, F., Montgomery, G. W., Poline, J. B., Porteous, D. J., Sisodiya, S. M., Starr, J. M., Sussmann, J., Toga, A. W., Veltman, D. J., Walter, H., Weiner, M. W., EPIGEN Consortium, IMAGENConsortium, Saguenay Youth Study Group, Bis, J. C., Ikram, M. A., Smith, A. V., Gudnason, V., Tzourio, C., Vernooij, M. W., Launer, L. J., DeCarli, C., Seshadri, S., Heart, C. f., Consortium, A. R. i. G. E. (., Andreassen, O. A., Apostolova, L. G., Bastin, M. E., Blangero, J., Brunner, H. G., Buckner, R. L., Cichon, S., Coppola, G., de Zubicaray, G. I., Deary, I. J., Donohoe, G., de Geus, E. J. C., Espeseth, T., Fernández, G., Glahn, D. C., Grabe, H. J., Hardy, J., Hulshoff Pol, H. E., Jenkinson, M., Kahn, R. S., McDonald, C., McIntosh, A. M., McMahon, F. J., McMahon, K. L., Meyer-Lindenberg, A., Morris, D. W., Müller-Myhsok, B., Nichols, T. E., Ophoff, R. A., Paus, T., Pausova, Z., Penninx, B. W., Sämann, P. G., Saykin, A. J., Schumann, G., Smoller, J. W., Wardlaw, J. M., Weale, M. E., Martin, N. G., Franke, B., Wright, M. J., Thompson, P. M., & the Enhancing Neuro Imaging Genetics through Meta-Analysis (ENIGMA) Consortium (2012). Identification of common variants associated with human hippocampal and intracranial volumes. Nature Genetics, 44, 552-561. doi:10.1038/ng.2250.

    Abstract

    Identifying genetic variants influencing human brain structures may reveal new biological mechanisms underlying cognition and neuropsychiatric illness. The volume of the hippocampus is a biomarker of incipient Alzheimer's disease and is reduced in schizophrenia, major depression and mesial temporal lobe epilepsy. Whereas many brain imaging phenotypes are highly heritable, identifying and replicating genetic influences has been difficult, as small effects and the high costs of magnetic resonance imaging (MRI) have led to underpowered studies. Here we report genome-wide association meta-analyses and replication for mean bilateral hippocampal, total brain and intracranial volumes from a large multinational consortium. The intergenic variant rs7294919 was associated with hippocampal volume (12q24.22; N = 21,151; P = 6.70 × 10(-16)) and the expression levels of the positional candidate gene TESC in brain tissue. Additionally, rs10784502, located within HMGA2, was associated with intracranial volume (12q14.3; N = 15,782; P = 1.12 × 10(-12)). We also identified a suggestive association with total brain volume at rs10494373 within DDR2 (1q23.3; N = 6,500; P = 5.81 × 10(-7)).
  • Udden, J., Ingvar, M., Hagoort, P., & Petersson, K. M. (2012). Implicit acquisition of grammars with crossed and nested non-adjacent dependencies: Investigating the push-down stack model. Cognitive Science, 36, 1078-1101. doi:10.1111/j.1551-6709.2012.01235.x.

    Abstract

    A recent hypothesis in empirical brain research on language is that the fundamental difference between animal and human communication systems is captured by the distinction between finite-state and more complex phrase-structure grammars, such as context-free and context-sensitive grammars. However, the relevance of this distinction for the study of language as a neurobiological system has been questioned and it has been suggested that a more relevant and partly analogous distinction is that between non-adjacent and adjacent dependencies. Online memory resources are central to the processing of non-adjacent dependencies as information has to be maintained across intervening material. One proposal is that an external memory device in the form of a limited push-down stack is used to process non-adjacent dependencies. We tested this hypothesis in an artificial grammar learning paradigm where subjects acquired non-adjacent dependencies implicitly. Generally, we found no qualitative differences between the acquisition of non-adjacent dependencies and adjacent dependencies. This suggests that although the acquisition of non-adjacent dependencies requires more exposure to the acquisition material, it utilizes the same mechanisms used for acquiring adjacent dependencies. We challenge the push-down stack model further by testing its processing predictions for nested and crossed multiple non-adjacent dependencies. The push-down stack model is partly supported by the results, and we suggest that stack-like properties are some among many natural properties characterizing the underlying neurophysiological mechanisms that implement the online memory resources used in language and structured sequence processing.
  • Van den Brink, D., Van Berkum, J. J. A., Bastiaansen, M. C. M., Tesink, C. M. J. Y., Kos, M., Buitelaar, J. K., & Hagoort, P. (2012). Empathy matters: ERP evidence for inter-individual differences in social language processing. Social, Cognitive and Affective Neuroscience, 7, 173-182. doi:10.1093/scan/nsq094.

