Peter Hagoort

Publications

Displaying 1 - 41 of 41
  • 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

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  • Acheson, D. J., & Hagoort, P. (2013). Stimulating the brain's language network: Syntactic ambiguity resolution after TMS to the IFG and MTG. Journal of Cognitive Neuroscience, 25(10), 1664-1677. doi:10.1162/jocn_a_00430.

    Abstract

    The posterior middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) are two critical nodes of the brain's language network. Previous neuroimaging evidence has supported a dissociation in language comprehension in which parts of the MTG are involved in the retrieval of lexical syntactic information and the IFG is involved in unification operations that maintain, select, and integrate multiple sources of information over time. In the present investigation, we tested for causal evidence of this dissociation by modulating activity in IFG and MTG using an offline TMS procedure: continuous theta-burst stimulation. Lexical–syntactic retrieval was manipulated by using sentences with and without a temporarily word-class (noun/verb) ambiguity (e.g., run). In one group of participants, TMS was applied to the IFG and MTG, and in a control group, no TMS was applied. Eye movements were recorded and quantified at two critical sentence regions: a temporarily ambiguous region and a disambiguating region. Results show that stimulation of the IFG led to a modulation of the ambiguity effect (ambiguous–unambiguous) at the disambiguating sentence region in three measures: first fixation durations, total reading times, and regressive eye movements into the region. Both IFG and MTG stimulation modulated the ambiguity effect for total reading times in the temporarily ambiguous sentence region relative to a control group. The current results demonstrate that an offline repetitive TMS protocol can have influences at a different point in time during online processing and provide causal evidence for IFG involvement in unification operations during sentence comprehension.
  • Hagoort, P. (2013). MUC (Memory, Unification, Control) and beyond. Frontiers in Psychology, 4: 416. doi:10.3389/fpsyg.2013.00416.

    Abstract

    A neurobiological model of language is discussed that overcomes the shortcomings of the classical Wernicke-Lichtheim-Geschwind model. It is based on a subdivision of language processing into three components: Memory, Unification, and Control. The functional components as well as the neurobiological underpinnings of the model are discussed. In addition, the need for extension of the model beyond the classical core regions for language is shown. Attentional networks as well as networks for inferential processing are crucial to realize language comprehension beyond single word processing and beyond decoding propositional content. It is shown that this requires the dynamic interaction between multiple brain regions.
  • Hagoort, P., & Poeppel, D. (2013). The infrastructure of the language-ready brain. In M. A. Arbib (Ed.), Language, music, and the brain: A mysterious relationship (pp. 233-255). Cambridge, MA: MIT Press.

    Abstract

    This chapter sketches in very general terms the cognitive architecture of both language comprehension and production, as well as the neurobiological infrastructure that makes the human brain ready for language. Focus is on spoken language, since that compares most directly to processing music. It is worth bearing in mind that humans can also interface with language as a cognitive system using sign and text (visual) as well as Braille (tactile); that is to say, the system can connect with input/output processes in any sensory modality. Language processing consists of a complex and nested set of subroutines to get from sound to meaning (in comprehension) or meaning to sound (in production), with remarkable speed and accuracy. The fi rst section outlines a selection of the major constituent operations, from fractionating the input into manageable units to combining and unifying information in the construction of meaning. The next section addresses the neurobiological infrastructure hypothesized to form the basis for language processing. Principal insights are summarized by building on the notion of “brain networks” for speech–sound processing, syntactic processing, and the construction of meaning, bearing in mind that such a neat three-way subdivision overlooks important overlap and shared mechanisms in the neural architecture subserving language processing. Finally, in keeping with the spirit of the volume, some possible relations are highlighted between language and music that arise from the infrastructure developed here. Our characterization of language and its neurobiological foundations is necessarily selective and brief. Our aim is to identify for the reader critical questions that require an answer to have a plausible cognitive neuroscience of language processing.
  • Hagoort, P., & Meyer, A. S. (2013). What belongs together goes together: the speaker-hearer perspective. A commentary on MacDonald's PDC account. Frontiers in Psychology, 4: 228. doi:10.3389/fpsyg.2013.00228.

