Publications

Displaying 1 - 96 of 96
  • Aparicio, X., Heidlmayr, K., & Isel, F. (2017). Inhibition Efficiency in Highly Proficient Bilinguals and Simultaneous Interpreters: Evidence from Language Switching and Stroop Tasks. Journal of Psycholinguistic Research, 46, 1427-1451. doi:10.1007/s10936-017-9501-3.

    Abstract

    The present behavioral study aimed to examine the impact of language control expertise on two domain-general control processes, i.e. active inhibition of competing representations and overcoming of inhibition. We compared how Simultaneous Interpreters (SI) and Highly Proficient Bilinguals—two groups assumed to differ in language control capacity—performed executive tasks involving specific inhibition processes. In Experiment 1 (language decision task), both active and overcoming of inhibition processes are involved, while in Experiment 2 (bilingual Stroop task) only interference suppression is supposed to be required. The results of Experiment 1 showed a language switching effect only for the highly proficient bilinguals, potentially because overcoming of inhibition requires more cognitive resources than in SI. Nevertheless, both groups performed similarly on the Stroop task in Experiment 2, which suggests that active inhibition may work similarly in both groups. These contrasting results suggest that overcoming of inhibition may be harder to master than active inhibition. Taken together, these data indicate that some executive control processes may be less sensitive to the degree of expertise in bilingual language control than others. Our findings lend support to psycholinguistic models of bilingualism postulating a higher-order mechanism regulating language activation.
  • Armeni, K., Willems, R. M., & Frank, S. (2017). Probabilistic language models in cognitive neuroscience: Promises and pitfalls. Neuroscience and Biobehavioral Reviews, 83, 579-588. doi:10.1016/j.neubiorev.2017.09.001.

    Abstract

    Cognitive neuroscientists of language comprehension study how neural computations relate to cognitive computations during comprehension. On the cognitive part of the equation, it is important that the computations and processing complexity are explicitly defined. Probabilistic language models can be used to give a computationally explicit account of language complexity during comprehension. Whereas such models have so far predominantly been evaluated against behavioral data, only recently have the models been used to explain neurobiological signals. Measures obtained from these models emphasize the probabilistic, information-processing view of language understanding and provide a set of tools that can be used for testing neural hypotheses about language comprehension. Here, we provide a cursory review of the theoretical foundations and example neuroimaging studies employing probabilistic language models. We highlight the advantages and potential pitfalls of this approach and indicate avenues for future research
  • De Boer, M., Kokal, I., Blokpoel, M., Liu, R., Stolk, A., Roelofs, K., Van Rooij, I., & Toni, I. (2017). Oxytocin modulates human communication by enhancing cognitive exploration. Psychoneuroendocrinology, 86, 64-72. doi:10.1016/j.psyneuen.2017.09.010.

    Abstract

    Oxytocin is a neuropeptide known to influence how humans share material resources. Here we explore whether oxytocin influences how we share knowledge. We focus on two distinguishing features of human communication, namely the ability to select communicative signals that disambiguate the many-to-many mappings that exist between a signal’s form and meaning, and adjustments of those signals to the presumed cognitive characteristics of the addressee (“audience design”). Fifty-five males participated in a randomized, double-blind, placebo controlled experiment involving the intranasal administration of oxytocin. The participants produced novel non-verbal communicative signals towards two different addressees, an adult or a child, in an experimentally-controlled live interactive setting. We found that oxytocin administration drives participants to generate signals of higher referential quality, i.e. signals that disambiguate more communicative problems; and to rapidly adjust those communicative signals to what the addressee understands. The combined effects of oxytocin on referential quality and audience design fit with the notion that oxytocin administration leads participants to explore more pervasively behaviors that can convey their intention, and diverse models of the addressees. These findings suggest that, besides affecting prosocial drive and salience of social cues, oxytocin influences how we share knowledge by promoting cognitive exploration
  • Bosker, H. R., & Kösem, A. (2017). An entrained rhythm's frequency, not phase, influences temporal sampling of speech. In Proceedings of Interspeech 2017 (pp. 2416-2420). doi:10.21437/Interspeech.2017-73.

    Abstract

    Brain oscillations have been shown to track the slow amplitude fluctuations in speech during comprehension. Moreover, there is evidence that these stimulus-induced cortical rhythms may persist even after the driving stimulus has ceased. However, how exactly this neural entrainment shapes speech perception remains debated. This behavioral study investigated whether and how the frequency and phase of an entrained rhythm would influence the temporal sampling of subsequent speech. In two behavioral experiments, participants were presented with slow and fast isochronous tone sequences, followed by Dutch target words ambiguous between as /ɑs/ “ash” (with a short vowel) and aas /a:s/ “bait” (with a long vowel). Target words were presented at various phases of the entrained rhythm. Both experiments revealed effects of the frequency of the tone sequence on target word perception: fast sequences biased listeners to more long /a:s/ responses. However, no evidence for phase effects could be discerned. These findings show that an entrained rhythm’s frequency, but not phase, influences the temporal sampling of subsequent speech. These outcomes are compatible with theories suggesting that sensory timing is evaluated relative to entrained frequency. Furthermore, they suggest that phase tracking of (syllabic) rhythms by theta oscillations plays a limited role in speech parsing.
  • Bouhali, F., Mongelli, V., & Cohen, L. (2017). Musical literacy shifts asymmetries in the ventral visual cortex. NeuroImage, 156, 445-455. doi:10.1016/j.neuroimage.2017.04.027.

    Abstract

    The acquisition of literacy has a profound impact on the functional specialization and lateralization of the visual cortex. Due to the overall lateralization of the language network, specialization for printed words develops in the left occipitotemporal cortex, allegedly inducing a secondary shift of visual face processing to the right, in literate as compared to illiterate subjects. Applying the same logic to the acquisition of high-level musical literacy, we predicted that, in musicians as compared to non-musicians, occipitotemporal activations should show a leftward shift for music reading, and an additional rightward push for face perception. To test these predictions, professional musicians and non-musicians viewed pictures of musical notation, faces, words, tools and houses in the MRI, and laterality was assessed in the ventral stream combining ROI and voxel-based approaches. The results supported both predictions, and allowed to locate the leftward shift to the inferior temporal gyrus and the rightward shift to the fusiform cortex. Moreover, these laterality shifts generalized to categories other than music and faces. Finally, correlation measures across subjects did not support a causal link between the leftward and rightward shifts. Thus the acquisition of an additional perceptual expertise extensively modifies the laterality pattern in the visual system

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  • Coco, M. I., Araujo, S., & Petersson, K. M. (2017). Disentangling stimulus plausibility and contextual congruency: Electro-physiological evidence for differential cognitive dynamics. Neuropsychologia, 96, 150-163. doi:10.1016/j.neuropsychologia.2016.12.008.

    Abstract

    Expectancy mechanisms are routinely used by the cognitive system in stimulus processing and in anticipation of appropriate responses. Electrophysiology research has documented negative shifts of brain activity when expectancies are violated within a local stimulus context (e.g., reading an implausible word in a sentence) or more globally between consecutive stimuli (e.g., a narrative of images with an incongruent end). In this EEG study, we examine the interaction between expectancies operating at the level of stimulus plausibility and at more global level of contextual congruency to provide evidence for, or against, a disassociation of the underlying processing mechanisms. We asked participants to verify the congruency of pairs of cross-modal stimuli (a sentence and a scene), which varied in plausibility. ANOVAs on ERP amplitudes in selected windows of interest show that congruency violation has longer-lasting (from 100 to 500 ms) and more widespread effects than plausibility violation (from 200 to 400 ms). We also observed critical interactions between these factors, whereby incongruent and implausible pairs elicited stronger negative shifts than their congruent counterpart, both early on (100–200 ms) and between 400–500 ms. Our results suggest that the integration mechanisms are sensitive to both global and local effects of expectancy in a modality independent manner. Overall, we provide novel insights into the interdependence of expectancy during meaning integration of cross-modal stimuli in a verification task
  • Dai, B., McQueen, J. M., Hagoort, P., & Kösem, A. (2017). Pure linguistic interference during comprehension of competing speech signals. The Journal of the Acoustical Society of America, 141, EL249-EL254. doi:10.1121/1.4977590.

    Abstract

    Speech-in-speech perception can be challenging because the processing of competing acoustic and linguistic information leads to informational masking. Here, a method is proposed to isolate the linguistic component of informational masking while keeping the distractor's acoustic information unchanged. Participants performed a dichotic listening cocktail-party task before and after training on 4-band noise-vocoded sentences that became intelligible through the training. Distracting noise-vocoded speech interfered more with target speech comprehension after training (i.e., when intelligible) than before training (i.e., when unintelligible) at −3 dB SNR. These findings confirm that linguistic and acoustic information have distinct masking effects during speech-in‐speech comprehension
  • Fitz, H., & Chang, F. (2017). Meaningful questions: The acquisition of auxiliary inversion in a connectionist model of sentence production. Cognition, 166, 225-250. doi:10.1016/j.cognition.2017.05.008.

    Abstract

    Nativist theories have argued that language involves syntactic principles which are unlearnable from the input children receive. A paradigm case of these innate principles is the structure dependence of auxiliary inversion in complex polar questions (Chomsky, 1968, 1975, 1980). Computational approaches have focused on the properties of the input in explaining how children acquire these questions. In contrast, we argue that messages are structured in a way that supports structure dependence in syntax. We demonstrate this approach within a connectionist model of sentence production (Chang, 2009) which learned to generate a range of complex polar questions from a structured message without positive exemplars in the input. The model also generated different types of error in development that were similar in magnitude to those in children (e.g., auxiliary doubling, Ambridge, Rowland, & Pine, 2008; Crain & Nakayama, 1987). Through model comparisons we trace how meaning constraints and linguistic experience interact during the acquisition of auxiliary inversion. Our results suggest that auxiliary inversion rules in English can be acquired without innate syntactic principles, as long as it is assumed that speakers who ask complex questions express messages that are structured into multiple propositions
  • Frank, S. L., & Willems, R. M. (2017). Word predictability and semantic similarity show distinct patterns of brain activity during language comprehension. Language, Cognition and Neuroscience, 32(9), 1192-1203. doi:10.1080/23273798.2017.1323109.

    Abstract

    We investigate the effects of two types of relationship between the words of a sentence or text – predictability and semantic similarity – by reanalysing electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) data from studies in which participants comprehend naturalistic stimuli. Each content word's predictability given previous words is quantified by a probabilistic language model, and semantic similarity to previous words is quantified by a distributional semantics model. Brain activity time-locked to each word is regressed on the two model-derived measures. Results show that predictability and semantic similarity have near identical N400 effects but are dissociated in the fMRI data, with word predictability related to activity in, among others, the visual word-form area, and semantic similarity related to activity in areas associated with the semantic network. This indicates that both predictability and similarity play a role during natural language comprehension and modulate distinct cortical regions.
  • Franken, M. K., Eisner, F., Schoffelen, J.-M., Acheson, D. J., Hagoort, P., & McQueen, J. M. (2017). Audiovisual recalibration of vowel categories. In Proceedings of Interspeech 2017 (pp. 655-658). doi:10.21437/Interspeech.2017-122.

    Abstract

    One of the most daunting tasks of a listener is to map a continuous auditory stream onto known speech sound categories and lexical items. A major issue with this mapping problem is the variability in the acoustic realizations of sound categories, both within and across speakers. Past research has suggested listeners may use visual information (e.g., lipreading) to calibrate these speech categories to the current speaker. Previous studies have focused on audiovisual recalibration of consonant categories. The present study explores whether vowel categorization, which is known to show less sharply defined category boundaries, also benefit from visual cues. Participants were exposed to videos of a speaker pronouncing one out of two vowels, paired with audio that was ambiguous between the two vowels. After exposure, it was found that participants had recalibrated their vowel categories. In addition, individual variability in audiovisual recalibration is discussed. It is suggested that listeners’ category sharpness may be related to the weight they assign to visual information in audiovisual speech perception. Specifically, listeners with less sharp categories assign more weight to visual information during audiovisual speech recognition.
  • Franken, M. K., Acheson, D. J., McQueen, J. M., Eisner, F., & Hagoort, P. (2017). Individual variability as a window on production-perception interactions in speech motor control. The Journal of the Acoustical Society of America, 142(4), 2007-2018. doi:10.1121/1.5006899.