    Abstract

    When an adult claims he cannot sleep without his teddy bear, people tend to react surprised. Language interpretation is, thus, influenced by social context, such as who the speaker is. The present study reveals inter-individual differences in brain reactivity to social aspects of language. Whereas women showed brain reactivity when stereotype-based inferences about a speaker conflicted with the content of the message, men did not. This sex difference in social information processing can be explained by a specific cognitive trait, one’s ability to empathize. Individuals who empathize to a greater degree revealed larger N400 effects (as well as a larger increase in γ-band power) to socially relevant information. These results indicate that individuals with high-empathizing skills are able to rapidly integrate information about the speaker with the content of the message, as they make use of voice-based inferences about the speaker to process language in a top-down manner. Alternatively, individuals with lower empathizing skills did not use information about social stereotypes in implicit sentence comprehension, but rather took a more bottom-up approach to the processing of these social pragmatic sentences.
  • Van Ackeren, M. J., Casasanto, D., Bekkering, H., Hagoort, P., & Rueschemeyer, S.-A. (2012). Pragmatics in action: Indirect requests engage theory of mind areas and the cortical motor network. Journal of Cognitive Neuroscience, 24, 2237-2247. doi:10.1162/jocn_a_00274.

    Abstract

    Research from the past decade has shown that understanding the meaning of words and utterances (i.e., abstracted symbols) engages the same systems we used to perceive and interact with the physical world in a content-specific manner. For example, understanding the word “grasp” elicits activation in the cortical motor network, that is, part of the neural substrate involved in planned and executing a grasping action. In the embodied literature, cortical motor activation during language comprehension is thought to reflect motor simulation underlying conceptual knowledge [note that outside the embodied framework, other explanations for the link between action and language are offered, e.g., Mahon, B. Z., & Caramazza, A. A critical look at the embodied cognition hypothesis and a new proposal for grouding conceptual content. Journal of Physiology, 102, 59–70, 2008; Hagoort, P. On Broca, brain, and binding: A new framework. Trends in Cognitive Sciences, 9, 416–423, 2005]. Previous research has supported the view that the coupling between language and action is flexible, and reading an action-related word form is not sufficient for cortical motor activation [Van Dam, W. O., van Dijk, M., Bekkering, H., & Rueschemeyer, S.-A. Flexibility in embodied lexical–semantic representations. Human Brain Mapping, doi: 10.1002/hbm.21365, 2011]. The current study goes one step further by addressing the necessity of action-related word forms for motor activation during language comprehension. Subjects listened to indirect requests (IRs) for action during an fMRI session. IRs for action are speech acts in which access to an action concept is required, although it is not explicitly encoded in the language. For example, the utterance “It is hot here!” in a room with a window is likely to be interpreted as a request to open the window. However, the same utterance in a desert will be interpreted as a statement. The results indicate (1) that comprehension of IR sentences activates cortical motor areas reliably more than comprehension of sentences devoid of any implicit motor information. This is true despite the fact that IR sentences contain no lexical reference to action. (2) Comprehension of IR sentences also reliably activates substantial portions of the theory of mind network, known to be involved in making inferences about mental states of others. The implications of these findings for embodied theories of language are discussed.
  • Wagensveld, B., Segers, E., Van Alphen, P. M., Hagoort, P., & Verhoeven, L. (2012). A neurocognitive perspective on rhyme awareness: The N450 rhyme effect. Brain Research, 1483, 63-70. doi:10.1016/j.brainres.2012.09.018.

    Abstract

    Rhyme processing is reflected in the electrophysiological signals of the brain as a negative deflection for non-rhyming as compared to rhyming stimuli around 450 ms after stimulus onset. Studies have shown that this N450 component is not solely sensitive to rhyme but also responds to other types of phonological overlap. In the present study, we examined whether the N450 component can be used to gain insight into the global similarity effect, indicating that rhyme judgment skills decrease when participants are presented with word pairs that share a phonological overlap but do not rhyme (e.g., bell–ball). We presented 20 adults with auditory rhyming, globally similar overlapping and unrelated word pairs. In addition to measuring behavioral responses by means of a yes/no button press, we also took EEG measures. The behavioral data showed a clear global similarity effect; participants judged overlapping pairs more slowly than unrelated pairs. However, the neural outcomes did not provide evidence that the N450 effect responds differentially to globally similar and unrelated word pairs, suggesting that globally similar and dissimilar non-rhyming pairs are processed in a similar fashion at the stage of early lexical access.
  • Wang, L., Jensen, O., Van den Brink, D., Weder, N., Schoffelen, J.-M., Magyari, L., Hagoort, P., & Bastiaansen, M. C. M. (2012). Beta oscillations relate to the N400m during language comprehension. Human Brain Mapping, 33, 2898-2912. doi:10.1002/hbm.21410.

    Abstract

    The relationship between the evoked responses (ERPs/ERFs) and the event-related changes in EEG/MEG power that can be observed during sentence-level language comprehension is as yet unclear. This study addresses a possible relationship between MEG power changes and the N400m component of the event-related field. Whole-head MEG was recorded while subjects listened to spoken sentences with incongruent (IC) or congruent (C) sentence endings. A clear N400m was observed over the left hemisphere, and was larger for the IC sentences than for the C sentences. A time–frequency analysis of power revealed a decrease in alpha and beta power over the left hemisphere in roughly the same time range as the N400m for the IC relative to the C condition. A linear regression analysis revealed a positive linear relationship between N400m and beta power for the IC condition, not for the C condition. No such linear relation was found between N400m and alpha power for either condition. The sources of the beta decrease were estimated in the LIFG, a region known to be involved in semantic unification operations. One source of the N400m was estimated in the left superior temporal region, which has been related to lexical retrieval. We interpret our data within a framework in which beta oscillations are inversely related to the engagement of task-relevant brain networks. The source reconstructions of the beta power suppression and the N400m effect support the notion of a dynamic communication between the LIFG and the left superior temporal region during language comprehension.
  • Wang, L., Bastiaansen, M. C. M., Yang, Y., & Hagoort, P. (2012). Information structure influences depth of syntactic processing: Event-related potential evidence for the Chomsky illusion. PLoS One, 7(10), e47917. doi:10.1371/journal.pone.0047917.

