Publications

Displaying 1 - 89 of 89
  • Aldosimani, M., Verdonschot, R. G., Iwamoto, Y., Nakazawa, M., Mallya, S. M., Kakimoto, N., Toyosawa, S., Kreiborg, S., & Murakami, S. (2022). Prognostic factors for lymph node metastasis from upper gingival carcinomas. Oral Radiology, 38(3), 389-396. doi:10.1007/s11282-021-00568-w.

    Abstract

    This study sought to identify tumor characteristics that associate with regional lymph node metastases in squamous cell carcinomas originating in the upper gingiva.
  • Arana, S. (2022). Abstract neural representations of language during sentence comprehension: Evidence from MEG and Behaviour. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Bauer, B. L. M. (2022). Counting systems. In A. Ledgeway, & M. Maiden (Eds.), The Cambridge Handbook of Romance Linguistics (pp. 459-488). Cambridge: Cambridge University Press.

    Abstract

    The Romance counting system is numerical – with residues of earlier systems whereby each commodity had its own unit of quantification – and decimal. Numeral formations beyond ‘10’ are compounds, combining two or more numerals that are in an arithmetical relation, typically that of addition and multiplication. Formal variation across the (standard) Romance languages and dialects and across historical stages involves the relative sequence of the composing elements, absence or presence of connectors, their synthetic vs. analytic nature, and the degree of grammatical marking. A number of ‘deviant’ numeral formations raise the question of borrowing vs independent development, such as vigesimals (featuring a base ‘20’ instead ‘10’) in certain Romance varieties and the teen and decad formations in Romanian. The other types of numeral in Romance, which derive from the unmarked and consistent cardinals, feature a significantly higher degree of formal complexity and variation involving Latin formants and tend toward analyticity. While Latin features prominently in the Romance counting system as a source of numeral formations and suffixes, it is only in Romance that the inherited decimal system reached its full potential, illustrating its increasing prominence, reflected not only in numerals, but also in language acquisition, sign language, and post-Revolution measuring systems.
  • Bauer, B. L. M. (2022). Finite verb + infinite + object in later Latin: Early brace constructions? In G. V. M. Haverling (Ed.), Studies on Late and Vulgar Latin in the Early 21st Century: Acts of the 12th International Colloquium "Latin vulgaire – Latin tardif (pp. 166-181). Uppsala: Acta Universitatis Upsaliensis.
  • Bocanegra, B. R., Poletiek, F. H., & Zwaan, R. A. (2022). Language concatenates perceptual features into representations during comprehension. Journal of Memory and Language, 127: 104355. doi:10.1016/j.jml.2022.104355.

    Abstract

    Although many studies have investigated the activation of perceptual representations during language comprehension, to our knowledge only one previous study has directly tested how perceptual features are combined into representations during comprehension. In their classic study, Potter and Faulconer [(1979). Understanding noun phrases. Journal of Verbal Learning and Verbal Behavior, 18, 509–521.] investigated the perceptual representation of adjective-noun combinations. However, their non-orthogonal design did not allow the differentiation between conjunctive vs. disjunctive representations. Using randomized orthogonal designs, we observe evidence for disjunctive perceptual representations when participants represent feature combinations simultaneously (in several experiments; N = 469), and we observe evidence for conjunctive perceptual representations when participants represent feature combinations sequentially (In several experiments; N = 628). Our findings show that the generation of conjunctive representations during comprehension depends on the concatenation of linguistic cues, and thus suggest the construction of elaborate perceptual representations may critically depend on language.
  • Carota, F., Schoffelen, J.-M., Oostenveld, R., & Indefrey, P. (2022). The time course of language production as revealed by pattern classification of MEG sensor data. The Journal of Neuroscience, 42(29), 5745-5754. doi:10.1523/JNEUROSCI.1923-21.2022.