    Abstract

    An important part of understanding speech motor control consists of capturing the interaction between speech production and speech perception. This study tests a prediction of theoretical frameworks that have tried to account for these interactions: if speech production targets are specified in auditory terms, individuals with better auditory acuity should have more precise speech targets, evidenced by decreased within-phoneme variability and increased between-phoneme distance. A study was carried out consisting of perception and production tasks in counterbalanced order. Auditory acuity was assessed using an adaptive speech discrimination task, while production variability was determined using a pseudo-word reading task. Analyses of the production data were carried out to quantify average within-phoneme variability as well as average between-phoneme contrasts. Results show that individuals not only vary in their production and perceptual abilities, but that better discriminators have more distinctive vowel production targets (that is, targets with less within-phoneme variability and greater between-phoneme distances), confirming the initial hypothesis. This association between speech production and perception did not depend on local phoneme density in vowel space. This study suggests that better auditory acuity leads to more precise speech production targets, which may be a consequence of auditory feedback affecting speech production over time.
  • Grabot, L., Kösem, A., Azizi, L., & Van Wassenhove, V. (2017). Prestimulus Alpha Oscillations and the Temporal Sequencing of Audio-visual Events. Journal of Cognitive Neuroscience, 29(9), 1566-1582. doi:10.1162/jocn_a_01145.

    Abstract

    Perceiving the temporal order of sensory events typically depends on participants' attentional state, thus likely on the endogenous fluctuations of brain activity. Using magnetoencephalography, we sought to determine whether spontaneous brain oscillations could disambiguate the perceived order of auditory and visual events presented in close temporal proximity, that is, at the individual's perceptual order threshold (Point of Subjective Simultaneity [PSS]). Two neural responses were found to index an individual's temporal order perception when contrasting brain activity as a function of perceived order (i.e., perceiving the sound first vs. perceiving the visual event first) given the same physical audiovisual sequence. First, average differences in prestimulus auditory alpha power indicated perceiving the correct ordering of audiovisual events irrespective of which sensory modality came first: a relatively low alpha power indicated perceiving auditory or visual first as a function of the actual sequence order. Additionally, the relative changes in the amplitude of the auditory (but not visual) evoked responses were correlated with participant's correct performance. Crucially, the sign of the magnitude difference in prestimulus alpha power and evoked responses between perceived audiovisual orders correlated with an individual's PSS. Taken together, our results suggest that spontaneous oscillatory activity cannot disambiguate subjective temporal order without prior knowledge of the individual's bias toward perceiving one or the other sensory modality first. Altogether, our results suggest that, under high perceptual uncertainty, the magnitude of prestimulus alpha (de)synchronization indicates the amount of compensation needed to overcome an individual's prior in the serial ordering and temporal sequencing of information
  • Hagoort, P. (2017). Don't forget neurobiology: An experimental approach to linguistic representation. Commentary on Branigan and Pickering "An experimental approach to linguistic representation". Behavioral and Brain Sciences, 40: e292. doi:10.1017/S0140525X17000401.

    Abstract

    Acceptability judgments are no longer acceptable as the holy grail for testing the nature of linguistic representations. Experimental and quantitative methods should be used to test theoretical claims in psycholinguistics. These methods should include not only behavior, but also the more recent possibilities to probe the neural codes for language-relevant representation
  • Hagoort, P. (2017). It is the facts, stupid. In J. Brockman, F. Van der Wa, & H. Corver (Eds.), Wetenschappelijke parels: het belangrijkste wetenschappelijke nieuws volgens 193 'briljante geesten'. Amsterdam: Maven Press.
  • Hagoort, P. (2017). The core and beyond in the language-ready brain. Neuroscience and Biobehavioral Reviews, 81, 194-204. doi:10.1016/j.neubiorev.2017.01.048.

    Abstract

    In this paper a general cognitive architecture of spoken language processing is specified. This is followed by an account of how this cognitive architecture is instantiated in the human brain. Both the spatial aspects of the networks for language are discussed, as well as the temporal dynamics and the underlying neurophysiology. A distinction is proposed between networks for coding/decoding linguistic information and additional networks for getting from coded meaning to speaker meaning, i.e. for making the inferences that enable the listener to understand the intentions of the speaker
  • Hagoort, P. (2017). The neural basis for primary and acquired language skills. In E. Segers, & P. Van den Broek (Eds.), Developmental Perspectives in Written Language and Literacy: In honor of Ludo Verhoeven (pp. 17-28). Amsterdam: Benjamins. doi:10.1075/z.206.02hag.

    Abstract

    Reading is a cultural invention that needs to recruit cortical infrastructure that was not designed for it (cultural recycling of cortical maps). In the case of reading both visual cortex and networks for speech processing are recruited. Here I discuss current views on the neurobiological underpinnings of spoken language 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. 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. The consequences of this architecture for reading are discussed.
  • Hartung, F. (2017). Getting under your skin: The role of perspective and simulation of experience in narrative comprehension. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Hartung, F., Hagoort, P., & Willems, R. M. (2017). Readers select a comprehension mode independent of pronoun: Evidence from fMRI during narrative comprehension. Brain and Language, 170, 29-38. doi:10.1016/j.bandl.2017.03.007.

    Abstract

    Perspective is a crucial feature for communicating about events. Yet it is unclear how linguistically encoded perspective relates to cognitive perspective taking. Here, we tested the effect of perspective taking with short literary stories. Participants listened to stories with 1st or 3rd person pronouns referring to the protagonist, while undergoing fMRI. When comparing action events with 1st and 3rd person pronouns, we found no evidence for a neural dissociation depending on the pronoun. A split sample approach based on the self-reported experience of perspective taking revealed 3 comprehension preferences. One group showed a strong 1st person preference, another a strong 3rd person preference, while a third group engaged in 1st and 3rd person perspective taking simultaneously. Comparing brain activations of the groups revealed different neural networks. Our results suggest that comprehension is perspective dependent, but not on the perspective suggested by the text, but on the reader’s (situational) preference
  • Hartung, F., Withers, P., Hagoort, P., & Willems, R. M. (2017). When fiction is just as real as fact: No differences in reading behavior between stories believed to be based on true or fictional events. Frontiers in Psychology, 8: 1618. doi:10.3389/fpsyg.2017.01618.

    Abstract

    Experiments have shown that compared to fictional texts, readers read factual texts faster and have better memory for described situations. Reading fictional texts on the other hand seems to improve memory for exact wordings and expressions. Most of these studies used a ‘newspaper’ versus ‘literature’ comparison. In the present study, we investigated the effect of reader’s expectation to whether information is true or fictional with a subtler manipulation by labelling short stories as either based on true or fictional events. In addition, we tested whether narrative perspective or individual preference in perspective taking affects reading true or fictional stories differently. In an online experiment, participants (final N=1742) read one story which was introduced as based on true events or as fictional (factor fictionality). The story could be narrated in either 1st or 3rd person perspective (factor perspective). We measured immersion in and appreciation of the story, perspective taking, as well as memory for events. We found no evidence that knowing a story is fictional or based on true events influences reading behavior or experiential aspects of reading. We suggest that it is not whether a story is true or fictional, but rather expectations towards certain reading situations (e.g. reading newspaper or literature) which affect behavior by activating appropriate reading goals. Results further confirm that narrative perspective partially influences perspective taking and experiential aspects of reading
  • Heyselaar, E., Hagoort, P., & Segaert, K. (2017). How social opinion influences syntactic processing – An investigation using virtual reality. PLoS One, 12(4): e0174405. doi:10.1371/journal.pone.0174405.
  • Heyselaar, E. (2017). Influences on the magnitude of syntactic priming. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Heyselaar, E., Hagoort, P., & Segaert, K. (2017). In dialogue with an avatar, language behavior is identical to dialogue with a human partner. Behavior Research Methods, 49(1), 46-60. doi:10.3758/s13428-015-0688-7.

    Abstract

    The use of virtual reality (VR) as a methodological tool is becoming increasingly popular in behavioral research as its flexibility allows for a wide range of applications. This new method has not been as widely accepted in the field of psycholinguistics, however, possibly due to the assumption that language processing during human-computer interactions does not accurately reflect human-human interactions. Yet at the same time there is a growing need to study human-human language interactions in a tightly controlled context, which has not been possible using existing methods. VR, however, offers experimental control over parameters that cannot be (as finely) controlled in the real world. As such, in this study we aim to show that human-computer language interaction is comparable to human-human language interaction in virtual reality. In the current study we compare participants’ language behavior in a syntactic priming task with human versus computer partners: we used a human partner, a human-like avatar with human-like facial expressions and verbal behavior, and a computer-like avatar which had this humanness removed. As predicted, our study shows comparable priming effects between the human and human-like avatar suggesting that participants attributed human-like agency to the human-like avatar. Indeed, when interacting with the computer-like avatar, the priming effect was significantly decreased. This suggests that when interacting with a human-like avatar, sentence processing is comparable to interacting with a human partner. Our study therefore shows that VR is a valid platform for conducting language research and studying dialogue interactions in an ecologically valid manner.
  • Heyselaar, E., Segaert, K., Walvoort, S. J., Kessels, R. P., & Hagoort, P. (2017). The role of nondeclarative memory in the skill for language: Evidence from syntactic priming in patients with amnesia. Neuropsychologia, 101, 97-105. doi:10.1016/j.neuropsychologia.2017.04.033.

    Abstract

    Syntactic priming, the phenomenon in which participants adopt the linguistic behaviour of their partner, is widely used in psycholinguistics to investigate syntactic operations. Although the phenomenon of syntactic priming is well documented, the memory system that supports the retention of this syntactic information long enough to influence future utterances, is not as widely investigated. We aim to shed light on this issue by assessing patients with Korsakoff's amnesia on an active-passive syntactic priming task and compare their performance to controls matched in age, education, and premorbid intelligence. Patients with Korsakoff's syndrome display deficits in all subdomains of declarative memory, yet their nondeclarative memory remains intact, making them an ideal patient group to determine which memory system supports syntactic priming. In line with the hypothesis that syntactic priming relies on nondeclarative memory, the patient group shows strong priming tendencies (12.6% passive structure repetition). Our healthy control group did not show a priming tendency, presumably due to cognitive interference between declarative and nondeclarative memory. We discuss the results in relation to amnesia, aging, and compensatory mechanisms.
  • Hirschmann, J., Schoffelen, J.-M., Schnitzler, A., & Van Gerven, M. A. J. (2017). Parkinsonian rest tremor can be detected accurately based on neuronal oscillations recorded from the subthalamic nucleus. Clinical Neurophysiology, 128, 2029-2036. doi:10.1016/j.clinph.2017.07.419.

    Abstract

    Objective: To investigate the possibility of tremor detection based on deep brain activity. Methods: We re-analyzed recordings of local field potentials (LFPs) from the subthalamic nucleus in 10 PD patients (12 body sides) with spontaneously fluctuating rest tremor. Power in several frequency bands was estimated and used as input to Hidden Markov Models (HMMs) which classified short data segments as either tremor-free rest or rest tremor. HMMs were compared to direct threshold application to individual power features. Results: Applying a threshold directly to band-limited power was insufficient for tremor detection (mean area under the curve [AUC] of receiver operating characteristic: 0.64, STD: 0.19). Multi-feature HMMs, in contrast, allowed for accurate detection (mean AUC: 0.82, STD: 0.15), using four power features obtained from a single contact pair. Within-patient training yielded better accuracy than across-patient training (0.84 vs. 0.78, p = 0.03), yet tremor could often be detected accurately with either approach. High frequency oscillations (>200 Hz) were the best performing individual feature. Conclusions: LFP-based markers of tremor are robust enough to allow for accurate tremor detection in short data segments, provided that appropriate statistical models are used. Significance: LFP-based markers of tremor could be useful control signals for closed-loop deep brain stimulation.
  • Ito, A., Martin, A. E., & Nieuwland, M. S. (2017). Why the A/AN prediction effect may be hard to replicate: A rebuttal to DeLong, Urbach & Kutas (2017). Language, Cognition and Neuroscience, 32(8), 974-983. doi:10.1080/23273798.2017.1323112.
  • Kita, S., Alibali, M. W., & Chu, M. (2017). How Do Gestures Influence Thinking and Speaking? The Gesture-for-Conceptualization Hypothesis. Psychological Review, 124(3), 245-266. doi:10.1037/rev0000059.

    Abstract

    People spontaneously produce gestures during speaking and thinking. The authors focus here on gestures that depict or indicate information related to the contents of concurrent speech or thought (i.e., representational gestures). Previous research indicates that such gestures have not only communicative functions, but also self-oriented cognitive functions. In this article, the authors propose a new theoretical framework, the gesture-for-conceptualization hypothesis, which explains the self-oriented functions of representational gestures. According to this framework, representational gestures affect cognitive processes in 4 main ways: gestures activate, manipulate, package, and explore spatio-motoric information for speaking and thinking. These four functions are shaped by gesture's ability to schematize information, that is, to focus on a small subset of available information that is potentially relevant to the task at hand. The framework is based on the assumption that gestures are generated from the same system that generates practical actions, such as object manipulation; however, gestures are distinct from practical actions in that they represent information. The framework provides a novel, parsimonious, and comprehensive account of the self-oriented functions of gestures. The authors discuss how the framework accounts for gestures that depict abstract or metaphoric content, and they consider implications for the relations between self-oriented and communicative functions of gestures
  • Kösem, A., & Van Wassenhove, V. (2017). Distinct contributions of low and high frequency neural oscillations to speech comprehension. Language, Cognition and Neuroscience, 32(5), 536-544. doi:10.1080/23273798.2016.1238495.