    Abstract

    Information structure facilitates communication between interlocutors by highlighting relevant information. It has previously been shown that information structure modulates the depth of semantic processing. Here we used event-related potentials to investigate whether information structure can modulate the depth of syntactic processing. In question-answer pairs, subtle (number agreement) or salient (phrase structure) syntactic violations were placed either in focus or out of focus through information structure marking. P600 effects to these violations reflect the depth of syntactic processing. For subtle violations, a P600 effect was observed in the focus condition, but not in the non-focus condition. For salient violations, comparable P600 effects were found in both conditions. These results indicate that information structure can modulate the depth of syntactic processing, but that this effect depends on the salience of the information. When subtle violations are not in focus, they are processed less elaborately. We label this phenomenon the Chomsky illusion.
  • Xiang, H., Dediu, D., Roberts, L., Van Oort, E., Norris, D., & Hagoort, P. (2012). The structural connectivity underpinning language aptitude, working memory and IQ in the perisylvian language network. Language Learning, 62(Supplement S2), 110-130. doi:10.1111/j.1467-9922.2012.00708.x.

    Abstract

    We carried out the first study on the relationship between individual language aptitude and structural connectivity of language pathways in the adult brain. We measured four components of language aptitude (vocabulary learning, VocL; sound recognition, SndRec; sound-symbol correspondence, SndSym; and grammatical inferencing, GrInf) using the LLAMA language aptitude test (Meara, 2005). Spatial working memory (SWM), verbal working memory (VWM) and IQ were also measured as control factors. Diffusion Tensor Imaging (DTI) was employed to investigate the structural connectivity of language pathways in the perisylvian language network. Principal Component Analysis (PCA) on behavioural measures suggests that a general ability might be important to the first stages of L2 acquisition. It also suggested that VocL, SndSy and SWM are more closely related to general IQ than SndRec and VocL, and distinguished the tasks specifically designed to tap into L2 acquisition (VocL, SndRec,SndSym and GrInf) from more generic measures (IQ, SWM and VWM). Regression analysis suggested significant correlations between most of these behavioural measures and the structural connectivity of certain language pathways, i.e., VocL and BA47-Parietal pathway, SndSym and inter-hemispheric BA45 pathway, GrInf and BA45-Temporal pathway and BA6-Temporal pathway, IQ and BA44-Parietal pathway, BA47-Parietal pathway, BA47-Temporal pathway and inter-hemispheric BA45 pathway, SWM and inter-hemispheric BA6 pathway and BA47-Parietal pathway, and VWM and BA47-Temporal pathway. These results are discussed in relation to relevant findings in the literature.
  • Zhu, Z., Hagoort, P., Zhang, J. X., Feng, G., Chen, H.-C., Bastiaansen, M. C. M., & Wang, S. (2012). The anterior left inferior frontal gyrus contributes to semantic unification. NeuroImage, 60, 2230-2237. doi:10.1016/j.neuroimage.2012.02.036.

    Abstract

    Semantic unification, the process by which small blocks of semantic information are combined into a coherent utterance, has been studied with various types of tasks. However, whether the brain activations reported in these studies are attributed to semantic unification per se or to other task-induced concomitant processes still remains unclear. The neural basis for semantic unification in sentence comprehension was examined using event-related potentials (ERP) and functional Magnetic Resonance Imaging (fMRI). The semantic unification load was manipulated by varying the goodness of fit between a critical word and its preceding context (in high cloze, low cloze and violation sentences). The sentences were presented in a serial visual presentation mode. The participants were asked to perform one of three tasks: semantic congruency judgment (SEM), silent reading for comprehension (READ), or font size judgment (FONT), in separate sessions. The ERP results showed a similar N400 amplitude modulation by the semantic unification load across all of the three tasks. The brain activations associated with the semantic unification load were found in the anterior left inferior frontal gyrus (aLIFG) in the FONT task and in a widespread set of regions in the other two tasks. These results suggest that the aLIFG activation reflects a semantic unification, which is different from other brain activations that may reflect task-specific strategic processing.

    Additional information

    Zhu_2012_suppl.dot
  • 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.

    Additional information

    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.

    Additional information

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

    Abstract

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

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

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

    A debated issue in the relationship between language and thought is how our linguistic abilities are involved in understanding the intentions of others (‘mentalizing’). The results of both theoretical and empirical work have been used to argue that linguistic, and more specifically, grammatical, abilities are crucial in representing the mental states of others. Here we contribute to this debate by investigating how damage to the language system influences the generation and understanding of intentional communicative behaviors. Four patients with pervasive language difficulties (severe global or agrammatic aphasia) engaged in an experimentally controlled non-verbal communication paradigm, which required signaling and understanding a communicative message. Despite their profound language problems they were able to engage in recipient design as well as intention recognition, showing similar indicators of mentalizing as have been observed in the neurologically healthy population. Our results show that aspects of the ability to communicate remain present even when core capacities of the language system are dysfunctional
  • Adank, P., Hagoort, P., & Bekkering, H. (2010). Imitation improves language comprehension. Psychological Science, 21, 1903-1909. doi:10.1177/0956797610389192.

