Peter Hagoort

Publications

Displaying 1 - 37 of 37
  • Acheson, D. J., & Hagoort, P. (2014). Twisting tongues to test for conflict monitoring in speech production. Frontiers in Human Neuroscience, 8: 206. doi:10.3389/fnhum.2014.00206.

    Abstract

    A number of recent studies have hypothesized that monitoring in speech production may occur via domain-general mechanisms responsible for the detection of response conflict. Outside of language, two ERP components have consistently been elicited in conflict-inducing tasks (e.g., the flanker task): the stimulus-locked N2 on correct trials, and the response-locked error-related negativity (ERN). The present investigation used these electrophysiological markers to test whether a common response conflict monitor is responsible for monitoring in speech and non-speech tasks. Electroencephalography (EEG) was recorded while participants performed a tongue twister (TT) task and a manual version of the flanker task. In the TT task, people rapidly read sequences of four nonwords arranged in TT and non-TT patterns three times. In the flanker task, people responded with a left/right button press to a center-facing arrow, and conflict was manipulated by the congruency of the flanking arrows. Behavioral results showed typical effects of both tasks, with increased error rates and slower speech onset times for TT relative to non-TT trials and for incongruent relative to congruent flanker trials. In the flanker task, stimulus-locked EEG analyses replicated previous results, with a larger N2 for incongruent relative to congruent trials, and a response-locked ERN. In the TT task, stimulus-locked analyses revealed broad, frontally-distributed differences beginning around 50 ms and lasting until just before speech initiation, with TT trials more negative than non-TT trials; response-locked analyses revealed an ERN. Correlation across these measures showed some correlations within a task, but little evidence of systematic cross-task correlation. Although the present results do not speak against conflict signals from the production system serving as cues to self-monitoring, they are not consistent with signatures of response conflict being mediated by a single, domain-general conflict monitor
  • Basnakova, J., Weber, K., Petersson, K. M., Van Berkum, J. J. A., & Hagoort, P. (2014). Beyond the language given: The neural correlates of inferring speaker meaning. Cerebral Cortex, 24(10), 2572-2578. doi:10.1093/cercor/bht112.

    Abstract

    Even though language allows us to say exactly what we mean, we often use language to say things indirectly, in a way that depends on the specific communicative context. For example, we can use an apparently straightforward sentence like "It is hard to give a good presentation" to convey deeper meanings, like "Your talk was a mess!" One of the big puzzles in language science is how listeners work out what speakers really mean, which is a skill absolutely central to communication. However, most neuroimaging studies of language comprehension have focused on the arguably much simpler, context-independent process of understanding direct utterances. To examine the neural systems involved in getting at contextually constrained indirect meaning, we used functional magnetic resonance imaging as people listened to indirect replies in spoken dialog. Relative to direct control utterances, indirect replies engaged dorsomedial prefrontal cortex, right temporo-parietal junction and insula, as well as bilateral inferior frontal gyrus and right medial temporal gyrus. This suggests that listeners take the speaker's perspective on both cognitive (theory of mind) and affective (empathy-like) levels. In line with classic pragmatic theories, our results also indicate that currently popular "simulationist" accounts of language comprehension fail to explain how listeners understand the speaker's intended message.
  • Cai, D., Fonteijn, H. M., Guadalupe, T., Zwiers, M., Wittfeld, K., Teumer, A., Hoogman, M., Arias Vásquez, A., Yang, Y., Buitelaar, J., Fernández, G., Brunner, H. G., Van Bokhoven, H., Franke, B., Hegenscheid, K., Homuth, G., Fisher, S. E., Grabe, H. J., Francks, C., & Hagoort, P. (2014). A genome wide search for quantitative trait loci affecting the cortical surface area and thickness of Heschl's gyrus. Genes, Brain and Behavior, 13, 675-685. doi:10.1111/gbb.12157.

    Abstract

    Heschl's gyrus (HG) is a core region of the auditory cortex whose morphology is highly variable across individuals. This variability has been linked to sound perception ability in both speech and music domains. Previous studies show that variations in morphological features of HG, such as cortical surface area and thickness, are heritable. To identify genetic variants that affect HG morphology, we conducted a genome-wide association scan (GWAS) meta-analysis in 3054 healthy individuals using HG surface area and thickness as quantitative traits. None of the single nucleotide polymorphisms (SNPs) showed association P values that would survive correction for multiple testing over the genome. The most significant association was found between right HG area and SNP rs72932726 close to gene DCBLD2 (3q12.1; P=2.77x10(-7)). This SNP was also associated with other regions involved in speech processing. The SNP rs333332 within gene KALRN (3q21.2; P=2.27x10(-6)) and rs143000161 near gene COBLL1 (2q24.3; P=2.40x10(-6)) were associated with the area and thickness of left HG, respectively. Both genes are involved in the development of the nervous system. The SNP rs7062395 close to the X-linked deafness gene POU3F4 was associated with right HG thickness (Xq21.1; P=2.38x10(-6)). This is the first molecular genetic analysis of variability in HG morphology
  • Chu, M., & Hagoort, P. (2014). Synchronization of speech and gesture: Evidence for interaction in action. Journal of Experimental Psychology: General, 143(4), 1726-1741. doi:10.1037/a0036281.