    Abstract

    In the last decade, the involvement of neural oscillatory mechanisms in speech comprehension has been increasingly investigated. Current evidence suggests that low-frequency and high-frequency neural entrainment to the acoustic dynamics of speech are linked to its analysis. One crucial question is whether acoustical processing primarily modulates neural entrainment, or whether entrainment instead reflects linguistic processing. Here, we review studies investigating the effect of linguistic manipulations on neural oscillatory activity. In light of the current findings, we argue that theta (3–8 Hz) entrainment may primarily reflect the analysis of the acoustic features of speech. In contrast, recent evidence suggests that delta (1–3 Hz) and high-frequency activity (>40 Hz) are reliable indicators of perceived linguistic representations. The interdependence between low-frequency and high-frequency neural oscillations, as well as their causal role on speech comprehension, is further discussed with regard to neurophysiological models of speech processing
  • Kunert, R., & Jongman, S. R. (2017). Entrainment to an auditory signal: Is attention involved? Journal of Experimental Psychology: General, 146(1), 77-88. doi:10.1037/xge0000246.

    Abstract

    Many natural auditory signals, including music and language, change periodically. The effect of such auditory rhythms on the brain is unclear however. One widely held view, dynamic attending theory, proposes that the attentional system entrains to the rhythm and increases attention at moments of rhythmic salience. In support, 2 experiments reported here show reduced response times to visual letter strings shown at auditory rhythm peaks, compared with rhythm troughs. However, we argue that an account invoking the entrainment of general attention should further predict rhythm entrainment to also influence memory for visual stimuli. In 2 pseudoword memory experiments we find evidence against this prediction. Whether a pseudoword is shown during an auditory rhythm peak or not is irrelevant for its later recognition memory in silence. Other attention manipulations, dividing attention and focusing attention, did result in a memory effect. This raises doubts about the suggested attentional nature of rhythm entrainment. We interpret our findings as support for auditory rhythm perception being based on auditory-motor entrainment, not general attention entrainment.
  • Kunert, R. (2017). Music and language comprehension in the brain. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Lam, N. H. L. (2017). Comprehending comprehension: Insights from neuronal oscillations on the neuronal basis of language. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Lam, K. J. Y., Bastiaansen, M. C. M., Dijkstra, T., & Rueschemeyer, S. A. (2017). Making sense: motor activation and action plausibility during sentence processing. Language, Cognition and Neuroscience, 32(5), 590-600. doi:10.1080/23273798.2016.1164323.

    Abstract

    The current electroencephalography study investigated the relationship between the motor and (language) comprehension systems by simultaneously measuring mu and N400 effects. Specifically, we examined whether the pattern of motor activation elicited by verbs depends on the larger sentential context. A robust N400 congruence effect confirmed the contextual manipulation of action plausibility, a form of semantic congruency. Importantly, this study showed that: (1) Action verbs elicited more mu power decrease than non-action verbs when sentences described plausible actions. Action verbs thus elicited more motor activation than non-action verbs. (2) In contrast, when sentences described implausible actions, mu activity was present but the difference between the verb types was not observed. The increased processing associated with a larger N400 thus coincided with mu activity in sentences describing implausible actions. Altogether, context-dependent motor activation appears to play a functional role in deriving context-sensitive meaning
  • Lewis, A. G., Schoffelen, J.-M., Hoffmann, C., Bastiaansen, M. C. M., & Schriefers, H. (2017). Discourse-level semantic coherence influences beta oscillatory dynamics and the N400 during sentence comprehension. Language, Cognition and Neuroscience, 32(5), 601-617. doi:10.1080/23273798.2016.1211300.

    Abstract

    In this study, we used electroencephalography to investigate the influence of discourse-level semantic coherence on electrophysiological signatures of local sentence-level processing. Participants read groups of four sentences that could either form coherent stories or were semantically unrelated. For semantically coherent discourses compared to incoherent ones, the N400 was smaller at sentences 2–4, while the visual N1 was larger at the third and fourth sentences. Oscillatory activity in the beta frequency range (13–21 Hz) was higher for coherent discourses. We relate the N400 effect to a disruption of local sentence-level semantic processing when sentences are unrelated. Our beta findings can be tentatively related to disruption of local sentence-level syntactic processing, but it cannot be fully ruled out that they are instead (or also) related to disrupted local sentence-level semantic processing. We conclude that manipulating discourse-level semantic coherence does have an effect on oscillatory power related to local sentence-level processing.
  • Lewis, A. G. (2017). Explorations of beta-band neural oscillations during language comprehension: Sentence processing and beyond. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Lockwood, G. (2017). Talking sense: The behavioural and neural correlates of sound symbolism. PhD Thesis, Radboud University, Nijmegen.
  • Lopopolo, A., Frank, S. L., Van den Bosch, A., & Willems, R. M. (2017). Using stochastic language models (SLM) to map lexical, syntactic, and phonological information processing in the brain. PLoS One, 12(5): e0177794. doi:10.1371/journal.pone.0177794.

    Abstract

    Language comprehension involves the simultaneous processing of information at the phonological, syntactic, and lexical level. We track these three distinct streams of information in the brain by using stochastic measures derived from computational language models to detect neural correlates of phoneme, part-of-speech, and word processing in an fMRI experiment. Probabilistic language models have proven to be useful tools for studying how language is processed as a sequence of symbols unfolding in time. Conditional probabilities between sequences of words are at the basis of probabilistic measures such as surprisal and perplexity which have been successfully used as predictors of several behavioural and neural correlates of sentence processing. Here we computed perplexity from sequences of words and their parts of speech, and their phonemic transcriptions. Brain activity time-locked to each word is regressed on the three model-derived measures. We observe that the brain keeps track of the statistical structure of lexical, syntactic and phonological information in distinct areas.

    Additional information

    Data availability
  • Martin, A. E., Huettig, F., & Nieuwland, M. S. (2017). Can structural priming answer the important questions about language? A commentary on Branigan and Pickering "An experimental approach to linguistic representation". Behavioral and Brain Sciences, 40: e304. doi:10.1017/S0140525X17000528.

    Abstract

    While structural priming makes a valuable contribution to psycholinguistics, it does not allow direct observation of representation, nor escape “source ambiguity.” Structural priming taps into implicit memory representations and processes that may differ from what is used online. We question whether implicit memory for language can and should be equated with linguistic representation or with language processing.
  • Nieuwland, M. S., & Martin, A. E. (2017). Neural oscillations and a nascent corticohippocampal theory of reference. Journal of Cognitive Neuroscience, 29(5), 896-910. doi:10.1162/jocn_a_01091.

    Abstract

    The ability to use words to refer to the world is vital to the communicative power of human language. In particular, the anaphoric use of words to refer to previously mentioned concepts (antecedents) allows dialogue to be coherent and meaningful. Psycholinguistic theory posits that anaphor comprehension involves reactivating a memory representation of the antecedent. Whereas this implies the involvement of recognition memory, or the mnemonic sub-routines by which people distinguish old from new, the neural processes for reference resolution are largely unknown. Here, we report time-frequency analysis of four EEG experiments to reveal the increased coupling of functional neural systems associated with referentially coherent expressions compared to referentially problematic expressions. Despite varying in modality, language, and type of referential expression, all experiments showed larger gamma-band power for referentially coherent expressions compared to referentially problematic expressions. Beamformer analysis in high-density Experiment 4 localised the gamma-band increase to posterior parietal cortex around 400-600 ms after anaphor-onset and to frontaltemporal cortex around 500-1000 ms. We argue that the observed gamma-band power increases reflect successful referential binding and resolution, which links incoming information to antecedents through an interaction between the brain’s recognition memory networks and frontal-temporal language network. We integrate these findings with previous results from patient and neuroimaging studies, and we outline a nascent cortico-hippocampal theory of reference.
  • Peeters, D., Snijders, T. M., Hagoort, P., & Ozyurek, A. (2017). Linking language to the visual world: Neural correlates of comprehending verbal reference to objects through pointing and visual cues. Neuropsychologia, 95, 21-29. doi:10.1016/j.neuropsychologia.2016.12.004.

    Abstract

    In everyday communication speakers often refer in speech and/or gesture to objects in their immediate environment, thereby shifting their addressee's attention to an intended referent. The neurobiological infrastructure involved in the comprehension of such basic multimodal communicative acts remains unclear. In an event-related fMRI study, we presented participants with pictures of a speaker and two objects while they concurrently listened to her speech. In each picture, one of the objects was singled out, either through the speaker's index-finger pointing gesture or through a visual cue that made the object perceptually more salient in the absence of gesture. A mismatch (compared to a match) between speech and the object singled out by the speaker's pointing gesture led to enhanced activation in left IFG and bilateral pMTG, showing the importance of these areas in conceptual matching between speech and referent. Moreover, a match (compared to a mismatch) between speech and the object made salient through a visual cue led to enhanced activation in the mentalizing system, arguably reflecting an attempt to converge on a jointly attended referent in the absence of pointing. These findings shed new light on the neurobiological underpinnings of the core communicative process of comprehending a speaker's multimodal referential act and stress the power of pointing as an important natural device to link speech to objects.
  • Rommers, J., Dickson, D. S., Norton, J. J. S., Wlotko, E. W., & Federmeier, K. D. (2017). Alpha and theta band dynamics related to sentential constraint and word expectancy. Language, Cognition and Neuroscience, 32(5), 576-589. doi:10.1080/23273798.2016.1183799.

    Abstract

    Despite strong evidence for prediction during language comprehension, the underlying mechanisms, and the extent to which they are specific to language, remain unclear. Re-analysing an event-related potentials study, we examined responses in the time-frequency domain to expected and unexpected (but plausible) words in strongly and weakly constraining sentences, and found results similar to those reported in nonverbal domains. Relative to expected words, unexpected words elicited an increase in the theta band (4–7 Hz) in strongly constraining contexts, suggesting the involvement of control processes to deal with the consequences of having a prediction disconfirmed. Prior to critical word onset, strongly constraining sentences exhibited a decrease in the alpha band (8–12 Hz) relative to weakly constraining sentences, suggesting that comprehenders can take advantage of predictive sentence contexts to prepare for the input. The results suggest that the brain recruits domain-general preparation and control mechanisms when making and assessing predictions during sentence comprehension
  • Rommers, J., Meyer, A. S., & Praamstra, P. (2017). Lateralized electrical brain activity reveals covert attention allocation during speaking. Neuropsychologia, 95, 101-110. doi:10.1016/j.neuropsychologia.2016.12.013.

    Abstract

    Speakers usually begin to speak while only part of the utterance has been planned. Earlier work has shown that speech planning processes are reflected in speakers’ eye movements as they describe visually presented objects. However, to-be-named objects can be processed to some extent before they have been fixated upon, presumably because attention can be allocated to objects covertly, without moving the eyes. The present study investigated whether EEG could track speakers’ covert attention allocation as they produced short utterances to describe pairs of objects (e.g., “dog and chair”). The processing difficulty of each object was varied by presenting it in upright orientation (easy) or in upside down orientation (difficult). Background squares flickered at different frequencies in order to elicit steady-state visual evoked potentials (SSVEPs). The N2pc component, associated with the focusing of attention on an item, was detectable not only prior to speech onset, but also during speaking. The time course of the N2pc showed that attention shifted to each object in the order of mention prior to speech onset. Furthermore, greater processing difficulty increased the time speakers spent attending to each object. This demonstrates that the N2pc can track covert attention allocation in a naming task. In addition, an effect of processing difficulty at around 200–350 ms after stimulus onset revealed early attention allocation to the second to-be-named object. The flickering backgrounds elicited SSVEPs, but SSVEP amplitude was not influenced by processing difficulty. These results help complete the picture of the coordination of visual information uptake and motor output during speaking.
  • Schoffelen, J.-M., Hulten, A., Lam, N. H. L., Marquand, A. F., Udden, J., & Hagoort, P. (2017). Frequency-specific directed interactions in the human brain network for language. Proceedings of the National Academy of Sciences of the United States of America, 114(30), 8083-8088. doi:10.1073/pnas.1703155114.