    Abstract

    Humans imitate each other during social interaction. This imitative behavior streamlines social interaction and aids in learning to replicate actions. However, the effect of imitation on action comprehension is unclear. This study investigated whether vocal imitation of an unfamiliar accent improved spoken-language comprehension. Following a pretraining accent comprehension test, participants were assigned to one of six groups. The baseline group received no training, but participants in the other five groups listened to accented sentences, listened to and repeated accented sentences in their own accent, listened to and transcribed accented sentences, listened to and imitated accented sentences, or listened to and imitated accented sentences without being able to hear their own vocalizations. Posttraining measures showed that accent comprehension was most improved for participants who imitated the speaker’s accent. These results show that imitation may aid in streamlining interaction by improving spoken-language comprehension under adverse listening conditions.
  • Baggio, G., Choma, T., Van Lambalgen, M., & Hagoort, P. (2010). Coercion and compositionality. Journal of Cognitive Neuroscience, 22, 2131-2140. doi:10.1162/jocn.2009.21303.

    Abstract

    Research in psycholinguistics and in the cognitive neuroscience of language has suggested that semantic and syntactic integration are associated with different neurophysiologic correlates, such as the N400 and the P600 in the ERPs. However, only a handful of studies have investigated the neural basis of the syntax–semantics interface, and even fewer experiments have dealt with the cases in which semantic composition can proceed independently of the syntax. Here we looked into one such case—complement coercion—using ERPs. We compared sentences such as, “The journalist wrote the article” with “The journalist began the article.” The second sentence seems to involve a silent semantic element, which is expressed in the first sentence by the head of the VP “wrote the article.” The second type of construction may therefore require the reader to infer or recover from memory a richer event sense of the VP “began the article,” such as began writing the article, and to integrate that into a semantic representation of the sentence. This operation is referred to as “complement coercion.” Consistently with earlier reading time, eye tracking, and MEG studies, we found traces of such additional computations in the ERPs: Coercion gives rise to a long-lasting negative shift, which differs at least in duration from a standard N400 effect. Issues regarding the nature of the computation involved are discussed in the light of a neurocognitive model of language processing and a formal semantic analysis of coercion.
  • Bastiaansen, M. C. M., Magyari, L., & Hagoort, P. (2010). Syntactic unification operations are reflected in oscillatory dynamics during on-line sentence comprehension. Journal of Cognitive Neuroscience, 22, 1333-1347. doi:10.1162/jocn.2009.21283.

    Abstract

    There is growing evidence suggesting that synchronization changes in the oscillatory neuronal dynamics in the EEG or MEG reflect the transient coupling and uncoupling of functional networks related to different aspects of language comprehension. In this work, we examine how sentence-level syntactic unification operations are reflected in the oscillatory dynamics of the MEG. Participants read sentences that were either correct, contained a word category violation, or were constituted of random word sequences devoid of syntactic structure. A time-frequency analysis of MEG power changes revealed three types of effects. The first type of effect was related to the detection of a (word category) violation in a syntactically structured sentence, and was found in the alpha and gamma frequency bands. A second type of effect was maximally sensitive to the syntactic manipulations: A linear increase in beta power across the sentence was present for correct sentences, was disrupted upon the occurrence of a word category violation, and was absent in syntactically unstructured random word sequences. We therefore relate this effect to syntactic unification operations. Thirdly, we observed a linear increase in theta power across the sentence for all syntactically structured sentences. The effects are tentatively related to the building of a working memory trace of the linguistic input. In conclusion, the data seem to suggest that syntactic unification is reflected by neuronal synchronization in the lower-beta frequency band.
  • Fournier, R., Gussenhoven, C., Jensen, O., & Hagoort, P. (2010). Lateralization of tonal and intonational pitch processing: An MEG study. Brain Research, 1328, 79-88. doi:10.1016/j.brainres.2010.02.053.

    Abstract

    An MEG experiment was carried out in order to compare the processing of lexical-tonal and intonational contrasts, based on the tonal dialect of Roermond (the Netherlands). A set of words with identical phoneme sequences but distinct pitch contours, which represented different lexical meanings or discourse meanings (statement vs. question), were presented to native speakers as well as to a control group of speakers of Standard Dutch, a non-tone language. The stimuli were arranged in a mismatch paradigm, under three experimental conditions: in the first condition (lexical), the pitch contour differences between standard and deviant stimuli reflected differences between lexical meanings; in the second condition (intonational), the stimuli differed in their discourse meaning; in the third condition (combined), they differed both in their lexical and discourse meaning. In all three conditions, native as well as non-native responses showed a clear MMNm (magnetic mismatch negativity) in a time window from 150 to 250 ms after the divergence point of standard and deviant pitch contours. In the lexical condition, a stronger response was found over the left temporal cortex of native as well as non-native speakers. In the intonational condition, the same activation pattern was observed in the control group, but not in the group of native speakers, who showed a right-hemisphere dominance instead. Finally, in the combined (lexical and intonational) condition, brain reactions appeared to represent the summation of the patterns found in the other two conditions. In sum, the lateralization of pitch processing is condition-dependent in the native group only, which suggests that language experience determines how processes should be distributed over both temporal cortices, according to the functions available in the grammar.
  • Groen, W. B., Tesink, C. M. J. Y., Petersson, K. M., Van Berkum, J. J. A., Van der Gaag, R. J., Hagoort, P., & Buitelaar, J. K. (2010). Semantic, factual, and social language comprehension in adolescents with autism: An fMRI study. Cerebral Cortex, 20(8), 1937-1945. doi:10.1093/cercor/bhp264.