    Abstract

    Language and action systems are highly interlinked. A critical piece of evidence is that speech and its accompanying gestures are tightly synchronized. Five experiments were conducted to test 2 hypotheses about the synchronization of speech and gesture. According to the interactive view, there is continuous information exchange between the gesture and speech systems, during both their planning and execution phases. According to the ballistic view, information exchange occurs only during the planning phases of gesture and speech, but the 2 systems become independent once their execution has been initiated. In all experiments, participants were required to point to and/or name a light that had just lit up. Virtual reality and motion tracking technologies were used to disrupt their gesture or speech execution. Participants delayed their speech onset when their gesture was disrupted. They did so even when their gesture was disrupted at its late phase and even when they received only the kinesthetic feedback of their gesture. Also, participants prolonged their gestures when their speech was disrupted. These findings support the interactive view and add new constraints on models of speech and gesture production
  • Cristia, A., Seidl, A., Junge, C., Soderstrom, M., & Hagoort, P. (2014). Predicting individual variation in language from infant speech perception measures. Child development, 85(4), 1330-1345. doi:10.1111/cdev.12193.

    Abstract

    There are increasing reports that individual variation in behavioral and neurophysiological measures of infant speech processing predicts later language outcomes, and specifically concurrent or subsequent vocabulary size. If such findings are held up under scrutiny, they could both illuminate theoretical models of language development and contribute to the prediction of communicative disorders. A qualitative, systematic review of this emergent literature illustrated the variety of approaches that have been used and highlighted some conceptual problems regarding the measurements. A quantitative analysis of the same data established that the bivariate relation was significant, with correlations of similar strength to those found for well-established nonlinguistic predictors of language. Further exploration of infant speech perception predictors, particularly from a methodological perspective, is recommended.
  • Dolscheid, S., Willems, R. M., Hagoort, P., & Casasanto, D. (2014). The relation of space and musical pitch in the brain. In P. Bello, M. Guarini, M. McShane, & B. Scassellati (Eds.), Proceedings of the 36th Annual Meeting of the Cognitive Science Society (CogSci 2014) (pp. 421-426). Austin, Tx: Cognitive Science Society.

    Abstract

    Numerous experiments show that space and musical pitch are
    closely linked in people's minds. However, the exact nature of
    space-pitch associations and their neuronal underpinnings are
    not well understood. In an fMRI experiment we investigated
    different types of spatial representations that may underlie
    musical pitch. Participants judged stimuli that varied in
    spatial height in both the visual and tactile modalities, as well
    as auditory stimuli that varied in pitch height. In order to
    distinguish between unimodal and multimodal spatial bases of
    musical pitch, we examined whether pitch activations were
    present in modality-specific (visual or tactile) versus
    multimodal (visual and tactile) regions active during spatial
    height processing. Judgments of musical pitch were found to
    activate unimodal visual areas, suggesting that space-pitch
    associations may involve modality-specific spatial
    representations, supporting a key assumption of embodied
    theories of metaphorical mental representation.
  • Guadalupe, T., Willems, R. M., Zwiers, M., Arias Vasquez, A., Hoogman, M., Hagoort, P., Fernández, G., Buitelaar, J., Franke, B., Fisher, S. E., & Francks, C. (2014). Differences in cerebral cortical anatomy of left- and right-handers. Frontiers in Psychology, 5: 261. doi:10.3389/fpsyg.2014.00261.

    Abstract

    The left and right sides of the human brain are specialized for different kinds of information processing, and much of our cognition is lateralized to an extent towards one side or the other. Handedness is a reflection of nervous system lateralization. Roughly ten percent of people are mixed- or left-handed, and they show an elevated rate of reductions or reversals of some cerebral functional asymmetries compared to right-handers. Brain anatomical correlates of left-handedness have also been suggested. However, the relationships of left-handedness to brain structure and function remain far from clear. We carried out a comprehensive analysis of cortical surface area differences between 106 left-handed subjects and 1960 right-handed subjects, measured using an automated method of regional parcellation (FreeSurfer, Destrieux atlas). This is the largest study sample that has so far been used in relation to this issue. No individual cortical region showed an association with left-handedness that survived statistical correction for multiple testing, although there was a nominally significant association with the surface area of a previously implicated region: the left precentral sulcus. Identifying brain structural correlates of handedness may prove useful for genetic studies of cerebral asymmetries, as well as providing new avenues for the study of relations between handedness, cerebral lateralization and cognition.
  • Guadalupe, T., Zwiers, M. P., Teumer, A., Wittfeld, K., Arias Vasquez, A., Hoogman, M., Hagoort, P., Fernández, G., Buitelaar, J., Hegenscheid, K., Völzke, H., Franke, B., Fisher, S. E., Grabe, H. J., & Francks, C. (2014). Measurement and genetics of human subcortical and hippocampal asymmetries in large datasets. Human Brain Mapping, 35(7), 3277-3289. doi:10.1002/hbm.22401.