    Abstract

    The brain’s remarkable capacity for language requires bidirectional interactions between functionally specialized brain regions. We used magnetoencephalography to investigate interregional interactions in the brain network for language while 102 participants were reading sentences. Using Granger causality analysis, we identified inferior frontal cortex and anterior temporal regions to receive widespread input and middle temporal regions to send widespread output. This fits well with the notion that these regions play a central role in language processing. Characterization of the functional topology of this network, using data-driven matrix factorization, which allowed for partitioning into a set of subnetworks, revealed directed connections at distinct frequencies of interaction. Connections originating from temporal regions peaked at alpha frequency, whereas connections originating from frontal and parietal regions peaked at beta frequency. These findings indicate that the information flow between language-relevant brain areas, which is required for linguistic processing, may depend on the contributions of distinct brain rhythms

    Additional information

    pnas.201703155SI.pdf
  • Schoot, L. (2017). Language processing in a conversation context. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Shitova, N., Roelofs, A., Schriefers, H., Bastiaansen, M., & Schoffelen, J.-M. (2017). Control adjustments in speaking: Electrophysiology of the Gratton effect in picture naming. Cortex, 92, 289-303. doi:10.1016/j.cortex.2017.04.017.

    Abstract

    Accumulating evidence suggests that spoken word production requires different amounts of top-down control depending on the prevailing circumstances. For example, during Stroop-like tasks, the interference in response time (RT) is typically larger following congruent trials than following incongruent trials. This effect is called the Gratton effect, and has been taken to reflect top-down control adjustments based on the previous trial type. Such control adjustments have been studied extensively in Stroop and Eriksen flanker tasks (mostly using manual responses), but not in the picture-word interference (PWI) task, which is a workhorse of language production research. In one of the few studies of the Gratton effect in PWI, Van Maanen and Van Rijn (2010) examined the effect in picture naming RTs during dual-task performance. Based on PWI effect differences between dual-task conditions, they argued that the functional locus of the PWI effect differs between post-congruent trials (i.e., locus in perceptual and conceptual encoding) and post-incongruent trials (i.e., locus in word planning). However, the dual-task procedure may have contaminated the results. We therefore performed an EEG study on the Gratton effect in a regular PWI task. We observed a PWI effect in the RTs, in the N400 component of the event-related brain potentials, and in the midfrontal theta power, regardless of the previous trial type. Moreover, the RTs, N400, and theta power reflected the Gratton effect. These results provide evidence that the PWI effect arises at the word planning stage following both congruent and incongruent trials, while the amount of top-down control changes depending on the previous trial type.
  • Silva, S., Inácio, F., Folia, V., & Petersson, K. M. (2017). Eye movements in implicit artificial grammar learning. Journal of Experimental Psychology: Learning, Memory, and Cognition, 43(9), 1387-1402. doi:10.1037/xlm0000350.

    Abstract

    Artificial grammar learning (AGL) has been probed with forced-choice behavioral tests (active tests). Recent attempts to probe the outcomes of learning (implicitly acquired knowledge) with eye-movement responses (passive tests) have shown null results. However, these latter studies have not tested for sensitivity effects, for example, increased eye movements on a printed violation. In this study, we tested for sensitivity effects in AGL tests with (Experiment 1) and without (Experiment 2) concurrent active tests (preference- and grammaticality classification) in an eye-tracking experiment. Eye movements discriminated between sequence types in passive tests and more so in active tests. The eye-movement profile did not differ between preference and grammaticality classification, and it resembled sensitivity effects commonly observed in natural syntax processing. Our findings show that the outcomes of implicit structured sequence learning can be characterized in eye tracking. More specifically, whole trial measures (dwell time, number of fixations) showed robust AGL effects, whereas first-pass measures (first-fixation duration) did not. Furthermore, our findings strengthen the link between artificial and natural syntax processing, and they shed light on the factors that determine performance differences in preference and grammaticality classification tests
  • Silva, S., Petersson, K. M., & Castro, S. L. (2017). The effects of ordinal load on incidental temporal learning. Quarterly Journal of Experimental Psychology, 70(4), 664-674. doi:10.1080/17470218.2016.1146909.

    Abstract

    How can we grasp the temporal structure of events? A few studies have indicated that representations of temporal structure are acquired when there is an intention to learn, but not when learning is incidental. Response-to-stimulus intervals, uncorrelated temporal structures, unpredictable ordinal information, and lack of metrical organization have been pointed out as key obstacles to incidental temporal learning, but the literature includes piecemeal demonstrations of learning under all these circumstances. We suggest that the unacknowledged effects of ordinal load may help reconcile these conflicting findings, ordinal load referring to the cost of identifying the sequence of events (e.g., tones, locations) where a temporal pattern is embedded. In a first experiment, we manipulated ordinal load into simple and complex levels. Participants learned ordinal-simple sequences, despite their uncorrelated temporal structure and lack of metrical organization. They did not learn ordinal-complex sequences, even though there were no response-to-stimulus intervals nor unpredictable ordinal information. In a second experiment, we probed learning of ordinal-complex sequences with strong metrical organization, and again there was no learning. We conclude that ordinal load is a key obstacle to incidental temporal learning. Further analyses showed that the effect of ordinal load is to mask the expression of temporal knowledge, rather than to prevent learning.
  • Silva, S., Folia, V., Hagoort, P., & Petersson, K. M. (2017). The P600 in Implicit Artificial Grammar Learning. Cognitive Science, 41(1), 137-157. doi:10.1111/cogs.12343.

    Abstract

    The suitability of the Artificial Grammar Learning (AGL) paradigm to capture relevant aspects of the acquisition of linguistic structures has been empirically tested in a number of EEG studies. Some have shown a syntax-related P600 component, but it has not been ruled out that the AGL P600 effect is a response to surface features (e.g., subsequence familiarity) rather than the underlying syntax structure. Therefore, in this study, we controlled for the surface characteristics of the test sequences (associative chunk strength) and recorded the EEG before (baseline preference classification) and after (preference and grammaticality classification) exposure to a grammar. A typical, centroparietal P600 effect was elicited by grammatical violations after exposure, suggesting that the AGL P600 effect signals a response to structural irregularities. Moreover, preference and grammaticality classification showed a qualitatively similar ERP profile, strengthening the idea that the implicit structural mere exposure paradigm in combination with preference classification is a suitable alternative to the traditional grammaticality classification test.
  • Simon, E., & Sjerps, M. J. (2017). Phonological category quality in the mental lexicon of child and adult learners. International Journal of Bilingualism, 21(4), 474-499. doi:10.1177/1367006915626589.

    Abstract

    Aims and objectives: The aim was to identify which criteria children use to decide on the category membership of native and non-native vowels, and to get insight into the organization of phonological representations in the bilingual mind. Methodology: The study consisted of two cross-language mispronunciation detection tasks in which L2 vowels were inserted into L1 words and vice versa. In Experiment 1, 10- to 12-year-old Dutch-speaking children were presented with Dutch words which were either pronounced with the target Dutch vowel or with an English vowel inserted in the Dutch consonantal frame. Experiment 2 was a mirror of the first, with English words which were pronounced “correctly” or which were “mispronounced” with a Dutch vowel. Data and analysis: Analyses focused on extent to which child and adult listeners accepted substitutions of Dutch vowels by English ones, and vice versa. Findings: The results of Experiment 1 revealed that between the age of ten and twelve children have well-established phonological vowel categories in their native language. However, Experiment 2 showed that in their non-native language, children tended to accept mispronounced items which involve sounds from their native language. At the same time, though, they did not fully rely on their native phonemic inventory because the children accepted most of the correctly pronounced English items. Originality: While many studies have examined native and non-native perception by infants and adults, studies on first and second language perception of school-age children are rare. This study adds to the body of literature aimed at expanding our knowledge in this area. Implications: The study has implications for models of the organization of the bilingual mind: while proficient adult non-native listeners generally have clearly separated sets of phonological representations for their two languages, for non-proficient child learners the L1 phonology still exerts a strong influence on the L2 phonology.
  • Soutschek, A., Burke, C. J., Beharelle, A. R., Schreiber, R., Weber, S. C., Karipidis, I. I., Ten Velden, J., Weber, B., Haker, H., Kalenscher, T., & Tobler, P. N. (2017). The dopaminergic reward system underpins gender differences in social preferences. Nature Human Behaviour, 1, 819-827. doi:10.1038/s41562-017-0226-y.

    Abstract

    Women are known to have stronger prosocial preferences than men, but it remains an open question as to how these behavioural differences arise from differences in brain functioning. Here, we provide a neurobiological account for the hypothesized gender difference. In a pharmacological study and an independent neuroimaging study, we tested the hypothesis that the neural reward system encodes the value of sharing money with others more strongly in women than in men. In the pharmacological study, we reduced receptor type-specific actions of dopamine, a neurotransmitter related to reward processing, which resulted in more selfish decisions in women and more prosocial decisions in men. Converging findings from an independent neuroimaging study revealed gender-related activity in neural reward circuits during prosocial decisions. Thus, the neural reward system appears to be more sensitive to prosocial rewards in women than in men, providing a neurobiological account for why women often behave more prosocially than men. A large body of evidence suggests that women are often more prosocial (for example, generous, altruistic and inequality averse) than men, at least when other factors such as reputation and strategic considerations are excluded1,2,3. This dissociation could result from cultural expectations and gender stereotypes, because in Western societies women are more strongly expected to be prosocial4,5,6 and sensitive to variations in social context than men1. It remains an open question, however, whether and how on a neurobiological level the social preferences of women and men arise from differences in brain functioning. The assumption of gender differences in social preferences predicts that the neural reward system’s sensitivity to prosocial and selfish rewards should differ between women and men. Specifically, the hypothesis would be that the neural reward system is more sensitive to prosocial than selfish rewards in women and more sensitive to selfish than prosocial rewards in men. The goal of the current study was to test in two independent experiments for the hypothesized gender differences on both a pharmacological and a haemodynamic level. In particular, we examined the functions of the neurotransmitter dopamine using a dopamine receptor antagonist, and the role of the striatum (a brain region strongly innervated by dopamine neurons) during social decision-making in women and men using neuroimaging. The neurotransmitter dopamine is thought to play a key role in neural reward processing7,8. Recent evidence suggests that dopaminergic activity is sensitive not only to rewards for oneself but to rewards for others as well9. The assumption that dopamine is sensitive to both self- and other-related outcomes is consistent with the finding that the striatum shows activation for both selfish and shared rewards10,11,12,13,14,15. The dopaminergic response may represent a net signal encoding the difference between the value of preferred and unpreferred rewards8. Regarding the hypothesized gender differences in social preferences, this account makes the following predictions. If women prefer shared (prosocial) outcomes2, women’s dopaminergic signals to shared rewards will be stronger than to non-shared (selfish) rewards, so reducing dopaminergic activity should bias women to make more selfish decisions. In line with this hypothesis, a functional imaging study reported enhanced striatal activation in female participants during charitable donations11. In contrast, if men prefer selfish over prosocial rewards, dopaminergic activity should be enhanced to selfish compared to prosocial rewards. In line with this view, upregulating dopaminergic activity in a sample of exclusively male participants increased selfish behaviour in a bargaining game16. Thus, contrary to the hypothesized effect in women, reducing dopaminergic neurotransmission should render men more prosocial. Taken together, the current study tested the following three predictions: we expected the dopaminergic reward system (1) to be more sensitive to prosocial than selfish rewards in women and (2) to be more sensitive to selfish than prosocial rewards in men. As a consequence of these two predictions, we also predicted (3) dopaminoceptive regions such as the striatum to show stronger activation to prosocial relative to selfish rewards in women than in men. To test these predictions, we conducted a pharmacological study in which we reduced dopaminergic neurotransmission with amisulpride. Amisulpride is a dopamine antagonist that is highly specific for dopaminergic D2/D3 receptors17. After receiving amisulpride or placebo, participants performed an interpersonal decision task18,19,20, in which they made choices between a monetary reward only for themselves (selfish reward option) and sharing money with others (prosocial reward option). We expected that blocking dopaminergic neurotransmission with amisulpride, relative to placebo, would result in fewer prosocial choices in women and more prosocial choices in men. To investigate whether potential gender-related effects of dopamine are selective for social decision-making, we also tested the effects of amisulpride on time preferences in a non-social control task that was matched to the interpersonal decision task in terms of choice structure. In addition, because dopaminergic neurotransmission plays a crucial role in brain regions involved in value processing, such as the striatum21, a gender-related role of dopaminergic activity for social decision-making should also be reflected by dissociable activity patterns in the striatum. Therefore, to further test our hypothesis, we investigated the neural correlates of social decision-making in a functional imaging study. In line with our predictions for the pharmacological study, we expected to find stronger striatum activity during prosocial relative to selfish decisions in women, whereas men should show enhanced activity in the striatum for selfish relative to prosocial choices.

    Additional information

    Supplementary Information
  • Ye, Z., Stolk, A., Toni, I., & Hagoort, P. (2017). Oxytocin modulates semantic integration in speech comprehension. Journal of Cognitive Neuroscience, 29, 267-276. doi:10.1162/jocn_a_01044.