    Abstract

    Language in high-functioning autism is characterized by pragmatic and semantic deficits, and people with autism have a reduced tendency to integrate information. Because the left and right inferior frontal (LIF and RIF) regions are implicated with integration of speaker information, world knowledge, and semantic knowledge, we hypothesized that abnormal functioning of the LIF and RIF regions might contribute to pragmatic and semantic language deficits in autism. Brain activation of sixteen 12- to 18-year-old, high-functioning autistic participants was measured with functional magnetic resonance imaging during sentence comprehension and compared with that of twenty-six matched controls. The content of the pragmatic sentence was congruent or incongruent with respect to the speaker characteristics (male/female, child/adult, and upper class/lower class). The semantic- and world-knowledge sentences were congruent or incongruent with respect to semantic expectancies and factual expectancies about the world, respectively. In the semanticknowledge and world-knowledge condition, activation of the LIF region did not differ between groups. In sentences that required integration of speaker information, the autism group showed abnormally reduced activation of the LIF region. The results suggest that people with autism may recruit the LIF region in a different manner in tasks that demand integration of social information.
  • Junge, C., Hagoort, P., Kooijman, V., & Cutler, A. (2010). Brain potentials for word segmentation at seven months predict later language development. In K. Franich, K. M. Iserman, & L. L. Keil (Eds.), Proceedings of the 34th Annual Boston University Conference on Language Development. Volume 1 (pp. 209-220). Somerville, MA: Cascadilla Press.
  • Junge, C., Cutler, A., & Hagoort, P. (2010). Ability to segment words from speech as a precursor of later language development: Insights from electrophysiological responses in the infant brain. In M. Burgess, J. Davey, C. Don, & T. McMinn (Eds.), Proceedings of 20th International Congress on Acoustics, ICA 2010. Incorporating Proceedings of the 2010 annual conference of the Australian Acoustical Society (pp. 3727-3732). Australian Acoustical Society, NSW Division.
  • Kos, M., Vosse, T. G., Van den Brink, D., & Hagoort, P. (2010). About edible restaurants: Conflicts between syntax and semantics as revealed by ERPs. Frontiers in Psychology, 1, E222. doi:10.3389/fpsyg.2010.00222.

    Abstract

    In order to investigate conflicts between semantics and syntax, we recorded ERPs, while participants read Dutch sentences. Sentences containing conflicts between syntax and semantics (Fred eats in a sandwich…/ Fred eats a restaurant…) elicited an N400. These results show that conflicts between syntax and semantics not necessarily lead to P600 effects and are in line with the processing competition account. According to this parallel account the syntactic and semantic processing streams are fully interactive and information from one level can influence the processing at another level. The relative strength of the cues of the processing streams determines which level is affected most strongly by the conflict. The processing competition account maintains the distinction between the N400 as index for semantic processing and the P600 as index for structural processing.
  • Noordzij, M. L., Newman-Norlund, S. E., De Ruiter, J. P., Hagoort, P., Levinson, S. C., & Toni, I. (2010). Neural correlates of intentional communication. Frontiers in Neuroscience, 4, E188. doi:10.3389/fnins.2010.00188.

    Abstract

    We know a great deal about the neurophysiological mechanisms supporting instrumental actions, i.e. actions designed to alter the physical state of the environment. In contrast, little is known about our ability to select communicative actions, i.e. actions directly designed to modify the mental state of another agent. We have recently provided novel empirical evidence for a mechanism in which a communicator selects his actions on the basis of a prediction of the communicative intentions that an addressee is most likely to attribute to those actions. The main novelty of those finding was that this prediction of intention recognition is cerebrally implemented within the intention recognition system of the communicator, is modulated by the ambiguity in meaning of the communicative acts, and not by their sensorimotor complexity. The characteristics of this predictive mechanism support the notion that human communicative abilities are distinct from both sensorimotor and linguistic processes.
  • Pijnacker, J., Geurts, B., Van Lambalgen, M., Buitelaar, J., & Hagoort, P. (2010). Exceptions and anomalies: An ERP study on context sensitivity in autism. Neuropsychologia, 48, 2940-2951. doi:10.1016/j.neuropsychologia.2010.06.003.

    Abstract

    Several studies have demonstrated that people with ASD and intact language skills still have problems processing linguistic information in context. Given this evidence for reduced sensitivity to linguistic context, the question arises how contextual information is actually processed by people with ASD. In this study, we used event-related brain potentials (ERPs) to examine context sensitivity in high-functioning adults with autistic disorder (HFA) and Asperger syndrome at two levels: at the level of sentence processing and at the level of solving reasoning problems. We found that sentence context as well as reasoning context had an immediate ERP effect in adults with Asperger syndrome, as in matched controls. Both groups showed a typical N400 effect and a late positive component for the sentence conditions, and a sustained negativity for the reasoning conditions. In contrast, the HFA group demonstrated neither an N400 effect nor a sustained negativity. However, the HFA group showed a late positive component which was larger for semantically anomalous sentences than congruent sentences. Because sentence context had a modulating effect in a later phase, semantic integration is perhaps less automatic in HFA, and presumably more elaborate processes are needed to arrive at a sentence interpretation.
  • De Ruiter, J. P., Noordzij, M. L., Newman-Norlund, S., Hagoort, P., Levinson, S. C., & Toni, I. (2010). Exploring the cognitive infrastructure of communication. Interaction studies, 11, 51-77. doi:10.1075/is.11.1.05rui.