    Abstract

    Functional and anatomical asymmetries are prevalent features of the human brain, linked to gender, handedness, and cognition. However, little is known about the neurodevelopmental processes involved. In zebrafish, asymmetries arise in the diencephalon before extending within the central nervous system. We aimed to identify genes involved in the development of subtle, left-right volumetric asymmetries of human subcortical structures using large datasets. We first tested the feasibility of measuring left-right volume differences in such large-scale samples, as assessed by two automated methods of subcortical segmentation (FSL|FIRST and FreeSurfer), using data from 235 subjects who had undergone MRI twice. We tested the agreement between the first and second scan, and the agreement between the segmentation methods, for measures of bilateral volumes of six subcortical structures and the hippocampus, and their volumetric asymmetries. We also tested whether there were biases introduced by left-right differences in the regional atlases used by the methods, by analyzing left-right flipped images. While many bilateral volumes were measured well (scan-rescan r = 0.6-0.8), most asymmetries, with the exception of the caudate nucleus, showed lower repeatabilites. We meta-analyzed genome-wide association scan results for caudate nucleus asymmetry in a combined sample of 3,028 adult subjects but did not detect associations at genome-wide significance (P < 5 × 10-8). There was no enrichment of genetic association in genes involved in left-right patterning of the viscera. Our results provide important information for researchers who are currently aiming to carry out large-scale genome-wide studies of subcortical and hippocampal volumes, and their asymmetries
  • Hagoort, P. (2014). Introduction to section on language and abstract thought. In M. S. Gazzaniga, & G. R. Mangun (Eds.), The cognitive neurosciences (5th ed., pp. 615-618). Cambridge, Mass: MIT Press.
  • Hagoort, P., & Levinson, S. C. (2014). Neuropragmatics. In M. S. Gazzaniga, & G. R. Mangun (Eds.), The cognitive neurosciences (5th ed., pp. 667-674). Cambridge, Mass: MIT Press.
  • Hagoort, P. (2014). Nodes and networks in the neural architecture for language: Broca's region and beyond. Current Opinion in Neurobiology, 28, 136-141. doi:10.1016/j.conb.2014.07.013.

    Abstract

    Current views on the neurobiological underpinnings of language are discussed that deviate in a number of ways from the classical Wernicke–Lichtheim–Geschwind model. More areas than Broca's and Wernicke's region are involved in language. Moreover, a division along the axis of language production and language comprehension does not seem to be warranted. Instead, for central aspects of language processing neural infrastructure is shared between production and comprehension. Three different accounts of the role of Broca's area in language are discussed. Arguments are presented in favor of a dynamic network view, in which the functionality of a region is co-determined by the network of regions in which it is embedded at particular moments in time. Finally, core regions of language processing need to interact with other networks (e.g. the attentional networks and the ToM network) to establish full functionality of language and communication.
  • Hagoort, P., & Indefrey, P. (2014). The neurobiology of language beyond single words. Annual Review of Neuroscience, 37, 347-362. doi:10.1146/annurev-neuro-071013-013847.

    Abstract

    A hallmark of human language is that we combine lexical building blocks retrieved from memory in endless new ways. This combinatorial aspect of language is referred to as unification. Here we focus on the neurobiological infrastructure for syntactic and semantic unification. Unification is characterized by a high-speed temporal profile including both prediction and integration of retrieved lexical elements. A meta-analysis of numerous neuroimaging studies reveals a clear dorsal/ventral gradient in both left inferior frontal cortex and left posterior temporal cortex, with dorsal foci for syntactic processing and ventral foci for semantic processing. In addition to core areas for unification, further networks need to be recruited to realize language-driven communication to its full extent. One example is the theory of mind network, which allows listeners and readers to infer the intended message (speaker meaning) from the coded meaning of the linguistic utterance. This indicates that sensorimotor simulation cannot handle all of language processing.
  • Heyselaar, E., Hagoort, P., & Segaert, K. (2014). In dialogue with an avatar, syntax production is identical compared to dialogue with a human partner. In P. Bello, M. Guarini, M. McShane, & B. Scassellati (Eds.), Proceedings of the 36th Annual Meeting of the Cognitive Science Society (CogSci 2014) (pp. 2351-2356). Austin, Tx: Cognitive Science Society.

    Abstract

    The use of virtual reality (VR) as a methodological tool is
    becoming increasingly popular in behavioural research due
    to its seemingly limitless possibilities. This new method has
    not been used frequently in the field of psycholinguistics,
    however, possibly due to the assumption that humancomputer
    interaction does not accurately reflect human-human
    interaction. In the current study we compare participants’
    language behaviour in a syntactic priming task with human
    versus avatar partners. Our study shows comparable priming
    effects between human and avatar partners (Human: 12.3%;
    Avatar: 12.6% for passive sentences) suggesting that VR is a
    valid platform for conducting language research and studying
    dialogue interactions.
  • Holler, J., Schubotz, L., Kelly, S., Hagoort, P., Schuetze, M., & Ozyurek, A. (2014). Social eye gaze modulates processing of speech and co-speech gesture. Cognition, 133, 692-697. doi:10.1016/j.cognition.2014.08.008.