    Abstract

    First paragraph:
    MacDonald (2013) proposes that distributional properties of language and processing biases in language comprehension can to a large extent be attributed to consequences of the language production process. In essence, the account is derived from the principle of least effort that was formulated by Zipf, among others (Zipf, 1949; Levelt, 2013). However, in Zipf's view the outcome of the least effort principle was a compromise between least effort for the speaker and least effort for the listener, whereas MacDonald puts most of the burden on the production process.
  • Holler, J., Schubotz, L., Kelly, S., Schuetze, M., Hagoort, P., & Ozyurek, A. (2013). Here's not looking at you, kid! Unaddressed recipients benefit from co-speech gestures when speech processing suffers. In M. Knauff, M. Pauen, I. Sebanz, & I. Wachsmuth (Eds.), Proceedings of the 35th Annual Meeting of the Cognitive Science Society (CogSci 2013) (pp. 2560-2565). Austin, TX: Cognitive Science Society. Retrieved from http://mindmodeling.org/cogsci2013/papers/0463/index.html.

    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 these different modalities, and how perceived communicative intentions, often signaled through visual signals, such as eye
    gaze, may influence this processing. We address this question by simulating a triadic communication context in which a
    speaker alternated her gaze between two different recipients. Participants thus viewed speech-only or speech+gesture
    object-related utterances when being addressed (direct gaze) or unaddressed (averted gaze). Two object images followed
    each message and participants’ task was to choose the object that matched the message. Unaddressed recipients responded significantly slower than addressees for speech-only
    utterances. However, perceiving the same speech accompanied by gestures sped them up to a level identical to
    that of addressees. That is, when speech processing suffers due to not being addressed, gesture processing remains intact and enhances the comprehension of a speaker’s message
  • Kooijman, V., Junge, C., Johnson, E. K., Hagoort, P., & Cutler, A. (2013). Predictive brain signals of linguistic development. Frontiers in Psychology, 4: 25. doi:10.3389/fpsyg.2013.00025.

    Abstract

    The ability to extract word forms from continuous speech is a prerequisite for constructing a vocabulary and emerges in the first year of life. Electrophysiological (ERP) studies of speech segmentation by 9- to 12-month-old listeners in several languages have found a left-localized negativity linked to word onset as a marker of word detection. We report an ERP study showing significant evidence of speech segmentation in Dutch-learning 7-month-olds. In contrast to the left-localized negative effect reported with older infants, the observed overall mean effect had a positive polarity. Inspection of individual results revealed two participant sub-groups: a majority showing a positive-going response, and a minority showing the left negativity observed in older age groups. We retested participants at age three, on vocabulary comprehension and word and sentence production. On every test, children who at 7 months had shown the negativity associated with segmentation of words from speech outperformed those who had produced positive-going brain responses to the same input. The earlier that infants show the left-localized brain responses typically indicating detection of words in speech, the better their early childhood language skills.
  • Kristensen, L. B., Wang, L., Petersson, K. M., & Hagoort, P. (2013). The interface between language and attention: Prosodic focus marking recruits a general attention network in spoken language comprehension. Cerebral Cortex, 23, 1836-1848. doi:10.1093/cercor/bhs164.

    Abstract

    In spoken language, pitch accent can mark certain information as focus, whereby more attentional resources are allocated to the focused information. Using functional magnetic resonance imaging, this study examined whether pitch accent, used for marking focus, recruited general attention networks during sentence comprehension. In a language task, we independently manipulated the prosody and semantic/pragmatic congruence of sentences. We found that semantic/pragmatic processing affected bilateral inferior and middle frontal gyrus. The prosody manipulation showed bilateral involvement of the superior/inferior parietal cortex, superior and middle temporal cortex, as well as inferior, middle, and posterior parts of the frontal cortex. We compared these regions with attention networks localized in an auditory spatial attention task. Both tasks activated bilateral superior/inferior parietal cortex, superior temporal cortex, and left precentral cortex. Furthermore, an interaction between prosody and congruence was observed in bilateral inferior parietal regions: for incongruent sentences, but not for congruent ones, there was a larger activation if the incongruent word carried a pitch accent, than if it did not. The common activations between the language task and the spatial attention task demonstrate that pitch accent activates a domain general attention network, which is sensitive to semantic/pragmatic aspects of language. Therefore, attention and language comprehension are highly interactive.