    Abstract

    Listeners interpret utterances by integrating information from multiple sources including word level semantics and world knowledge. When the semantics of an expression is inconsistent with his or her knowledge about the world, the listener may have to search through the conceptual space for alternative possible world scenarios that can make the expression more acceptable. Such cognitive exploration requires considerable computational resources and might depend on motivational factors. This study explores whether and how oxytocin, a neuropeptide known to influence socialmotivation by reducing social anxiety and enhancing affiliative tendencies, can modulate the integration of world knowledge and sentence meanings. The study used a betweenparticipant double-blind randomized placebo-controlled design. Semantic integration, indexed with magnetoencephalography through the N400m marker, was quantified while 45 healthymale participants listened to sentences that were either congruent or incongruent with facts of the world, after receiving intranasally delivered oxytocin or placebo. Compared with congruent sentences, world knowledge incongruent sentences elicited a stronger N400m signal from the left inferior frontal and anterior temporal regions and medial pFC (the N400m effect) in the placebo group. Oxytocin administration significantly attenuated the N400meffect at both sensor and cortical source levels throughout the experiment, in a state-like manner. Additional electrophysiological markers suggest that the absence of the N400m effect in the oxytocin group is unlikely due to the lack of early sensory or semantic processing or a general downregulation of attention. These findings suggest that oxytocin drives listeners to resolve challenges of semantic integration, possibly by promoting the cognitive exploration of alternative possible world scenarios.
  • Takashima, A., Bakker, I., Van Hell, J. G., Janzen, G., & McQueen, J. M. (2017). Interaction between episodic and semantic memory networks in the acquisition and consolidation of novel spoken words. Brain and Language, 167, 44-60. doi:10.1016/j.bandl.2016.05.009.

    Abstract

    When a novel word is learned, its memory representation is thought to undergo a process of consolidation and integration. In this study, we tested whether the neural representations of novel words change as a function of consolidation by observing brain activation patterns just after learning and again after a delay of one week. Words learned with meanings were remembered better than those learned without meanings. Both episodic (hippocampus-dependent) and semantic (dependent on distributed neocortical areas) memory systems were utilised during recognition of the novel words. The extent to which the two systems were involved changed as a function of time and the amount of associated information, with more involvement of both systems for the meaningful words than for the form-only words after the one-week delay. These results suggest that the reason the meaningful words were remembered better is that their retrieval can benefit more from these two complementary memory systems
  • Tan, Y., Martin, R. C., & Van Dyke, J. A. (2017). Semantic and syntactic interference in sentence comprehension: A comparison of working memory models. Frontiers in Psychology, 8: 198. doi:10.3389/fpsyg.2017.00198.

    Abstract

    This study investigated the nature of the underlying working memory system supporting sentence processing through examining individual differences in sensitivity to retrieval interference effects during sentence comprehension. Interference effects occur when readers incorrectly retrieve sentence constituents which are similar to those required during integrative processes. We examined interference arising from a partial match between distracting constituents and syntactic and semantic cues, and related these interference effects to performance on working memory, short-term memory (STM), vocabulary, and executive function tasks. For online sentence comprehension, as measured by self-paced reading, the magnitude of individuals' syntactic interference effects was predicted by general WM capacity and the relation remained significant when partialling out vocabulary, indicating that the effects were not due to verbal knowledge. For offline sentence comprehension, as measured by responses to comprehension questions, both general WM capacity and vocabulary knowledge interacted with semantic interference for comprehension accuracy, suggesting that both general WM capacity and the quality of semantic representations played a role in determining how well interference was resolved offline. For comprehension question reaction times, a measure of semantic STM capacity interacted with semantic but not syntactic interference. However, a measure of phonological capacity (digit span) and a general measure of resistance to response interference (Stroop effect) did not predict individuals' interference resolution abilities in either online or offline sentence comprehension. The results are discussed in relation to the multiple capacities account of working memory (e.g., Martin and Romani, 1994; Martin and He, 2004), and the cue-based retrieval parsing approach (e.g., Lewis et al., 2006; Van Dyke et al., 2014). While neither approach was fully supported, a possible means of reconciling the two approaches and directions for future research are proposed.
  • Tsuji, S., Fikkert, P., Minagawa, Y., Dupoux, E., Filippin, L., Versteegh, M., Hagoort, P., & Cristia, A. (2017). The more, the better? Behavioral and neural correlates of frequent and infrequent vowel exposure. Developmental Psychobiology, 59, 603-612. doi:10.1002/dev.21534.

    Abstract

    A central assumption in the perceptual attunement literature holds that exposure to a speech sound contrast leads to improvement in native speech sound processing. However, whether the amount of exposure matters for this process has not been put to a direct test. We elucidated indicators of frequency-dependent perceptual attunement by comparing 5–8-month-old Dutch infants’ discrimination of tokens containing a highly frequent [hɪt-he:t] and a highly infrequent [hʏt-hø:t] native vowel contrast as well as a non-native [hɛt-hæt] vowel contrast in a behavioral visual habituation paradigm (Experiment 1). Infants discriminated both native contrasts similarly well, but did not discriminate the non-native contrast. We sought further evidence for subtle differences in the processing of the two native contrasts using near-infrared spectroscopy and a within-participant design (Experiment 2). The neuroimaging data did not provide additional evidence that responses to native contrasts are modulated by frequency of exposure. These results suggest that even large differences in exposure to a native contrast may not directly translate to behavioral and neural indicators of perceptual attunement, raising the possibility that frequency of exposure does not influence improvements in discriminating native contrasts.

    Additional information

    dev21534-sup-0001-SuppInfo-S1.docx
  • Udden, J., Snijders, T. M., Fisher, S. E., & Hagoort, P. (2017). A common variant of the CNTNAP2 gene is associated with structural variation in the left superior occipital gyrus. Brain and Language, 172, 16-21. doi:10.1016/j.bandl.2016.02.003.

    Abstract

    The CNTNAP2 gene encodes a cell-adhesion molecule that influences the properties of neural networks and the morphology and density of neurons and glial cells. Previous studies have shown association of CNTNAP2 variants with language-related phenotypes in health and disease. Here, we report associations of a common CNTNAP2 polymorphism (rs7794745) with variation in grey matter in a region in the dorsal visual stream. We tried to replicate an earlier study on 314 subjects by Tan and colleagues (2010), but now in a substantially larger group of more than 1700 subjects. Carriers of the T allele showed reduced grey matter volume in left superior occipital gyrus, while we did not replicate associations with grey matter volume in other regions identified by Tan et al (2010). Our work illustrates the importance of independent replication in neuroimaging genetic studies of language-related candidate genes.
  • Udden, J., Ingvar, M., Hagoort, P., & Petersson, K. M. (2017). Broca’s region: A causal role in implicit processing of grammars with crossed non-adjacent dependencies. Cognition, 164, 188-198. doi:10.1016/j.cognition.2017.03.010.

    Abstract

    Non-adjacent dependencies are challenging for the language learning machinery and are acquired later than adjacent dependencies. In this transcranial magnetic stimulation (TMS) study, we show that participants successfully discriminated between grammatical and non-grammatical sequences after having implicitly acquired an artificial language with crossed non-adjacent dependencies. Subsequent to transcranial magnetic stimulation of Broca’s region, discrimination was impaired compared to when a language-irrelevant control region (vertex) was stimulated. These results support the view that Broca’s region is engaged in structured sequence processing and extend previous functional neuroimaging results on artificial grammar learning (AGL) in two directions: first, the results establish that Broca’s region is a causal component in the processing of non-adjacent dependencies, and second, they show that implicit processing of non-adjacent dependencies engages Broca’s region. Since patients with lesions in Broca’s region do not always show grammatical processing difficulties, the result that Broca’s region is causally linked to processing of non-adjacent dependencies is a step towards clarification of the exact nature of syntactic deficits caused by lesions or perturbation to Broca’s region. Our findings are consistent with previous results and support a role for Broca’s region in general structured sequence processing, rather than a specific role for the processing of hierarchically organized sentence structure.
  • Van der Ven, F., Takashima, A., Segers, A., & Verhoeven, L. (2017). Semantic priming in Dutch children: Word meaning integration and study modality effects. Language Learning, 67(3), 546-568. doi:10.1111/lang.12235.
  • Van Bergen, G., & Flecken, M. (2017). Putting things in new places: Linguistic experience modulates the predictive power of placement verb semantics. Journal of Memory and Language, 92, 26-42. doi:10.1016/j.jml.2016.05.003.

    Abstract

    A central question regarding predictive language processing concerns the extent to which linguistic experience modulates the process. We approached this question by investigating sentence processing in advanced second language (L2) users with different native language (L1) backgrounds. Using a visual world eye tracking paradigm, we investigated to what extent L1 and L2 participants showed anticipatory eye movements to objects while listening to Dutch placement event descriptions. L2 groups differed in the degree of similarity between Dutch and their L1 with respect to placement verb semantics: German, like Dutch, specifies object position in placement verbs (put.STAND vs. put.LIE), whereas English and French typically leave position underspecified (put). Results showed that German L2 listeners, like native Dutch listeners, anticipate objects that match the verbally encoded position immediately upon encountering the verb. French/English L2 participants, however, did not show any prediction effects, despite proper understanding of Dutch placement verbs. Our findings suggest that prior experience with a specific semantic contrast in one’s L1 facilitates prediction in L2, and hence adds to the evidence that linguistic experience modulates predictive sentence processing
  • Van Ekert, J., Wegman, J., Jansen, C., Takashima, A., & Janzen, G. (2017). The dynamics of memory consolidation of landmarks. Hippocampus, 27(4), 303-404. doi:10.1002/hipo.22698.

    Abstract

    Navigating through space is fundamental to human nature and requires the ability to retrieve relevant information from the remote past. With the passage of time, some memories become generic, capturing only a sense of familiarity. Yet, others maintain precision, even when acquired decades ago. Understanding the dynamics of memory consolidation is a major challenge to neuroscientists. Using functional magnetic resonance imaging, we systematically examined the effects of time and spatial context on the neural representation of landmark recognition memory. An equal number of male and female subjects (males N = 10, total N = 20) watched a route through a large-scale virtual environment. Landmarks occurred at navigationally relevant and irrelevant locations along the route. Recognition memory for landmarks was tested directly following encoding, 24 h later and 30 days later. Surprisingly, changes over time in the neural representation of navigationally relevant landmarks differed between males and females. In males, relevant landmarks selectively engaged the parahippocampal gyrus (PHG) regardless of the age of the memory. In females, the response to relevant landmarks gradually diminished with time in the PHG but strengthened progressively in the inferior frontal gyrus (IFG). Based on what is known about the functioning of the PHG and IFG, the findings of this study suggest that males maintain access to the initially formed spatial representation of landmarks whereas females become strongly dependent on a verbal representation of landmarks with time. Our findings yield a clear objective for future studies
  • Vanlangendonck, F. (2017). Finding common ground: On the neural mechanisms of communicative language production. PhD Thesis, Radboud University, Nijmegen.
  • Varma, S., Takashima, A., Krewinkel, S., Van Kooten, M., Fu, L., Medendorp, W. P., Kessels, R. P. C., & Daselaar, S. M. (2017). Non-interfering effects of active post-encoding tasks on episodic memory consolidation in humans. Frontiers in Behavioral Neuroscience, 11: 54. doi:10.3389/fnbeh.2017.00054.

    Abstract

    So far, studies that investigated interference effects of post-learning processes on episodic memory consolidation in humans have used tasks involving only complex and meaningful information. Such tasks require reallocation of general or encoding-specific resources away from consolidation-relevant activities. The possibility that interference can be elicited using a task that heavily taxes our limited brain resources, but has low semantic and hippocampal related long-term memory processing demands, has never been tested. We address this question by investigating whether consolidation could persist in parallel with an active, encoding-irrelevant, minimally semantic task, regardless of its high resource demands for cognitive processing. We distinguish the impact of such a task on consolidation based on whether it engages resources that are: (1) general/executive, or (2) specific/overlapping with the encoding modality. Our experiments compared subsequent memory performance across two post-encoding consolidation periods: quiet wakeful rest and a cognitively demanding n-Back task. Across six different experiments (total N = 176), we carefully manipulated the design of the n-Back task to target general or specific resources engaged in the ongoing consolidation process. In contrast to previous studies that employed interference tasks involving conceptual stimuli and complex processing demands, we did not find any differences between n-Back and rest conditions on memory performance at delayed test, using both recall and recognition tests. Our results indicate that: (1) quiet, wakeful rest is not a necessary prerequisite for episodic memory consolidation; and (2) post-encoding cognitive engagement does not interfere with memory consolidation when task-performance has minimal semantic and hippocampally-based episodic memory processing demands. We discuss our findings with reference to resource and reactivation-led interference theories
  • Allen, S., Ozyurek, A., Kita, S., Brown, A., Furman, R., Ishizuka, T., & Fujii, M. (2007). Language-specific and universal influences in children's syntactic packaging of manner and path: A comparison of English, Japanese, and Turkish. Cognition, 102, 16-48. doi:10.1016/j.cognition.2005.12.006.