    Abstract

    In human face-to-face communication, language comprehension is a multi-modal, situated activity. However, little is known about how we combine information from different modalities during comprehension, and how perceived communicative intentions, often signaled through visual signals, influence this process. We explored this question by simulating a multi-party communication context in which a speaker alternated her gaze between two recipients. Participants viewed speech-only or speech + gesture object-related messages when being addressed (direct gaze) or unaddressed (gaze averted to other participant). They were then asked to choose which of two object images matched the speaker’s preceding message. Unaddressed recipients responded significantly more slowly than addressees for speech-only utterances. However, perceiving the same speech accompanied by gestures sped unaddressed recipients up to a level identical to that of addressees. That is, when unaddressed recipients’ speech processing suffers, gestures can enhance the comprehension of a speaker’s message. We discuss our findings with respect to two hypotheses attempting to account for how social eye gaze may modulate multi-modal language comprehension.
  • Junge, C., Cutler, A., & Hagoort, P. (2014). Successful word recognition by 10-month-olds given continuous speech both at initial exposure and test. Infancy, 19(2), 179-193. doi:10.1111/infa.12040.

    Abstract

    Most words that infants hear occur within fluent speech. To compile a vocabulary, infants therefore need to segment words from speech contexts. This study is the first to investigate whether infants (here: 10-month-olds) can recognize words when both initial exposure and test presentation are in continuous speech. Electrophysiological evidence attests that this indeed occurs: An increased extended negativity (word recognition effect) appears for familiarized target words relative to control words. This response proved constant at the individual level: Only infants who showed this negativity at test had shown such a response, within six repetitions after first occurrence, during familiarization.
  • Levy, J., Hagoort, P., & Démonet, J.-F. (2014). A neuronal gamma oscillatory signature during morphological unification in the left occipitotemporal junction. Human Brain Mapping, 35, 5847-5860. doi:10.1002/hbm.22589.

    Abstract

    Morphology is the aspect of language concerned with the internal structure of words. In the past decades, a large body of masked priming (behavioral and neuroimaging) data has suggested that the visual word recognition system automatically decomposes any morphologically complex word into a stem and its constituent morphemes. Yet the reliance of morphology on other reading processes (e.g., orthography and semantics), as well as its underlying neuronal mechanisms are yet to be determined. In the current magnetoencephalography study, we addressed morphology from the perspective of the unification framework, that is, by applying the Hold/Release paradigm, morphological unification was simulated via the assembly of internal morphemic units into a whole word. Trials representing real words were divided into words with a transparent (true) or a nontransparent (pseudo) morphological relationship. Morphological unification of truly suffixed words was faster and more accurate and additionally enhanced induced oscillations in the narrow gamma band (60–85 Hz, 260–440 ms) in the left posterior occipitotemporal junction. This neural signature could not be explained by a mere automatic lexical processing (i.e., stem perception), but more likely it related to a semantic access step during the morphological unification process. By demonstrating the validity of unification at the morphological level, this study contributes to the vast empirical evidence on unification across other language processes. Furthermore, we point out that morphological unification relies on the retrieval of lexical semantic associations via induced gamma band oscillations in a cerebral hub region for visual word form processing.
  • Schoot, L., Menenti, L., Hagoort, P., & Segaert, K. (2014). A little more conversation - The influence of communicative context on syntactic priming in brain and behavior. Frontiers in Psychology, 5: 208. doi:10.3389/fpsyg.2014.00208.

    Abstract

    We report on an fMRI syntactic priming experiment in which we measure brain activity for participants who communicate with another participant outside the scanner. We investigated whether syntactic processing during overt language production and comprehension is influenced by having a (shared) goal to communicate. Although theory suggests this is true, the nature of this influence remains unclear. Two hypotheses are tested: i. syntactic priming effects (fMRI and RT) are stronger for participants in the communicative context than for participants doing the same experiment in a non-communicative context, and ii. syntactic priming magnitude (RT) is correlated with the syntactic priming magnitude of the speaker’s communicative partner. Results showed that across conditions, participants were faster to produce sentences with repeated syntax, relative to novel syntax. This behavioral result converged with the fMRI data: we found repetition suppression effects in the left insula extending into left inferior frontal gyrus (BA 47/45), left middle temporal gyrus (BA 21), left inferior parietal cortex (BA 40), left precentral gyrus (BA 6), bilateral precuneus (BA 7), bilateral supplementary motor cortex (BA 32/8) and right insula (BA 47). We did not find support for the first hypothesis: having a communicative intention does not increase the magnitude of syntactic priming effects (either in the brain or in behavior) per se. We did find support for the second hypothesis: if speaker A is strongly/weakly primed by speaker B, then speaker B is primed by speaker A to a similar extent. We conclude that syntactic processing is influenced by being in a communicative context, and that the nature of this influence is bi-directional: speakers are influenced by each other.
  • Segaert, K., Weber, K., Cladder-Micus, M., & Hagoort, P. (2014). The influence of verb-bound syntactic preferences on the processing of syntactic structures. Journal of Experimental Psychology: Learning, Memory, and Cognition, 40(5), 1448-1460. doi:10.1037/a0036796.

    Abstract

    Speakers sometimes repeat syntactic structures across sentences, a phenomenon called syntactic priming. We investigated the influence of verb-bound syntactic preferences on syntactic priming effects in response choices and response latencies for German ditransitive sentences. In the response choices we found inverse preference effects: There were stronger syntactic priming effects for primes in the less preferred structure, given the syntactic preference of the prime verb. In the response latencies we found positive preference effects: There were stronger syntactic priming effects for primes in the more preferred structure, given the syntactic preference of the prime verb. These findings provide further support for the idea that syntactic processing is lexically guided.
  • Simanova, I., Hagoort, P., Oostenveld, R., & Van Gerven, M. A. J. (2014). Modality-independent decoding of semantic information from the human brain. Cerebral Cortex, 24, 426-434. doi:10.1093/cercor/bhs324.