    Additional information

    Kirstensen_Cer_Cor_Suppl_Mat.doc
  • Meyer, A. S., & Hagoort, P. (2013). What does it mean to predict one's own utterances? [Commentary on Pickering & Garrod]. Behavioral and Brain Sciences, 36, 367-368. doi:10.1017/S0140525X12002786.

    Abstract

    Many authors have recently highlighted the importance of prediction for language comprehension. Pickering & Garrod (P&G) are the first to propose a central role for prediction in language production. This is an intriguing idea, but it is not clear what it means for speakers to predict their own utterances, and how prediction during production can be empirically distinguished from production proper.
  • Peeters, D., Chu, M., Holler, J., Ozyurek, A., & Hagoort, P. (2013). Getting to the point: The influence of communicative intent on the kinematics of pointing gestures. In M. Knauff, M. Pauen, N. Sebanz, & I. Wachsmuth (Eds.), Proceedings of the 35th Annual Meeting of the Cognitive Science Society (CogSci 2013) (pp. 1127-1132). Austin, TX: Cognitive Science Society.

    Abstract

    In everyday communication, people not only use speech but
    also hand gestures to convey information. One intriguing
    question in gesture research has been why gestures take the
    specific form they do. Previous research has identified the
    speaker-gesturer’s communicative intent as one factor
    shaping the form of iconic gestures. Here we investigate
    whether communicative intent also shapes the form of
    pointing gestures. In an experimental setting, twenty-four
    participants produced pointing gestures identifying a referent
    for an addressee. The communicative intent of the speakergesturer
    was manipulated by varying the informativeness of
    the pointing gesture. A second independent variable was the
    presence or absence of concurrent speech. As a function of their communicative intent and irrespective of the presence of speech, participants varied the durations of the stroke and the post-stroke hold-phase of their gesture. These findings add to our understanding of how the communicative context influences the form that a gesture takes.
  • Segaert, K., Kempen, G., Petersson, K. M., & Hagoort, P. (2013). Syntactic priming and the lexical boost effect during sentence production and sentence comprehension: An fMRI study. Brain and Language, 124, 174-183. doi:10.1016/j.bandl.2012.12.003.

    Abstract

    Behavioral syntactic priming effects during sentence comprehension are typically observed only if both the syntactic structure and lexical head are repeated. In contrast, during production syntactic priming occurs with structure repetition alone, but the effect is boosted by repetition of the lexical head. We used fMRI to investigate the neuronal correlates of syntactic priming and lexical boost effects during sentence production and comprehension. The critical measure was the magnitude of fMRI adaptation to repetition of sentences in active or passive voice, with or without verb repetition. In conditions with repeated verbs, we observed adaptation to structure repetition in the left IFG and MTG, for active and passive voice. However, in the absence of repeated verbs, adaptation occurred only for passive sentences. None of the fMRI adaptation effects yielded differential effects for production versus comprehension, suggesting that sentence comprehension and production are subserved by the same neuronal infrastructure for syntactic processing.

    Additional information

    Segaert_Supplementary_data_2013.docx
  • Segaert, K., Weber, K., De Lange, F., Petersson, K. M., & Hagoort, P. (2013). The suppression of repetition enhancement: A review of fMRI studies. Neuropsychologia, 51, 59-66. doi:10.1016/j.neuropsychologia.2012.11.006.

    Abstract

    Repetition suppression in fMRI studies is generally thought to underlie behavioural facilitation effects (i.e., priming) and it is often used to identify the neuronal representations associated with a stimulus. However, this pays little heed to the large number of repetition enhancement effects observed under similar conditions. In this review, we identify several cognitive variables biasing repetition effects in the BOLD response towards enhancement instead of suppression. These variables are stimulus recognition, learning, attention, expectation and explicit memory. We also evaluate which models can account for these repetition effects and come to the conclusion that there is no one single model that is able to embrace all repetition enhancement effects. Accumulation, novel network formation as well as predictive coding models can all explain subsets of repetition enhancement effects.
  • Stolk, A., Verhagen, L., Schoffelen, J.-M., Oostenveld, R., Blokpoel, M., Hagoort, P., van Rooij, I., & Tonia, I. (2013). Neural mechanisms of communicative innovation. Proceedings of the National Academy of Sciences of the United States of America, 110(36), 14574-14579. doi:10.1073/pnas.1303170110.