    Abstract

    Different languages map semantic elements of spatial relations onto different lexical and syntactic units. These crosslinguistic differences raise important questions for language development in terms of how this variation is learned by children. We investigated how Turkish-, English-, and Japanese-speaking children (mean age 3;8) package the semantic elements of Manner and Path onto syntactic units when both the Manner and the Path of the moving Figure occur simultaneously and are salient in the event depicted. Both universal and language-specific patterns were evident in our data. Children used the semantic-syntactic mappings preferred by adult speakers of their own languages, and even expressed subtle syntactic differences that encode different relations between Manner and Path in the same way as their adult counterparts (i.e., Manner causing vs. incidental to Path). However, not all types of semantics-syntax mappings were easy for children to learn (e.g., expressing Manner and Path elements in two verbal clauses). In such cases, Turkish- and Japanese-speaking children frequently used syntactic patterns that were not typical in the target language but were similar to patterns used by English-speaking children, suggesting some universal influence. Thus, both language-specific and universal tendencies guide the development of complex spatial expressions.
  • Bramão, I., Mendonça, A., Faísca, L., Ingvar, M., Petersson, K. M., & Reis, A. (2007). The impact of reading and writing skills on a visuo-motor integration task: A comparison between illiterate and literate subjects. Journal of the International Neuropsychological Society, 13(2), 359-364. doi:10.1017/S1355617707070440.

    Abstract

    Previous studies have shown a significant association between reading skills and the performance on visuo-motor tasks. In order to clarify whether reading and writing skills modulate non-linguistic domains, we investigated the performance of two literacy groups on a visuo-motor integration task with non-linguistic stimuli. Twenty-one illiterate participants and twenty matched literate controls were included in the experiment. Subjects were instructed to use the right or the left index finger to point to and touch a randomly presented target on the right or left side of a touch screen. The results showed that the literate subjects were significantly faster in detecting and touching targets on the left compared to the right side of the screen. In contrast, the presentation side did not affect the performance of the illiterate group. These results lend support to the idea that having acquired reading and writing skills, and thus a preferred left-to-right reading direction, influences visual scanning. (JINS, 2007, 13, 359–364
  • Furman, R., & Ozyurek, A. (2007). Development of interactional discourse markers: Insights from Turkish children's and adults' narratives. Journal of Pragmatics, 39(10), 1742-1757. doi:10.1016/j.pragma.2007.01.008.

    Abstract

    Discourse markers (DMs) are linguistic elements that index different relations and coherence between units of talk (Schiffrin, Deborah, 1987. Discourse Markers. Cambridge University Press, Cambridge). Most research on the development of these forms has focused on conversations rather than narratives and furthermore has not directly compared children's use of DMs to adult usage. This study examines the development of three DMs (şey ‘uuhh’, yani ‘I mean’, işte ‘y’know’) that mark interactional levels of discourse in oral Turkish narratives in 60 Turkish children (3-, 5- and 9-year-olds) and 20 Turkish-speaking adults. The results show that the frequency and functions of DMs change with age. Children learn şey, which mainly marks exchange level structures, earliest. However, yani and işte have multi-functions such as marking both information states and participation frameworks and are consequently learned later. Children also use DMs with different functions than adults. Overall, the results show that learning to use interactional DMs in narratives is complex and goes beyond age 9, especially for multi-functional DMs that index an interplay of discourse coherence at different levels.
  • Gisselgard, J., Uddén, J., Ingvar, M., & Petersson, K. M. (2007). Disruption of order information by irrelevant items: A serial recognition paradigm. Acta Psychologica, 124(3), 356-369. doi:10.1016/j.actpsy.2006.04.002.

    Abstract

    Irrelevant speech effect (ISE) is defined as a decrement in visually presented digit-list short-term memory performance due to exposure to irrelevant auditory material. Perhaps the most successful theoretical explanation of the effect is the changing state hypothesis. This hypothesis explains the effect in terms of confusion between amodal serial order cues, and represents a view based on the interference caused by the processing of similar order information of the visual and auditory materials. An alternative view suggests that the interference occurs as a consequence of the similarity between the visual and auditory contents of the stimuli. An important argument for the former view is the observation that ISE is almost exclusively observed in tasks that require memory for serial order. However, most short-term memory tasks require that both item and order information be retained in memory. An ideal task to investigate the sensitivity of maintenance of serial order to irrelevant speech would be one that calls upon order information but not item information. One task that is particularly suited to address this issue is serial recognition. In a typical serial recognition task, a list of items is presented and then probed by the same list in which the order of two adjacent items has been transposed. Due to the re-presentation of the encoding string, serial recognition requires primarily the serial order to be maintained while the content of the presented items is deemphasized. In demonstrating a highly significant ISE of changing versus steady-state auditory items in a serial recognition task, the present finding lends support for and extends previous empirical findings suggesting that irrelevant speech has the potential to interfere with the coding of the order of the items to be memorized.
  • Gürcanli, Ö., Nakipoglu Demiralp, M., & Ozyurek, A. (2007). Shared information and argument omission in Turkish. In H. Caunt-Nulton, S. Kulatilake, & I. Woo (Eds.), Proceedings of the 31st Annual Boston University Conference on Language Developement (pp. 267-273). Somerville, Mass: Cascadilla Press.
  • Hagoort, P., & Van Berkum, J. J. A. (2007). Beyond the sentence given. Philosophical Transactions of the Royal Society. Series B: Biological Sciences, 362, 801-811.

    Abstract

    A central and influential idea among researchers of language is that our language faculty is organized according to Fregean compositionality, which states that the meaning of an utterance is a function of the meaning of its parts and of the syntactic rules by which these parts are combined. Since the domain of syntactic rules is the sentence, the implication of this idea is that language interpretation takes place in a two-step fashion. First, the meaning of a sentence is computed. In a second step, the sentence meaning is integrated with information from prior discourse, world knowledge, information about the speaker and semantic information from extra-linguistic domains such as co-speech gestures or the visual world. Here, we present results from recordings of event-related brain potentials that are inconsistent with this classical two-step model of language interpretation. Our data support a one-step model in which knowledge about the context and the world, concomitant information from other modalities, and the speaker are brought to bear immediately, by the same fast-acting brain system that combines the meanings of individual words into a message-level representation. Underlying the one-step model is the immediacy assumption, according to which all available information will immediately be used to co-determine the interpretation of the speaker's message. Functional magnetic resonance imaging data that we collected indicate that Broca's area plays an important role in semantic unification. Language comprehension involves the rapid incorporation of information in a 'single unification space', coming from a broader range of cognitive domains than presupposed in the standard two-step model of interpretation.
  • Hagoort, P. (2007). The memory, unification, and control (MUC) model of language. In T. Sakamoto (Ed.), Communicating skills of intention (pp. 259-291). Tokyo: Hituzi Syobo.
  • Hagoort, P. (2007). The memory, unification, and control (MUC) model of language. In A. S. Meyer, L. Wheeldon, & A. Krott (Eds.), Automaticity and control in language processing (pp. 243-270). Hove: Psychology Press.
  • Hald, L. A., Steenbeek-Planting, E. G., & Hagoort, P. (2007). The interaction of discourse context and world knowledge in online sentence comprehension: Evidence from the N400. Brain Research, 1146, 210-218. doi:10.1016/j.brainres.2007.02.054.

    Abstract

    In an ERP experiment we investigated how the recruitment and integration of world knowledge information relate to the integration of information within a current discourse context. Participants were presented with short discourse contexts which were followed by a sentence that contained a critical word that was correct or incorrect based on general world knowledge and the supporting discourse context, or was more or less acceptable based on the combination of general world knowledge and the specific local discourse context. Relative to the critical word in the correct world knowledge sentences following a neutral discourse, all other critical words elicited an N400 effect that began at about 300 ms after word onset. However, the magnitude of the N400 effect varied in a way that suggests an interaction between world knowledge and discourse context. The results indicate that both world knowledge and discourse context have an effect on sentence interpretation, but neither overrides the other.
  • Janzen, G., Wagensveld, B., & Van Turennout, M. (2007). Neural representation of navigational relevance is rapidly induced and long lasting. Cerebral Cortex, 17(4), 975-981. doi:10.1093/cercor/bhl008.

    Abstract

    Successful navigation is facilitated by the presence of landmarks. Previous functional magnetic resonance imaging (fMRI) evidence indicated that the human parahippocampal gyrus automatically distinguishes between landmarks placed at navigationally relevant (decision points) and irrelevant locations (nondecision points). This storage of navigational relevance can provide a neural mechanism underlying successful navigation. However, an efficient wayfinding mechanism requires that important spatial information is learned quickly and maintained over time. The present study investigates whether the representation of navigational relevance is modulated by time and practice. Participants learned 2 film sequences through virtual mazes containing objects at decision and at nondecision points. One maze was shown one time, and the other maze was shown 3 times. Twenty-four hours after study, event-related fMRI data were acquired during recognition of the objects. The results showed that activity in the parahippocampal gyrus was increased for objects previously placed at decision points as compared with objects placed at nondecision points. The decision point effect was not modulated by the number of exposures to the mazes and independent of explicit memory functions. These findings suggest a persistent representation of navigationally relevant information, which is stable after only one exposure to an environment. These rapidly induced and long-lasting changes in object representation provide a basis for successful wayfinding.
  • Janzen, G., & Weststeijn, C. G. (2007). Neural representation of object location and route direction: An event-related fMRI study. Brain Research, 1165, 116-125. doi:10.1016/j.brainres.2007.05.074.

    Abstract

    The human brain distinguishes between landmarks placed at navigationally relevant and irrelevant locations. However, to provide a successful wayfinding mechanism not only landmarks but also the routes between them need to be stored. We examined the neural representation of a memory for route direction and a memory for relevant landmarks. Healthy human adults viewed objects along a route through a virtual maze. Event-related functional magnetic resonance imaging (fMRI) data were acquired during a subsequent subliminal priming recognition task. Prime-objects either preceded or succeeded a target-object on a preciously learned route. Our results provide evidence that the parahippocampal gyri distinguish between relevant and irrelevant landmarks whereas the inferior parietal gyrus, the anterior cingulate gyrus as well as the right caudate nucleus are involved in the coding of route direction. These data show that separated memory systems store different spatial information. A memory for navigationally relevant object information and a memory for route direction exist.
  • Kelly, S. D., & Ozyurek, A. (Eds.). (2007). Gesture, language, and brain [Special Issue]. Brain and Language, 101(3).
  • Kita, S., & Ozyurek, A. (2007). How does spoken language shape iconic gestures? In S. Duncan, J. Cassel, & E. Levy (Eds.), Gesture and the dynamic dimension of language (pp. 67-74). Amsterdam: Benjamins.
  • Kita, S., Ozyurek, A., Allen, S., Brown, A., Furman, R., & Ishizuka, T. (2007). Relations between syntactic encoding and co-speech gestures: Implications for a model of speech and gesture production. Language and Cognitive Processes, 22(8), 1212-1236. doi:10.1080/01690960701461426.

    Abstract

    Gestures that accompany speech are known to be tightly coupled with speech production. However little is known about the cognitive processes that underlie this link. Previous cross-linguistic research has provided preliminary evidence for online interaction between the two systems based on the systematic co-variation found between how different languages syntactically package Manner and Path information of a motion event and how gestures represent Manner and Path. Here we elaborate on this finding by testing whether speakers within the same language gesturally express Manner and Path differently according to their online choice of syntactic packaging of Manner and Path, or whether gestural expression is pre-determined by a habitual conceptual schema congruent with the linguistic typology. Typologically congruent and incongruent syntactic structures for expressing Manner and Path (i.e., in a single clause or multiple clauses) were elicited from English speakers. We found that gestural expressions were determined by the online choice of syntactic packaging rather than by a habitual conceptual schema. It is therefore concluded that speech and gesture production processes interface online at the conceptual planning phase. Implications of the findings for models of speech and gesture production are discussed
  • Marklund, P., Fransson, P., Cabeza, R., Petersson, K. M., Ingvar, M., & Nyberg, L. (2007). Sustained and transient neural modulations in prefrontal cortex related to declarative long-term memory, working memory, and attention. Cortex, 43(1), 22-37. doi:10.1016/S0010-9452(08)70443-X.