    Abstract

    An ability to decode semantic information from fMRI spatial patterns has been demonstrated in previous studies mostly for 1 specific input modality. In this study, we aimed to decode semantic category independent of the modality in which an object was presented. Using a searchlight method, we were able to predict the stimulus category from the data while participants performed a semantic categorization task with 4 stimulus modalities (spoken and written names, photographs, and natural sounds). Significant classification performance was achieved in all 4 modalities. Modality-independent decoding was implemented by training and testing the searchlight method across modalities. This allowed the localization of those brain regions, which correctly discriminated between the categories, independent of stimulus modality. The analysis revealed large clusters of voxels in the left inferior temporal cortex and in frontal regions. These voxels also allowed category discrimination in a free recall session where subjects recalled the objects in the absence of external stimuli. The results show that semantic information can be decoded from the fMRI signal independently of the input modality and have clear implications for understanding the functional mechanisms of semantic memory.
  • Stolk, A., Noordzij, M. L., Verhagen, L., Volman, I., Schoffelen, J.-M., Oostenveld, R., Hagoort, P., & Toni, I. (2014). Cerebral coherence between communicators marks the emergence of meaning. Proceedings of the National Academy of Sciences of the United States of America, 111, 18183-18188. doi:10.1073/pnas.1414886111.

    Abstract

    How can we understand each other during communicative interactions? An influential suggestion holds that communicators are primed by each other’s behaviors, with associative mechanisms automatically coordinating the production of communicative signals and the comprehension of their meanings. An alternative suggestion posits that mutual understanding requires shared conceptualizations of a signal’s use, i.e., “conceptual pacts” that are abstracted away from specific experiences. Both accounts predict coherent neural dynamics across communicators, aligned either to the occurrence of a signal or to the dynamics of conceptual pacts. Using coherence spectral-density analysis of cerebral activity simultaneously measured in pairs of communicators, this study shows that establishing mutual understanding of novel signals synchronizes cerebral dynamics across communicators’ right temporal lobes. This interpersonal cerebral coherence occurred only within pairs with a shared communicative history, and at temporal scales independent from signals’ occurrences. These findings favor the notion that meaning emerges from shared conceptualizations of a signal’s use.
  • Stolk, A., Noordzij, M. L., Volman, I., Verhagen, L., Overeem, S., van Elswijk, G., Bloem, B., Hagoort, P., & Toni, I. (2014). Understanding communicative actions: A repetitive TMS study. Cortex, 51, 25-34. doi:10.1016/j.cortex.2013.10.005.

    Abstract

    Despite the ambiguity inherent in human communication, people are remarkably efficient in establishing mutual understanding. Studying how people communicate in novel settings provides a window into the mechanisms supporting the human competence to rapidly generate and understand novel shared symbols, a fundamental property of human communication. Previous work indicates that the right posterior superior temporal sulcus (pSTS) is involved when people understand the intended meaning of novel communicative actions. Here, we set out to test whether normal functioning of this cerebral structure is required for understanding novel communicative actions using inhibitory low-frequency repetitive transcranial magnetic stimulation (rTMS). A factorial experimental design contrasted two tightly matched stimulation sites (right pSTS vs. left MT+, i.e. a contiguous homotopic task-relevant region) and tasks (a communicative task vs. a visual tracking task that used the same sequences of stimuli). Overall task performance was not affected by rTMS, whereas changes in task performance over time were disrupted according to TMS site and task combinations. Namely, rTMS over pSTS led to a diminished ability to improve action understanding on the basis of recent communicative history, while rTMS over MT+ perturbed improvement in visual tracking over trials. These findings qualify the contributions of the right pSTS to human communicative abilities, showing that this region might be necessary for incorporating previous knowledge, accumulated during interactions with a communicative partner, to constrain the inferential process that leads to action understanding.
  • Takashima, A., Wagensveld, B., Van Turennout, M., Zwitserlood, P., Hagoort, P., & Verhoeven, L. (2014). Training-induced neural plasticity in visual-word decoding and the role of syllables. Neuropsychologia, 61, 299-314. doi:10.1016/j.neuropsychologia.2014.06.017.