    Abstract

    Human referential communication is often thought as coding-decoding a set of symbols, neglecting that establishing shared meanings requires a computational mechanism powerful enough to mutually negotiate them. Sharing the meaning of a novel symbol might rely on similar conceptual inferences across communicators or on statistical similarities in their sensorimotor behaviors. Using magnetoencephalography, we assess spectral, temporal, and spatial characteristics of neural activity evoked when people generate and understand novel shared symbols during live communicative interactions. Solving those communicative problems induced comparable changes in the spectral profile of neural activity of both communicators and addressees. This shared neuronal up-regulation was spatially localized to the right temporal lobe and the ventromedial prefrontal cortex and emerged already before the occurrence of a specific communicative problem. Communicative innovation relies on neuronal computations that are shared across generating and understanding novel shared symbols, operating over temporal scales independent from transient sensorimotor behavior.
  • Thompson-Schill, S., Hagoort, P., Dominey, P. F., Honing, H., Koelsch, S., Ladd, D. R., Lerdahl, F., Levinson, S. C., & Steedman, M. (2013). Multiple levels of structure in language and music. In M. A. Arbib (Ed.), Language, music, and the brain: A mysterious relationship (pp. 289-303). Cambridge, MA: MIT Press.

    Abstract

    A forum devoted to the relationship between music and language begins with an implicit assumption: There is at least one common principle that is central to all human musical systems and all languages, but that is not characteristic of (most) other domains. Why else should these two categories be paired together for analysis? We propose that one candidate for a common principle is their structure. In this chapter, we explore the nature of that structure—and its consequences for psychological and neurological processing mechanisms—within and across these two domains.
  • Van Leeuwen, T. M., Hagoort, P., & Händel, B. F. (2013). Real color captures attention and overrides spatial cues in grapheme-color synesthetes but not in controls. Neuropsychologia, 51(10), 1802-1813. doi:10.1016/j.neuropsychologia.2013.06.024.

    Abstract

    Grapheme-color synesthetes perceive color when reading letters or digits. We investigated oscillatory brain signals of synesthetes vs. controls using magnetoencephalography. Brain oscillations specifically in the alpha band (∼10 Hz) have two interesting features: alpha has been linked to inhibitory processes and can act as a marker for attention. The possible role of reduced inhibition as an underlying cause of synesthesia, as well as the precise role of attention in synesthesia is widely discussed. To assess alpha power effects due to synesthesia, synesthetes as well as matched controls viewed synesthesia-inducing graphemes, colored control graphemes, and non-colored control graphemes while brain activity was recorded. Subjects had to report a color change at the end of each trial which allowed us to assess the strength of synesthesia in each synesthete. Since color (synesthetic or real) might allocate attention we also included an attentional cue in our paradigm which could direct covert attention. In controls the attentional cue always caused a lateralization of alpha power with a contralateral decrease and ipsilateral alpha increase over occipital sensors. In synesthetes, however, the influence of the cue was overruled by color: independent of the attentional cue, alpha power decreased contralateral to the color (synesthetic or real). This indicates that in synesthetes color guides attention. This was confirmed by reaction time effects due to color, i.e. faster RTs for the color side independent of the cue. Finally, the stronger the observed color dependent alpha lateralization, the stronger was the manifestation of synesthesia as measured by congruency effects of synesthetic colors on RTs. Behavioral and imaging results indicate that color induces a location-specific, automatic shift of attention towards color in synesthetes but not in controls. We hypothesize that this mechanism can facilitate coupling of grapheme and color during the development of synesthesia.
  • Wagensveld, B., Van Alphen, P. M., Segers, E., Hagoort, P., & Verhoeven, L. (2013). The neural correlates of rhyme awareness in preliterate and literate children. Clinical Neurophysiology, 124, 1336-1345. doi:10.1016/j.clinph.2013.01.022.