    Abstract

    Common activations in prefrontal cortex (PFC) during episodic and semantic long-term memory (LTM) tasks have been hypothesized to reflect functional overlap in terms of working memory (WM) and cognitive control. To evaluate a WM account of LTM-general activations, the present study took into consideration that cognitive task performance depends on the dynamic operation of multiple component processes, some of which are stimulus-synchronous and transient in nature; and some that are engaged throughout a task in a sustained fashion. PFC and WM may be implicated in both of these temporally independent components. To elucidate these possibilities we employed mixed blocked/event-related functional magnetic resonance imaging (fMRI) procedures to assess the extent to which sustained or transient activation patterns overlapped across tasks indexing episodic and semantic LTM, attention (ATT), and WM. Within PFC, ventrolateral and medial areas exhibited sustained activity across all tasks, whereas more anterior regions including right frontopolar cortex were commonly engaged in sustained processing during the three memory tasks. These findings do not support a WM account of sustained frontal responses during LTM tasks, but instead suggest that the pattern that was common to all tasks reflects general attentional set/vigilance, and that the shared WM-LTM pattern mediates control processes related to upholding task set. Transient responses during the three memory tasks were assessed relative to ATT to isolate item-specific mnemonic processes and were found to be largely distinct from sustained effects. Task-specific effects were observed for each memory task. In addition, a common item response for all memory tasks involved left dorsolateral PFC (DLPFC). The latter response might be seen as reflecting WM processes during LTM retrieval. Thus, our findings suggest that a WM account of shared PFC recruitment in LTM tasks holds for common transient item-related responses rather than sustained state-related responses that are better seen as reflecting more general attentional/control processes.
  • Menenti, L., & Burani, C. (2007). What causes the effect of age of acquisition in lexical processing? Quarterly Journal of Experimental Psychology, 60(5), 652-660. doi:10.1080/17470210601100126.

    Abstract

    Three hypotheses for effects of age of acquisition (AoA) in lexical processing are compared: the cumulative frequency hypothesis (frequency and AoA both influence the number of encounters with a word, which influences processing speed), the semantic hypothesis (early-acquired words are processed faster because they are more central in the semantic network), and the neural network model (early-acquired words are faster because they are acquired when a network has maximum plasticity). In a regression study of lexical decision (LD) and semantic categorization (SC) in Italian and Dutch, contrary to the cumulative frequency hypothesis, AoA coefficients were larger than frequency coefficients, and, contrary to the semantic hypothesis, the effect of AoA was not larger in SC than in LD. The neural network model was supported.
  • Nieuwland, M. S., Petersson, K. M., & Van Berkum, J. J. A. (2007). On sense and reference: Examining the functional neuroanatomy of referential processing. NeuroImage, 37(3), 993-1004. doi:10.1016/j.neuroimage.2007.05.048.

    Abstract

    In an event-related fMRI study, we examined the cortical networks involved in establishing reference during language comprehension. We compared BOLD responses to sentences containing referentially ambiguous pronouns (e.g., “Ronald told Frank that he…”), referentially failing pronouns (e.g., “Rose told Emily that he…”) or coherent pronouns. Referential ambiguity selectively recruited medial prefrontal regions, suggesting that readers engaged in problem-solving to select a unique referent from the discourse model. Referential failure elicited activation increases in brain regions associated with morpho-syntactic processing, and, for those readers who took failing pronouns to refer to unmentioned entities, additional regions associated with elaborative inferencing were observed. The networks activated by these two referential problems did not overlap with the network activated by a standard semantic anomaly. Instead, we observed a double dissociation, in that the systems activated by semantic anomaly are deactivated by referential ambiguity, and vice versa. This inverse coupling may reflect the dynamic recruitment of semantic and episodic processing to resolve semantically or referentially problematic situations. More generally, our findings suggest that neurocognitive accounts of language comprehension need to address not just how we parse a sentence and combine individual word meanings, but also how we determine who's who and what's what during language comprehension.
  • Nieuwland, M. S., Otten, M., & Van Berkum, J. J. A. (2007). Who are you talking about? Tracking discourse-level referential processing with event-related brain potentials. Journal of Cognitive Neuroscience, 19(2), 228-236. doi:10.1162/jocn.2007.19.2.228.

    Abstract

    In this event-related brain potentials (ERPs) study, we explored the possibility to selectively track referential ambiguity during spoken discourse comprehension. Earlier ERP research has shown that referentially ambiguous nouns (e.g., “the girl” in a two-girl context) elicit a frontal, sustained negative shift relative to unambiguous control words. In the current study, we examined whether this ERP effect reflects “deep” situation model ambiguity or “superficial” textbase ambiguity. We contrasted these different interpretations by investigating whether a discourse-level semantic manipulation that prevents referential ambiguity also averts the elicitation of a referentially induced ERP effect. We compared ERPs elicited by nouns that were referentially nonambiguous but were associated with two discourse entities (e.g., “the girl” with two girls introduced in the context, but one of which has died or left the scene), with referentially ambiguous and nonambiguous control words. Although temporally referentially ambiguous nouns elicited a frontal negative shift compared to control words, the “double bound” but referentially nonambiguous nouns did not. These results suggest that it is possible to selectively track referential ambiguity with ERPs at the level that is most relevant to discourse comprehension, the situation model.
  • Otten, M., Nieuwland, M. S., & Van Berkum, J. J. A. (2007). Great expectations: Specific lexical anticipation influences the processing of spoken language. BMC Neuroscience, 8: 89. doi:10.1186/1471-2202-8-89.

    Abstract

    Background Recently several studies have shown that people use contextual information to make predictions about the rest of the sentence or story as the text unfolds. Using event related potentials (ERPs) we tested whether these on-line predictions are based on a message-based representation of the discourse or on simple automatic activation by individual words. Subjects heard short stories that were highly constraining for one specific noun, or stories that were not specifically predictive but contained the same prime words as the predictive stories. To test whether listeners make specific predictions critical nouns were preceded by an adjective that was inflected according to, or in contrast with, the gender of the expected noun. Results When the message of the preceding discourse was predictive, adjectives with an unexpected gender-inflection evoked a negative deflection over right-frontal electrodes between 300 and 600 ms. This effect was not present in the prime control context, indicating that the prediction mismatch does not hinge on word-based priming but is based on the actual message of the discourse. Conclusions When listening to a constraining discourse people rapidly make very specific predictions about the remainder of the story, as the story unfolds. These predictions are not simply based on word-based automatic activation, but take into account the actual message of the discourse.
  • Otten, M., & Van Berkum, J. J. A. (2007). What makes a discourse constraining? Comparing the effects of discourse message and scenario fit on the discourse-dependent N400 effect. Brain Research, 1153, 166-177. doi:10.1016/j.brainres.2007.03.058.

    Abstract

    A discourse context provides a reader with a great deal of information that can provide constraints for further language processing, at several different levels. In this experiment we used event-related potentials (ERPs) to explore whether discourse-generated contextual constraints are based on the precise message of the discourse or, more `loosely', on the scenario suggested by one or more content words in the text. Participants read constraining stories whose precise message rendered a particular word highly predictable ("The manager thought that the board of directors should assemble to discuss the issue. He planned a...[meeting]") as well as non-constraining control stories that were only biasing in virtue of the scenario suggested by some of the words ("The manager thought that the board of directors need not assemble to discuss the issue. He planned a..."). Coherent words that were inconsistent with the message-level expectation raised in a constraining discourse (e.g., "session" instead of "meeting") elicited a classic centroparietal N400 effect. However, when the same words were only inconsistent with the scenario loosely suggested by earlier words in the text, they elicited a different negativity around 400 ms, with a more anterior, left-lateralized maximum. The fact that the discourse-dependent N400 effect cannot be reduced to scenario-mediated priming reveals that it reflects the rapid use of precise message-level constraints in comprehension. At the same time, the left-lateralized negativity in non-constraining stories suggests that, at least in the absence of strong message-level constraints, scenario-mediated priming does also rapidly affect comprehension.
  • Ozyurek, A., & Kelly, S. D. (2007). Gesture, language, and brain. Brain and Language, 101(3), 181-185. doi:10.1016/j.bandl.2007.03.006.
  • Ozyurek, A. (2007). Processing of multi-modal semantic information: Insights from cross-linguistic comparisons and neurophysiological recordings. In T. Sakamoto (Ed.), Communicating skills of intention (pp. 131-142). Tokyo: Hituzi Syobo Publishing.
  • Ozyurek, A., Willems, R. M., Kita, S., & Hagoort, P. (2007). On-line integration of semantic information from speech and gesture: Insights from event-related brain potentials. Journal of Cognitive Neuroscience, 19(4), 605-616. doi:10.1162/jocn.2007.19.4.605.

    Abstract

    During language comprehension, listeners use the global semantic representation from previous sentence or discourse context to immediately integrate the meaning of each upcoming word into the unfolding message-level representation. Here we investigate whether communicative gestures that often spontaneously co-occur with speech are processed in a similar fashion and integrated to previous sentence context in the same way as lexical meaning. Event-related potentials were measured while subjects listened to spoken sentences with a critical verb (e.g., knock), which was accompanied by an iconic co-speech gesture (i.e., KNOCK). Verbal and/or gestural semantic content matched or mismatched the content of the preceding part of the sentence. Despite the difference in the modality and in the specificity of meaning conveyed by spoken words and gestures, the latency, amplitude, and topographical distribution of both word and gesture mismatches are found to be similar, indicating that the brain integrates both types of information simultaneously. This provides evidence for the claim that neural processing in language comprehension involves the simultaneous incorporation of information coming from a broader domain of cognition than only verbal semantics. The neural evidence for similar integration of information from speech and gesture emphasizes the tight interconnection between speech and co-speech gestures.
  • Petersson, K. M., Silva, C., Castro-Caldas, A., Ingvar, M., & Reis, A. (2007). Literacy: A cultural influence on functional left-right differences in the inferior parietal cortex. European Journal of Neuroscience, 26(3), 791-799. doi:10.1111/j.1460-9568.2007.05701.x.

    Abstract

    The current understanding of hemispheric interaction is limited. Functional hemispheric specialization is likely to depend on both genetic and environmental factors. In the present study we investigated the importance of one factor, literacy, for the functional lateralization in the inferior parietal cortex in two independent samples of literate and illiterate subjects. The results show that the illiterate group are consistently more right-lateralized than their literate controls. In contrast, the two groups showed a similar degree of left-right differences in early speech-related regions of the superior temporal cortex. These results provide evidence suggesting that a cultural factor, literacy, influences the functional hemispheric balance in reading and verbal working memory-related regions. In a third sample, we investigated grey and white matter with voxel-based morphometry. The results showed differences between literacy groups in white matter intensities related to the mid-body region of the corpus callosum and the inferior parietal and parietotemporal regions (literate > illiterate). There were no corresponding differences in the grey matter. This suggests that the influence of literacy on brain structure related to reading and verbal working memory is affecting large-scale brain connectivity more than grey matter per se.
  • Qin, S., Piekema, C., Petersson, K. M., Han, B., Luo, J., & Fernández, G. (2007). Probing the transformation of discontinuous associations into episodic memory: An event-related fMRI study. NeuroImage, 38(1), 212-222. doi:10.1016/j.neuroimage.2007.07.020.

    Abstract

    Using event-related functional magnetic resonance imaging, we identified brain regions involved in storing associations of events discontinuous in time into long-term memory. Participants were scanned while memorizing item-triplets including simultaneous and discontinuous associations. Subsequent memory tests showed that participants remembered both types of associations equally well. First, by constructing the contrast between the subsequent memory effects for discontinuous associations and simultaneous associations, we identified the left posterior parahippocampal region, dorsolateral prefrontal cortex, the basal ganglia, posterior midline structures, and the middle temporal gyrus as being specifically involved in transforming discontinuous associations into episodic memory. Second, we replicated that the prefrontal cortex and the medial temporal lobe (MTL) especially the hippocampus are involved in associative memory formation in general. Our findings provide evidence for distinct neural operation(s) that supports the binding and storing discontinuous associations in memory. We suggest that top-down signals from the prefrontal cortex and MTL may trigger reactivation of internal representation in posterior midline structures of the first event, thus allowing it to be associated with the second event. The dorsolateral prefrontal cortex together with basal ganglia may support this encoding operation by executive and binding processes within working memory, and the posterior parahippocampal region may play a role in binding and memory formation.
  • Reis, A., Faísca, L., Mendonça, S., Ingvar, M., & Petersson, K. M. (2007). Semantic interference on a phonological task in illiterate subjects. Scandinavian Journal of Psychology, 48(1), 69-74. doi:10.1111/j.1467-9450.2006.00544.x.