    Abstract

    To investigate the neural underpinnings of word decoding, and how it changes as a function of repeated exposure, we trained Dutch participants repeatedly over the course of a month of training to articulate a set of novel disyllabic input strings written in Greek script to avoid the use of familiar orthographic representations. The syllables in the input were phonotactically legal combinations but non-existent in the Dutch language, allowing us to assess their role in novel word decoding. Not only trained disyllabic pseudowords were tested but also pseudowords with recombined patterns of syllables to uncover the emergence of syllabic representations. We showed that with extensive training, articulation became faster and more accurate for the trained pseudowords. On the neural level, the initial stage of decoding was reflected by increased activity in visual attention areas of occipito-temporal and occipito-parietal cortices, and in motor coordination areas of the precentral gyrus and the inferior frontal gyrus. After one month of training, memory representations for holistic information (whole word unit) were established in areas encompassing the angular gyrus, the precuneus and the middle temporal gyrus. Syllabic representations also emerged through repeated training of disyllabic pseudowords, such that reading recombined syllables of the trained pseudowords showed similar brain activation to trained pseudowords and were articulated faster than novel combinations of letter strings used in the trained pseudowords.
  • Van Leeuwen, T. M., Petersson, K. M., Langner, O., Rijpkema, M., & Hagoort, P. (2014). Color specificity in the human V4 complex: An fMRI repetition suppression study. In T. D. Papageorgiou, G. I. Cristopoulous, & S. M. Smirnakis (Eds.), Advanced Brain Neuroimaging Topics in Health and Disease - Methods and Applications (pp. 275-295). Rijeka, Croatia: Intech. doi:10.5772/58278.
  • Van Leeuwen, T. M., Lamers, M. J. A., Petersson, K. M., Gussenhoven, C., Poser, B., & Hagoort, P. (2014). Phonological markers of information structure: An fMRI study. Neuropsychologia, 58(1), 64-74. doi:10.1016/j.neuropsychologia.2014.03.017.

    Abstract

    In this fMRI study we investigate the neural correlates of information structure integration during sentence comprehension in Dutch. We looked into how prosodic cues (pitch accents) that signal the information status of constituents to the listener (new information) are combined with other types of information during the unification process. The difficulty of unifying the prosodic cues into overall sentence meaning was manipulated by constructing sentences in which the pitch accent did (focus-accent agreement), and sentences in which the pitch accent did not (focus-accent disagreement) match the expectations for focus constituents of the sentence. In case of a mismatch, the load on unification processes increases. Our results show two anatomically distinct effects of focus-accent disagreement, one located in the posterior left inferior frontal gyrus (LIFG, BA6/44), and one in the more anterior-ventral LIFG (BA 47/45). Our results confirm that information structure is taken into account during unification, and imply an important role for the LIFG in unification processes, in line with previous fMRI studies.

    Additional information

    mmc1.doc
  • Bastiaansen, M. C. M., & Hagoort, P. (2003). Event-induced theta responses as a window on the dynamics of memory. Cortex, 39(4-5), 967-972. doi:10.1016/S0010-9452(08)70873-6.

    Abstract

    An important, but often ignored distinction in the analysis of EEG signals is that between evoked activity and induced activity. Whereas evoked activity reflects the summation of transient post-synaptic potentials triggered by an event, induced activity, which is mainly oscillatory in nature, is thought to reflect changes in parameters controlling dynamic interactions within and between brain structures. We hypothesize that induced activity may yield information about the dynamics of cell assembly formation, activation and subsequent uncoupling, which may play a prominent role in different types of memory operations. We then describe a number of analysis tools that can be used to study the reactivity of induced rhythmic activity, both in terms of amplitude changes and of phase variability.

    We briefly discuss how alpha, gamma and theta rhythms are thought to be generated, paying special attention to the hypothesis that the theta rhythm reflects dynamic interactions between the hippocampal system and the neocortex. This hypothesis would imply that studying the reactivity of scalp-recorded theta may provide a window on the contribution of the hippocampus to memory functions.

    We review studies investigating the reactivity of scalp-recorded theta in paradigms engaging episodic memory, spatial memory and working memory. In addition, we review studies that relate theta reactivity to processes at the interface of memory and language. Despite many unknowns, the experimental evidence largely supports the hypothesis that theta activity plays a functional role in cell assembly formation, a process which may constitute the neural basis of memory formation and retrieval. The available data provide only highly indirect support for the hypothesis that scalp-recorded theta yields information about hippocampal functioning. It is concluded that studying induced rhythmic activity holds promise as an additional important way to study brain function.
  • Hagoort, P., Wassenaar, M., & Brown, C. M. (2003). Syntax-related ERP-effects in Dutch. Cognitive Brain Research, 16(1), 38-50. doi:10.1016/S0926-6410(02)00208-2.

    Abstract

    In two studies subjects were required to read Dutch sentences that in some cases contained a syntactic violation, in other cases a semantic violation. All syntactic violations were word category violations. The design excluded differential contributions of expectancy to influence the syntactic violation effects. The syntactic violations elicited an Anterior Negativity between 300 and 500 ms. This negativity was bilateral and had a frontal distribution. Over posterior sites the same violations elicited a P600/SPS starting at about 600 ms. The semantic violations elicited an N400 effect. The topographic distribution of the AN was more frontal than the distribution of the classical N400 effect, indicating that the underlying generators of the AN and the N400 are, at least to a certain extent, non-overlapping. Experiment 2 partly replicated the design of Experiment 1, but with differences in rate of presentation and in the distribution of items over subjects, and without semantic violations. The word category violations resulted in the same effects as were observed in Experiment 1, showing that they were independent of some of the specific parameters of Experiment 1. The discussion presents a tentative account of the functional differences in the triggering conditions of the AN and the P600/SPS.
  • Hagoort, P., Wassenaar, M., & Brown, C. M. (2003). Real-time semantic compensation in patients with agrammatic comprehension: Electrophysiological evidence for multiple-route plasticity. Proceedings of the National Academy of Sciences of the United States of America, 100(7), 4340-4345. doi:10.1073/pnas.0230613100.