    Abstract

    Human communication is often thought about in terms of transmitted messages in a conventional code like a language. But communication requires a specialized interactive intelligence. Senders have to be able to perform recipient design, while receivers need to be able to do intention recognition, knowing that recipient design has taken place. To study this interactive intelligence in the lab, we developed a new task that taps directly into the underlying abilities to communicate in the absence of a conventional code. We show that subjects are remarkably successful communicators under these conditions, especially when senders get feedback from receivers. Signaling is accomplished by the manner in which an instrumental action is performed, such that instrumentally dysfunctional components of an action are used to convey communicative intentions. The findings have important implications for the nature of the human communicative infrastructure, and the task opens up a line of experimentation on human communication.
  • Simanova, I., Van Gerven, M., Oostenveld, R., & Hagoort, P. (2010). Identifying object categories from event-related EEG: Toward decoding of conceptual representations. Plos One, 5(12), E14465. doi:10.1371/journal.pone.0014465.

    Abstract

    Multivariate pattern analysis is a technique that allows the decoding of conceptual information such as the semantic category of a perceived object from neuroimaging data. Impressive single-trial classification results have been reported in studies that used fMRI. Here, we investigate the possibility to identify conceptual representations from event-related EEG based on the presentation of an object in different modalities: its spoken name, its visual representation and its written name. We used Bayesian logistic regression with a multivariate Laplace prior for classification. Marked differences in classification performance were observed for the tested modalities. Highest accuracies (89% correctly classified trials) were attained when classifying object drawings. In auditory and orthographical modalities, results were lower though still significant for some subjects. The employed classification method allowed for a precise temporal localization of the features that contributed to the performance of the classifier for three modalities. These findings could help to further understand the mechanisms underlying conceptual representations. The study also provides a first step towards the use of concept decoding in the context of real-time brain-computer interface applications.
  • Snijders, T. M., Petersson, K. M., & Hagoort, P. (2010). Effective connectivity of cortical and subcortical regions during unification of sentence structure. NeuroImage, 52, 1633-1644. doi:10.1016/j.neuroimage.2010.05.035.

    Abstract

    In a recent fMRI study we showed that left posterior middle temporal gyrus (LpMTG) subserves the retrieval of a word's lexical-syntactic properties from the mental lexicon (long-term memory), while left posterior inferior frontal gyrus (LpIFG) is involved in unifying (on-line integration of) this information into a sentence structure (Snijders et al., 2009). In addition, the right IFG, right MTG, and the right striatum were involved in the unification process. Here we report results from a psychophysical interactions (PPI) analysis in which we investigated the effective connectivity between LpIFG and LpMTG during unification, and how the right hemisphere areas and the striatum are functionally connected to the unification network. LpIFG and LpMTG both showed enhanced connectivity during the unification process with a region slightly superior to our previously reported LpMTG. Right IFG better predicted right temporal activity when unification processes were more strongly engaged, just as LpIFG better predicted left temporal activity. Furthermore, the striatum showed enhanced coupling to LpIFG and LpMTG during unification. We conclude that bilateral inferior frontal and posterior temporal regions are functionally connected during sentence-level unification. Cortico-subcortical connectivity patterns suggest cooperation between inferior frontal and striatal regions in performing unification operations on lexical-syntactic representations retrieved from LpMTG.
  • Van Leeuwen, T. M., Petersson, K. M., & Hagoort, P. (2010). Synaesthetic colour in the brain: Beyond colour areas. A functional magnetic resonance imaging study of synaesthetes and matched controls. PLoS One, 5(8), E12074. doi:10.1371/journal.pone.0012074.

    Abstract

    Background: In synaesthesia, sensations in a particular modality cause additional experiences in a second, unstimulated modality (e.g., letters elicit colour). Understanding how synaesthesia is mediated in the brain can help to understand normal processes of perceptual awareness and multisensory integration. In several neuroimaging studies, enhanced brain activity for grapheme-colour synaesthesia has been found in ventral-occipital areas that are also involved in real colour processing. Our question was whether the neural correlates of synaesthetically induced colour and real colour experience are truly shared. Methodology/Principal Findings: First, in a free viewing functional magnetic resonance imaging (fMRI) experiment, we located main effects of synaesthesia in left superior parietal lobule and in colour related areas. In the left superior parietal lobe, individual differences between synaesthetes (projector-associator distinction) also influenced brain activity, confirming the importance of the left superior parietal lobe for synaesthesia. Next, we applied a repetition suppression paradigm in fMRI, in which a decrease in the BOLD (blood-oxygenated-level-dependent) response is generally observed for repeated stimuli. We hypothesized that synaesthetically induced colours would lead to a reduction in BOLD response for subsequently presented real colours, if the neural correlates were overlapping. We did find BOLD suppression effects induced by synaesthesia, but not within the colour areas. Conclusions/Significance: Because synaesthetically induced colours were not able to suppress BOLD effects for real colour, we conclude that the neural correlates of synaesthetic colour experience and real colour experience are not fully shared. We propose that synaesthetic colour experiences are mediated by higher-order visual pathways that lie beyond the scope of classical, ventral-occipital visual areas. Feedback from these areas, in which the left parietal cortex is likely to play an important role, may induce V4 activation and the percept of synaesthetic colour.
  • Willems, R. M., Hagoort, P., & Casasanto, D. (2010). Body-specific representations of action verbs: Neural evidence from right- and left-handers. Psychological Science, 21, 67-74. doi:10.1177/0956797609354072.