    Abstract

    Previous research suggests that learning an alphabetic written language influences aspects of the auditory-verbal language system. In this study, we examined whether literacy influences the notion of words as phonological units independent of lexical semantics in literate and illiterate subjects. Subjects had to decide which item in a word- or pseudoword pair was phonologically longest. By manipulating the relationship between referent size and phonological length in three word conditions (congruent, neutral, and incongruent) we could examine to what extent subjects focused on form rather than meaning of the stimulus material. Moreover, the pseudoword condition allowed us to examine global phonological awareness independent of lexical semantics. The results showed that literate performed significantly better than illiterate subjects in the neutral and incongruent word conditions as well as in the pseudoword condition. The illiterate group performed least well in the incongruent condition and significantly better in the pseudoword condition compared to the neutral and incongruent word conditions and suggest that performance on phonological word length comparisons is dependent on literacy. In addition, the results show that the illiterate participants are able to perceive and process phonological length, albeit less well than the literate subjects, when no semantic interference is present. In conclusion, the present results confirm and extend the finding that illiterate subjects are biased towards semantic-conceptual-pragmatic types of cognitive processing.
  • De Ruiter, J. P., Noordzij, M. L., Newman-Norlund, S., Hagoort, P., & Toni, I. (2007). On the origins of intentions. In P. Haggard, Y. Rossetti, & M. Kawato (Eds.), Sensorimotor foundations of higher cognition (pp. 593-610). Oxford: Oxford University Press.
  • Snijders, T. M., Kooijman, V., Cutler, A., & Hagoort, P. (2007). Neurophysiological evidence of delayed segmentation in a foreign language. Brain Research, 1178, 106-113. doi:10.1016/j.brainres.2007.07.080.

    Abstract

    Previous studies have shown that segmentation skills are language-specific, making it difficult to segment continuous speech in an unfamiliar language into its component words. Here we present the first study capturing the delay in segmentation and recognition in the foreign listener using ERPs. We compared the ability of Dutch adults and of English adults without knowledge of Dutch (‘foreign listeners’) to segment familiarized words from continuous Dutch speech. We used the known effect of repetition on the event-related potential (ERP) as an index of recognition of words in continuous speech. Our results show that word repetitions in isolation are recognized with equivalent facility by native and foreign listeners, but word repetitions in continuous speech are not. First, words familiarized in isolation are recognized faster by native than by foreign listeners when they are repeated in continuous speech. Second, when words that have previously been heard only in a continuous-speech context re-occur in continuous speech, the repetition is detected by native listeners, but is not detected by foreign listeners. A preceding speech context facilitates word recognition for native listeners, but delays or even inhibits word recognition for foreign listeners. We propose that the apparent difference in segmentation rate between native and foreign listeners is grounded in the difference in language-specific skills available to the listeners.
  • Takashima, A., Nieuwenhuis, I. L. C., Rijpkema, M., Petersson, K. M., Jensen, O., & Fernández, G. (2007). Memory trace stabilization leads to large-scale changes in the retrieval network: A functional MRI study on associative memory. Learning & Memory, 14, 472-479. doi:10.1101/lm.605607.

    Abstract

    Spaced learning with time to consolidate leads to more stabile memory traces. However, little is known about the neural correlates of trace stabilization, especially in humans. The present fMRI study contrasted retrieval activity of two well-learned sets of face-location associations, one learned in a massed style and tested on the day of learning (i.e., labile condition) and another learned in a spaced scheme over the course of one week (i.e., stabilized condition). Both sets of associations were retrieved equally well, but the retrieval of stabilized association was faster and accompanied by large-scale changes in the network supporting retrieval. Cued recall of stabilized as compared with labile associations was accompanied by increased activity in the precuneus, the ventromedial prefrontal cortex, the bilateral temporal pole, and left temporo–parietal junction. Conversely, memory representational areas such as the fusiform gyrus for faces and the posterior parietal cortex for locations did not change their activity with stabilization. The changes in activation in the precuneus, which also showed increased connectivity with the fusiform area, are likely to be related to the spatial nature of our task. The activation increase in the ventromedial prefrontal cortex, on the other hand, might reflect a general function in stabilized memory retrieval. This area might succeed the hippocampus in linking distributed neocortical representations.
  • Tendolkar, I., Arnold, J., Petersson, K. M., Weis, S., Brockhaus-Dumke, A., Van Eijndhoven, P., Buitelaar, J., & Fernández, G. (2007). Probing the neural correlates of associative memory formation: A parametrically analyzed event-related functional MRI study. Brain Research, 1142, 159-168. doi:10.1016/j.brainres.2007.01.040.

    Abstract

    The medial temporal lobe (MTL) is crucial for declarative memory formation, but the function of its subcomponents in associative memory formation remains controversial. Most functional imaging studies on this topic are based on a stepwise approach comparing a condition with and one without associative encoding. Extending this approach we applied additionally a parametric analysis by varying the amount of associative memory formation. We found a hippocampal subsequent memory effect of almost similar magnitude regardless of the amount of associations formed. By contrast, subsequent memory effects in rhinal and parahippocampal cortices were parametrically and positively modulated by the amount of associations formed. Our results indicate that the parahippocampal region supports associative memory formation as tested here and the hippocampus adds a general mnemonic operation. This pattern of results might suggest a new interpretation. Instead of having either a fixed division of labor between the hippocampus (associative memory formation) and the rhinal cortex (non-associative memory formation) or a functionally unitary MTL system, in which all substructures are contributing to memory formation in a similar way, we propose that the location where associations are formed within the MTL depends on the kind of associations bound: If visual single-dimension associations, as used here, can already be integrated within the parahippocampal region, the hippocampus might add a general purpose mnemonic operation only. In contrast, if associations have to be formed across widely distributed neocortical representations, the hippocampus may provide a binding operation in order to establish a coherent memory.
  • Van Berkum, J. J. A., Koornneef, A. W., Otten, M., & Nieuwland, M. S. (2007). Establishing reference in language comprehension: An electrophysiological perspective. Brain Research, 1146, 158-171. doi:10.1016/j.brainres.2006.06.091.

    Abstract

    The electrophysiology of language comprehension has long been dominated by research on syntactic and semantic integration. However, to understand expressions like "he did it" or "the little girl", combining word meanings in accordance with semantic and syntactic constraints is not enough--readers and listeners also need to work out what or who is being referred to. We review our event-related brain potential research on the processes involved in establishing reference, and present a new experiment in which we examine when and how the implicit causality associated with specific interpersonal verbs affects the interpretation of a referentially ambiguous pronoun. The evidence suggests that upon encountering a singular noun or pronoun, readers and listeners immediately inspect their situation model for a suitable discourse entity, such that they can discriminate between having too many, too few, or exactly the right number of referents within at most half a second. Furthermore, our implicit causality findings indicate that a fragment like "David praised Linda because..." can immediately foreground a particular referent, to the extent that a subsequent "he" is at least initially construed as a syntactic error. In all, our brain potential findings suggest that referential processing is highly incremental, and not necessarily contingent upon the syntax. In addition, they demonstrate that we can use ERPs to relatively selectively keep track of how readers and listeners establish reference.
  • Van Alphen, P. M. (2007). Prevoicing in Dutch initial plosives: Production, perception, and word recognition. In J. van de Weijer, & E. van der Torre (Eds.), Voicing in Dutch (pp. 99-124). Amsterdam: Benjamins.

    Abstract

    Prevoicing is the presence of vocal fold vibration during the closure of initial voiced plosives (negative VOT). The presence or absence of prevoicing is generally used to describe the voicing distinction in Dutch initial plosives. However, a phonetic study showed that prevoicing is frequently absent in Dutch. This article discusses the role of prevoicing in the production and perception of Dutch plosives. Furthermore, two cross-modal priming experiments are presented that examined the effect of prevoicing variation on word recognition. Both experiments showed no difference between primes with 12, 6 or 0 periods of prevoicing, even though a third experiment indicated that listeners could discriminate these words. These results are discussed in light of another priming experiment that did show an effect of the absence of prevoicing, but only when primes had a voiceless word competitor. Phonetic detail appears to influence lexical access only when it helps to distinguish between lexical candidates.
  • Van Alphen, P. M., De Bree, E., Fikkert, P., & Wijnen, F. (2007). The role of metrical stress in comprehension and production of Dutch children at risk of dyslexia. In Proceedings of Interspeech 2007 (pp. 2313-2316). Adelaide: Causal Productions.

    Abstract

    The present study compared the role of metrical stress in comprehension and production of three-year-old children with a familial risk of dyslexia with that of normally developing children. A visual fixation task with stress (mis-)matches in bisyllabic words, as well as a non-word repetition task with bisyllabic targets were presented to the control and at-risk children. Results show that the at-risk group is less sensitive to stress mismatches in word recognition than the control group. Correct production of metrical stress patterns did not differ significantly between the groups, but the percentages of phonemes produced correctly were lower for the at-risk than the control group. The findings indicate that processing of metrical stress patterns is not impaired in at-risk children, but that the at-risk group cannot exploit metrical stress in word recognition
  • Wassenaar, M., & Hagoort, P. (2007). Thematic role assignment in patients with Broca's aphasia: Sentence-picture matching electrified. Neuropsychologia, 45(4), 716-740. doi:10.1016/j.neuropsychologia.2006.08.016.

    Abstract

    An event-related brain potential experiment was carried out to investigate on-line thematic role assignment during sentence–picture matching in patients with Broca's aphasia. Subjects were presented with a picture that was followed by an auditory sentence. The sentence either matched the picture or mismatched the visual information depicted. Sentences differed in complexity, and ranged from simple active semantically irreversible sentences to passive semantically reversible sentences. ERPs were recorded while subjects were engaged in sentence–picture matching. In addition, reaction time and accuracy were measured. Three groups of subjects were tested: Broca patients (N = 10), non-aphasic patients with a right hemisphere (RH) lesion (N = 8), and healthy aged-matched controls (N = 15). The results of this study showed that, in neurologically unimpaired individuals, thematic role assignment in the context of visual information was an immediate process. This in contrast to patients with Broca's aphasia who demonstrated no signs of on-line sensitivity to the picture–sentence mismatches. The syntactic contribution to the thematic role assignment process seemed to be diminished given the reduction and even absence of P600 effects. Nevertheless, Broca patients showed some off-line behavioral sensitivity to the sentence–picture mismatches. The long response latencies of Broca's aphasics make it likely that off-line response strategies were used.
  • Willems, R. M., & Hagoort, P. (2007). Neural evidence for the interplay between language, gesture, and action: A review. Brain and Language, 101(3), 278-289. doi:10.1016/j.bandl.2007.03.004.

    Abstract

    Co-speech gestures embody a form of manual action that is tightly coupled to the language system. As such, the co-occurrence of speech and co-speech gestures is an excellent example of the interplay between language and action. There are, however, other ways in which language and action can be thought of as closely related. In this paper we will give an overview of studies in cognitive neuroscience that examine the neural underpinnings of links between language and action. Topics include neurocognitive studies of motor representations of speech sounds, action-related language, sign language and co-speech gestures. It will be concluded that there is strong evidence on the interaction between speech and gestures in the brain. This interaction however shares general properties with other domains in which there is interplay between language and action.
  • Willems, R. M., Ozyurek, A., & Hagoort, P. (2007). When language meets action: The neural integration of gesture and speech. Cerebral Cortex, 17(10), 2322-2333. doi:10.1093/cercor/bhl141.

    Abstract

    Although generally studied in isolation, language and action often co-occur in everyday life. Here we investigated one particular form of simultaneous language and action, namely speech and gestures that speakers use in everyday communication. In a functional magnetic resonance imaging study, we identified the neural networks involved in the integration of semantic information from speech and gestures. Verbal and/or gestural content could be integrated easily or less easily with the content of the preceding part of speech. Premotor areas involved in action observation (Brodmann area [BA] 6) were found to be specifically modulated by action information "mismatching" to a language context. Importantly, an increase in integration load of both verbal and gestural information into prior speech context activated Broca's area and adjacent cortex (BA 45/47). A classical language area, Broca's area, is not only recruited for language-internal processing but also when action observation is integrated with speech. These findings provide direct evidence that action and language processing share a high-level neural integration system.
  • Van Berkum, J. J. A., Brown, C. M., & Hagoort, P. (1999). Early referential context effects in sentence processing: Evidence from event-related brain potentials. Journal of Memory and Language, 41(2), 147-182. doi:10.1006/jmla.1999.2641.

    Abstract

    An event-related brain potentials experiment was carried out to examine the interplay of referential and structural factors during sentence processing in discourse. Subjects read (Dutch) sentences beginning like “David told the girl that … ” in short story contexts that had introduced either one or two referents for a critical singular noun phrase (“the girl”). The waveforms showed that within 280 ms after onset of the critical noun the reader had already determined whether the noun phrase had a unique referent in earlier discourse. Furthermore, this referential information was immediately used in parsing the rest of the sentence, which was briefly ambiguous between a complement clause (“ … that there would be some visitors”) and a relative clause (“ … that had been on the phone to hang up”). A consistent pattern of P600/SPS effects elicited by various subsequent disambiguations revealed that a two-referent discourse context had led the parser to initially pursue the relative-clause alternative to a larger extent than a one-referent context. Together, the results suggest that during the processing of sentences in discourse, structural and referential sources of information interact on a word-by-word basis.

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