    Abstract

    To understand spoken language requires that the brain provides rapid access to different kinds of knowledge, including the sounds and meanings of words, and syntax. Syntax specifies constraints on combining words in a grammatically well formed manner. Agrammatic patients are deficient in their ability to use these constraints, due to a lesion in the perisylvian area of the languagedominant hemisphere. We report a study on real-time auditory sentence processing in agrammatic comprehenders, examining
    their ability to accommodate damage to the language system. We recorded event-related brain potentials (ERPs) in agrammatic comprehenders, nonagrammatic aphasics, and age-matched controls. When listening to sentences with grammatical violations, the agrammatic aphasics did not show the same syntax-related ERP effect as the two other subject groups. Instead, the waveforms of the agrammatic aphasics were dominated by a meaning-related ERP effect, presumably reflecting their attempts to achieve understanding by the use of semantic constraints. These data demonstrate that although agrammatic aphasics are impaired in their ability to exploit syntactic information in real time, they can reduce the consequences of a syntactic deficit by exploiting a semantic route. They thus provide evidence for the compensation of a syntactic deficit by a stronger reliance on another route in mapping
    sound onto meaning. This is a form of plasticity that we refer to as multiple-route plasticity.
  • Hagoort, P. (2003). De verloving tussen neurowetenschap en psychologie. In K. Hilberdink (Ed.), Interdisciplinariteit in de geesteswetenschappen (pp. 73-81). Amsterdam: KNAW.
  • Hagoort, P. (2003). Die einzigartige, grösstenteils aber unbewusste Fähigkeit der Menschen zu sprachlicher Kommunikation. In G. Kaiser (Ed.), Jahrbuch 2002-2003 / Wissenschaftszentrum Nordrhein-Westfalen (pp. 33-46). Düsseldorf: Wissenschaftszentrum Nordrhein-Westfalen.
  • Hagoort, P. (2003). Functional brain imaging. In W. J. Frawley (Ed.), International encyclopedia of linguistics (pp. 142-145). New York: Oxford University Press.
  • Hagoort, P. (2003). How the brain solves the binding problem for language: A neurocomputational model of syntactic processing. NeuroImage, 20(suppl. 1), S18-S29. doi:10.1016/j.neuroimage.2003.09.013.

    Abstract

    Syntax is one of the components in the architecture of language processing that allows the listener/reader to bind single-word information into a unified interpretation of multiword utterances. This paper discusses ERP effects that have been observed in relation to syntactic processing. The fact that these effects differ from the semantic N400 indicates that the brain honors the distinction between semantic and syntactic binding operations. Two models of syntactic processing attempt to account for syntax-related ERP effects. One type of model is serial, with a first phase that is purely syntactic in nature (syntax-first model). The other type of model is parallel and assumes that information immediately guides the interpretation process once it becomes available. This is referred to as the immediacy model. ERP evidence is presented in support of the latter model. Next, an explicit computational model is proposed to explain the ERP data. This Unification Model assumes that syntactic frames are stored in memory and retrieved on the basis of the spoken or written word form input. The syntactic frames associated with the individual lexical items are unified by a dynamic binding process into a structural representation that spans the whole utterance. On the basis of a meta-analysis of imaging studies on syntax, it is argued that the left posterior inferior frontal cortex is involved in binding syntactic frames together, whereas the left superior temporal cortex is involved in retrieval of the syntactic frames stored in memory. Lesion data that support the involvement of this left frontotemporal network in syntactic processing are discussed.
  • Hagoort, P. (2003). Interplay between syntax and semantics during sentence comprehension: ERP effects of combining syntactic and semantic violations. Journal of Cognitive Neuroscience, 15(6), 883-899. doi:10.1162/089892903322370807.

    Abstract

    This study investigated the effects of combined semantic and syntactic violations in relation to the effects of single semantic and single syntactic violations on language-related event-related brain potential (ERP) effects (N400 and P600/ SPS). Syntactic violations consisted of a mismatch in grammatical gender or number features of the definite article and the noun in sentence-internal or sentence-final noun phrases (NPs). Semantic violations consisted of semantically implausible adjective–noun combinations in the same NPs. Combined syntactic and semantic violations were a summation of these two respective violation types. ERPs were recorded while subjects read the sentences with the different types of violations and the correct control sentences. ERP effects were computed relative to ERPs elicited by the sentence-internal or sentence-final nouns. The size of the N400 effect to the semantic violation was increased by an additional syntactic violation (the syntactic boost). In contrast, the size of the P600/ SPS to the syntactic violation was not affected by an additional semantic violation. This suggests that in the absence of syntactic ambiguity, the assignment of syntactic structure is independent of semantic context. However, semantic integration is influenced by syntactic processing. In the sentence-final position, additional global processing consequences were obtained as a result of earlier violations in the sentence. The resulting increase in the N400 amplitude to sentence-final words was independent of the nature of the violation. A speeded anomaly detection task revealed that it takes substantially longer to detect semantic than syntactic anomalies. These results are discussed in relation to the latency and processing characteristics of the N400 and P600/SPS effects. Overall, the results reveal an asymmetry in the interplay between syntax and semantics during on-line sentence comprehension.
  • De Lange, F. P., Hagoort, P., & Toni, I. (2003). Differential fronto-parietal contributions to visual and motor imagery. NeuroImage, 19(2), e2094-e2095.