    Abstract

    According to theories of embodied cognition, understanding a verb like throw involves unconsciously simulating the action of throwing, using areas of the brain that support motor planning. If understanding action words involves mentally simulating one’s 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, we used functional magnetic resonance imaging to compare premotor activity correlated with action verb understanding in right- and left-handers. Righthanders preferentially activated the left premotor cortex during lexical decisions on manual-action verbs (compared with nonmanual-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 lefthanders, who perform actions differently, use correspondingly different areas of the brain for representing action verb meanings.
  • Willems, R. M., Peelen, M. V., & Hagoort, P. (2010). Cerebral lateralization of face-selective and body-selective visual areas depends on handedness. Cerebral Cortex, 20, 1719-1725. doi:10.1093/cercor/bhp234.

    Abstract

    The left-hemisphere dominance for language is a core example of the functional specialization of the cerebral hemispheres. The degree of left-hemisphere dominance for language depends on hand preference: Whereas the majority of right-handers show left-hemispheric language lateralization, this number is reduced in left-handers. Here, we assessed whether handedness analogously has an influence upon lateralization in the visual system. Using functional magnetic resonance imaging, we localized 4 more or less specialized extrastriate areas in left- and right-handers, namely fusiform face area (FFA), extrastriate body area (EBA), fusiform body area (FBA), and human motion area (human middle temporal [hMT]). We found that lateralization of FFA and EBA depends on handedness: These areas were right lateralized in right-handers but not in left-handers. A similar tendency was observed in FBA but not in hMT. We conclude that the relationship between handedness and hemispheric lateralization extends to functionally lateralized parts of visual cortex, indicating a general coupling between cerebral lateralization and handedness. Our findings indicate that hemispheric specialization is not fixed but can vary considerably across individuals even in areas engaged relatively early in the visual system.
  • Willems, R. M., De Boer, M., De Ruiter, J. P., Noordzij, M. L., Hagoort, P., & Toni, I. (2010). A dissociation between linguistic and communicative abilities in the human brain. Psychological Science, 21, 8-14. doi:10.1177/0956797609355563.

    Abstract

    Although language is an effective vehicle for communication, it is unclear how linguistic and communicative abilities relate to each other. Some researchers have argued that communicative message generation involves perspective taking (mentalizing), and—crucially—that mentalizing depends on language. We employed a verbal communication paradigm to directly test whether the generation of a communicative action relies on mentalizing and whether the cerebral bases of communicative message generation are distinct from parts of cortex sensitive to linguistic variables. We found that dorsomedial prefrontal cortex, a brain area consistently associated with mentalizing, was sensitive to the communicative intent of utterances, irrespective of linguistic difficulty. In contrast, left inferior frontal cortex, an area known to be involved in language, was sensitive to the linguistic demands of utterances, but not to communicative intent. These findings show that communicative and linguistic abilities rely on cerebrally (and computationally) distinct mechanisms
  • Willems, R. M., & Hagoort, P. (2010). Cortical motor contributions to language understanding. In L. Hermer (Ed.), Reciprocal interactions among early sensory and motor areas and higher cognitive networks (pp. 51-72). Kerala, India: Research Signpost Press.

    Abstract

    Here we review evidence from cognitive neuroscience for a tight relation between language and action in the brain. We focus on two types of relation between language and action. First, we investigate whether the perception of speech and speech sounds leads to activation of parts of the cortical motor system also involved in speech production. Second, we evaluate whether understanding action-related language involves the activation of parts of the motor system. We conclude that whereas there is considerable evidence that understanding language can involve parts of our motor cortex, this relation is best thought of as inherently flexible. As we explain, the exact nature of the input as well as the intention with which language is perceived influences whether and how motor cortex plays a role in language processing.
  • Willems, R. M., Toni, I., Hagoort, P., & Casasanto, D. (2010). Neural dissociations between action verb understanding and motor imagery. Journal of Cognitive Neuroscience, 22(10), 2387-2400. doi:10.1162/jocn.2009.21386.

    Abstract

    According to embodied theories of language, people understand a verb like throw, at least in part, by mentally simulating throwing. This implicit simulation is often assumed to be similar or identical to motor imagery. Here we used fMRI totest whether implicit simulations of actions during language understanding involve the same cortical motor regions as explicit motor imagery Healthy participants were presented with verbs related to hand actions (e.g., to throw) and nonmanual actions (e.g., to kneel). They either read these verbs (lexical decision task) or actively imagined performing the actions named by the verbs (imagery task). Primary motor cortex showd effector-specific activation during imagery, but not during lexical decision. Parts of premotor cortex distinguished manual from nonmanual actions during both lexical decision and imagery, but there was no overlap or correlation between regions activated during the two tasks. These dissociations suggest that implicit simulation and explicit imagery cued by action verbs may involve different types of motor representations and that the construct of “mental simulation” should be distinguished from “mental imagery” in embodied theories of language.
  • Xiang, H.-D., Fonteijn, H. M., Norris, D. G., & Hagoort, P. (2010). Topographical functional connectivity pattern in the perisylvian language networks. Cerebral Cortex, 20, 549-560. doi:10.1093/cercor/bhp119.