    Abstract

    Objective Most rhyme awareness assessments do not encompass measures of the global similarity effect (i.e., children who are able to perform simple rhyme judgments get confused when presented with globally similar non-rhyming pairs). The present study examines the neural nature of this effect by studying the N450 rhyme effect. Methods Behavioral and electrophysiological responses of Dutch pre-literate kindergartners and literate second graders were recorded while they made rhyme judgments of word pairs in three conditions; phonologically rhyming (e.g., wijn-pijn), overlapping non-rhyming (e.g., pen-pijn) and unrelated non-rhyming pairs (e.g., boom-pijn). Results Behaviorally, both groups had difficulty judging overlapping but not rhyming and unrelated pairs. The neural data of second graders showed overlapping pairs were processed in a similar fashion as unrelated pairs; both showed a more negative deflection of the N450 component than rhyming items. Kindergartners did not show a typical N450 rhyme effect. However, some other interesting ERP differences were observed, indicating preliterates are sensitive to rhyme at a certain level. Significance Rhyme judgments of globally similar items rely on the same process as rhyme judgments of rhyming and unrelated items. Therefore, incorporating a globally similar condition in rhyme assessments may lead to a more in-depth measure of early phonological awareness skills. Highlights Behavioral and electrophysiological responses were recorded while (pre)literate children made rhyme judgments of rhyming, overlapping and unrelated words. Behaviorally both groups had difficulty judging overlapping pairs as non-rhyming while overlapping and unrelated neural patterns were similar in literates. Preliterates show a different pattern indicating a developing phonological system.
  • Wang, L., Bastiaansen, M. C. M., Yang, Y., & Hagoort, P. (2013). ERP evidence on the interaction between information structure and emotional salience of words. Cognitive, Affective and Behavioral Neuroscience, 13, 297-310. doi:10.3758/s13415-012-0146-2.

    Abstract

    Both emotional words and words focused by information structure can capture attention. This study examined the interplay between emotional salience and information structure in modulating attentional resources in the service of integrating emotional words into sentence context. Event-related potentials (ERPs) to affectively negative, neutral, and positive words, which were either focused or nonfocused in question–answer pairs, were evaluated during sentence comprehension. The results revealed an early negative effect (90–200 ms), a P2 effect, as well as an effect in the N400 time window, for both emotional salience and information structure. Moreover, an interaction between emotional salience and information structure occurred within the N400 time window over right posterior electrodes, showing that information structure influences the semantic integration only for neutral words, but not for emotional words. This might reflect the fact that the linguistic salience of emotional words can override the effect of information structure on the integration of words into context. The interaction provides evidence for attention–emotion interactions at a later stage of processing. In addition, the absence of interaction in the early time window suggests that the processing of emotional information is highly automatic and independent of context. The results suggest independent attention capture systems of emotional salience and information structure at the early stage but an interaction between them at a later stage, during the semantic integration of words.
  • Wang, L., Zhu, Z., Bastiaansen, M. C. M., Hagoort, P., & Yang, Y. (2013). Recognizing the emotional valence of names: An ERP study. Brain and Language, 125, 118-127. doi:10.1016/j.bandl.2013.01.006.

    Abstract

    Unlike common nouns, person names refer to unique entities and generally have a referring function. We used event-related potentials to investigate the time course of identifying the emotional meaning of nouns and names. The emotional valence of names and nouns were manipulated separately. The results show early N1 effects in response to emotional valence only for nouns. This might reflect automatic attention directed towards emotional stimuli. The absence of such an effect for names supports the notion that the emotional meaning carried by names is accessed after word recognition and person identification. In addition, both names with negative valence and emotional nouns elicited late positive effects, which have been associated with evaluation of emotional significance. This positive effect started earlier for nouns than for names, but with similar durations. Our results suggest that distinct neural systems are involved in the retrieval of names’ and nouns’ emotional meaning.

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