    Abstract

    Mental imagery is a cognitive process crucial to human reasoning. Numerous studies have characterized specific
    instances of this cognitive ability, as evoked by visual imagery (VI) or motor imagery (MI) tasks. However, it
    remains unclear which neural resources are shared between VI and MI, and which are exclusively related to MI.
    To address this issue, we have used fMRI to measure human brain activity during performance of VI and MI
    tasks. Crucially, we have modulated the imagery process by manipulating the degree of mental rotation necessary
    to solve the tasks. We focused our analysis on changes in neural signal as a function of the degree of mental
    rotation in each task.
  • Swaab, T., Brown, C. M., & Hagoort, P. (2003). Understanding words in sentence contexts: The time course of ambiguity resolution. Brain and Language, 86(2), 326-343. doi:10.1016/S0093-934X(02)00547-3.

    Abstract

    Spoken language comprehension requires rapid integration of information from multiple linguistic sources. In the present study we addressed the temporal aspects of this integration process by focusing on the time course of the selection of the appropriate meaning of lexical ambiguities (“bank”) in sentence contexts. Successful selection of the contextually appropriate meaning of the ambiguous word is dependent upon the rapid binding of the contextual information in the sentence to the appropriate meaning of the ambiguity. We used the N400 to identify the time course of this binding process. The N400 was measured to target words that followed three types of context sentences. In the concordant context, the sentence biased the meaning of the sentence-final ambiguous word so that it was related to the target. In the discordant context, the sentence context biased the meaning so that it was not related to the target. In the unrelated control condition, the sentences ended in an unambiguous noun that was unrelated to the target. Half of the concordant sentences biased the dominant meaning, and the other half biased the subordinate meaning of the sentence-final ambiguous words. The ISI between onset of the target word and offset of the sentence-final word of the context sentence was 100 ms in one version of the experiment, and 1250 ms in the second version. We found that (i) the lexically dominant meaning is always partly activated, independent of context, (ii) initially both dominant and subordinate meaning are (partly) activated, which suggests that contextual and lexical factors both contribute to sentence interpretation without context completely overriding lexical information, and (iii) strong lexical influences remain present for a relatively long period of time.
  • Van Berkum, J. J. A., Zwitserlood, P., Hagoort, P., & Brown, C. M. (2003). When and how do listeners relate a sentence to the wider discourse? Evidence from the N400 effect. Cognitive Brain Research, 17(3), 701-718. doi:10.1016/S0926-6410(03)00196-4.

    Abstract

    In two ERP experiments, we assessed the impact of discourse-level information on the processing of an unfolding spoken sentence. Subjects listened to sentences like Jane told her brother that he was exceptionally quick/slow, designed such that the alternative critical words were equally acceptable within the local sentence context. In Experiment 1, these sentences were embedded in a discourse that rendered one of the critical words anomalous (e.g. because Jane’s brother had in fact done something very quickly). Relative to the coherent alternative, these discourse-anomalous words elicited a standard N400 effect that started at 150–200 ms after acoustic word onset. Furthermore, when the same sentences were heard in isolation in Experiment 2, the N400 effect disappeared. The results demonstrate that our listeners related the unfolding spoken words to the wider discourse extremely rapidly, after having heard the first two or three phonemes only, and in many cases well before the end of the word. In addition, the identical nature of discourse- and sentence-dependent N400 effects suggests that from the perspective of the word-elicited comprehension process indexed by the N400, the interpretive context delineated by a single unfolding sentence and a larger discourse is functionally identical.
  • Van Turennout, M., Schmitt, B., & Hagoort, P. (2003). When words come to mind: Electrophysiological insights on the time course of speaking and understanding words. In N. O. Schiller, & A. S. Meyer (Eds.), Phonetics and phonology in language comprehension and production: Differences and similarities (pp. 241-278). Berlin: Mouton de Gruyter.
  • Van Berkum, J. J. A., Brown, C. M., Hagoort, P., & Zwitserlood, P. (2003). Event-related brain potentials reflect discourse-referential ambiguity in spoken language comprehension. Psychophysiology, 40(2), 235-248. doi:10.1111/1469-8986.00025.

    Abstract

    In two experiments, we explored the use of event-related brain potentials to selectively track the processes that establish reference during spoken language comprehension. Subjects listened to stories in which a particular noun phrase like "the girl" either uniquely referred to a single referent mentioned in the earlier discourse, or ambiguously referred to two equally suitable referents. Referentially ambiguous nouns ("the girl" with two girls introduced in the discourse context) elicited a frontally dominant and sustained negative shift in brain potentials, emerging within 300–400 ms after acoustic noun onset. The early onset of this effect reveals that reference to a discourse entity can be established very rapidly. Its morphology and distribution suggest that at least some of the processing consequences of referential ambiguity may involve an increased demand on memory resources. Furthermore, because this referentially induced ERP effect is very different from that of well-known ERP effects associated with the semantic (N400) and syntactic (e.g., P600/SPS) aspects of language comprehension, it suggests that ERPs can be used to selectively keep track of three major processes involved in the comprehension of an unfolding piece of discourse.

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