    Abstract

    We performed a resting-state functional connectivity study to investigate directly the functional correlations within the perisylvian language networks by seeding from 3 subregions of Broca's complex (pars opercularis, pars triangularis, and pars orbitalis) and their right hemisphere homologues. A clear topographical functional connectivity pattern in the left middle frontal, parietal, and temporal areas was revealed for the 3 left seeds. This is the first demonstration that a functional connectivity topology can be observed in the perisylvian language networks. The results support the assumption of the functional division for phonology, syntax, and semantics of Broca's complex as proposed by the memory, unification, and control (MUC) model and indicated a topographical functional organization in the perisylvian language networks, which suggests a possible division of labor for phonological, syntactic, and semantic function in the left frontal, parietal, and temporal areas.
  • 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.
  • Hagoort, P. (2007). The memory, unification, and control (MUC) model of language. In T. Sakamoto (Ed.), Communicating skills of intention (pp. 259-291). Tokyo: Hituzi Syobo.
  • Hagoort, P. (2007). The memory, unification, and control (MUC) model of language. In A. S. Meyer, L. Wheeldon, & A. Krott (Eds.), Automaticity and control in language processing (pp. 243-270). Hove: Psychology Press.
  • Hagoort, P., & Van Berkum, J. J. A. (2007). Beyond the sentence given. Philosophical Transactions of the Royal Society. Series B: Biological Sciences, 362, 801-811.

    Abstract

    A central and influential idea among researchers of language is that our language faculty is organized according to Fregean compositionality, which states that the meaning of an utterance is a function of the meaning of its parts and of the syntactic rules by which these parts are combined. Since the domain of syntactic rules is the sentence, the implication of this idea is that language interpretation takes place in a two-step fashion. First, the meaning of a sentence is computed. In a second step, the sentence meaning is integrated with information from prior discourse, world knowledge, information about the speaker and semantic information from extra-linguistic domains such as co-speech gestures or the visual world. Here, we present results from recordings of event-related brain potentials that are inconsistent with this classical two-step model of language interpretation. Our data support a one-step model in which knowledge about the context and the world, concomitant information from other modalities, and the speaker are brought to bear immediately, by the same fast-acting brain system that combines the meanings of individual words into a message-level representation. Underlying the one-step model is the immediacy assumption, according to which all available information will immediately be used to co-determine the interpretation of the speaker's message. Functional magnetic resonance imaging data that we collected indicate that Broca's area plays an important role in semantic unification. Language comprehension involves the rapid incorporation of information in a 'single unification space', coming from a broader range of cognitive domains than presupposed in the standard two-step model of interpretation.
  • Hald, L. A., Steenbeek-Planting, E. G., & Hagoort, P. (2007). The interaction of discourse context and world knowledge in online sentence comprehension: Evidence from the N400. Brain Research, 1146, 210-218. doi:10.1016/j.brainres.2007.02.054.

    Abstract

    In an ERP experiment we investigated how the recruitment and integration of world knowledge information relate to the integration of information within a current discourse context. Participants were presented with short discourse contexts which were followed by a sentence that contained a critical word that was correct or incorrect based on general world knowledge and the supporting discourse context, or was more or less acceptable based on the combination of general world knowledge and the specific local discourse context. Relative to the critical word in the correct world knowledge sentences following a neutral discourse, all other critical words elicited an N400 effect that began at about 300 ms after word onset. However, the magnitude of the N400 effect varied in a way that suggests an interaction between world knowledge and discourse context. The results indicate that both world knowledge and discourse context have an effect on sentence interpretation, but neither overrides the other.
  • Ozyurek, A., Willems, R. M., Kita, S., & Hagoort, P. (2007). On-line integration of semantic information from speech and gesture: Insights from event-related brain potentials. Journal of Cognitive Neuroscience, 19(4), 605-616. doi:10.1162/jocn.2007.19.4.605.

    Abstract

    During language comprehension, listeners use the global semantic representation from previous sentence or discourse context to immediately integrate the meaning of each upcoming word into the unfolding message-level representation. Here we investigate whether communicative gestures that often spontaneously co-occur with speech are processed in a similar fashion and integrated to previous sentence context in the same way as lexical meaning. Event-related potentials were measured while subjects listened to spoken sentences with a critical verb (e.g., knock), which was accompanied by an iconic co-speech gesture (i.e., KNOCK). Verbal and/or gestural semantic content matched or mismatched the content of the preceding part of the sentence. Despite the difference in the modality and in the specificity of meaning conveyed by spoken words and gestures, the latency, amplitude, and topographical distribution of both word and gesture mismatches are found to be similar, indicating that the brain integrates both types of information simultaneously. This provides evidence for the claim that neural processing in language comprehension involves the simultaneous incorporation of information coming from a broader domain of cognition than only verbal semantics. The neural evidence for similar integration of information from speech and gesture emphasizes the tight interconnection between speech and co-speech gestures.

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