    Abstract

    Language production involves a complex set of computations, from conceptualization to articulation, which are thought to engage cascading neural events in the language network. However, recent neuromagnetic evidence suggests simultaneous meaning-to-speech mapping in picture naming tasks, as indexed by early parallel activation of frontotemporal regions to lexical semantic, phonological, and articulatory information. Here we investigate the time course of word production, asking to what extent such “earliness” is a distinctive property of the associated spatiotemporal dynamics. Using MEG, we recorded the neural signals of 34 human subjects (26 males) overtly naming 134 images from four semantic object categories (animals, foods, tools, clothes). Within each category, we covaried word length, as quantified by the number of syllables contained in a word, and phonological neighborhood density to target lexical and post-lexical phonological/phonetic processes. Multivariate pattern analyses searchlights in sensor space distinguished the stimulus-locked spatiotemporal responses to object categories early on, from 150 to 250 ms after picture onset, whereas word length was decoded in left frontotemporal sensors at 250-350 ms, followed by the latency of phonological neighborhood density (350-450 ms). Our results suggest a progression of neural activity from posterior to anterior language regions for the semantic and phonological/phonetic computations preparing overt speech, thus supporting serial cascading models of word production
  • Carter, G., & Nieuwland, M. S. (2022). Predicting definite and indefinite referents during discourse comprehension: Evidence from event‐related potentials. Cognitive Science, 46(2): e13092. doi:10.1111/cogs.13092.

    Abstract

    Linguistic predictions may be generated from and evaluated against a representation of events and referents described in the discourse. Compatible with this idea, recent work shows that predictions about novel noun phrases include their definiteness. In the current follow-up study, we ask whether people engage similar prediction-related processes for definite and indefinite referents. This question is relevant for linguistic theories that imply a processing difference between definite and indefinite noun phrases, typically because definiteness is thought to require a uniquely identifiable referent in the discourse. We addressed this question in an event-related potential (ERP) study (N = 48) with preregistration of data acquisition, preprocessing, and Bayesian analysis. Participants read Dutch mini-stories with a definite or indefinite novel noun phrase (e.g., “het/een huis,” the/a house), wherein (in)definiteness of the article was either expected or unexpected and the noun was always strongly expected. Unexpected articles elicited enhanced N400s, but unexpectedly indefinite articles also elicited a positive ERP effect at frontal channels compared to expectedly indefinite articles. We tentatively link this effect to an antiuniqueness violation, which may force people to introduce a new referent over and above the already anticipated one. Interestingly, expectedly definite nouns elicited larger N400s than unexpectedly definite nouns (replicating a previous surprising finding) and indefinite nouns. Although the exact nature of these noun effects remains unknown, expectedly definite nouns may have triggered the strongest semantic activation because they alone refer to specific and concrete referents. In sum, results from both the articles and nouns clearly demonstrate that definiteness marking has a rapid effect on processing, counter to recent claims regarding definiteness processing.
  • Coopmans, C. W., De Hoop, H., Kaushik, K., Hagoort, P., & Martin, A. E. (2022). Hierarchy in language interpretation: Evidence from behavioural experiments and computational modelling. Language, Cognition and Neuroscience, 37(4), 420-439. doi:10.1080/23273798.2021.1980595.

    Abstract

    It has long been recognised that phrases and sentences are organised hierarchically, but many computational models of language treat them as sequences of words without computing constituent structure. Against this background, we conducted two experiments which showed that participants interpret ambiguous noun phrases, such as second blue ball, in terms of their abstract hierarchical structure rather than their linear surface order. When a neural network model was tested on this task, it could simulate such “hierarchical” behaviour. However, when we changed the training data such that they were not entirely unambiguous anymore, the model stopped generalising in a human-like way. It did not systematically generalise to novel items, and when it was trained on ambiguous trials, it strongly favoured the linear interpretation. We argue that these models should be endowed with a bias to make generalisations over hierarchical structure in order to be cognitively adequate models of human language.
  • Coopmans, C. W., De Hoop, H., Hagoort, P., & Martin, A. E. (2022). Effects of structure and meaning on cortical tracking of linguistic units in naturalistic speech. Neurobiology of Language, 3(3), 386-412. doi:10.1162/nol_a_00070.

    Abstract

    Recent research has established that cortical activity “tracks” the presentation rate of syntactic phrases in continuous speech, even though phrases are abstract units that do not have direct correlates in the acoustic signal. We investigated whether cortical tracking of phrase structures is modulated by the extent to which these structures compositionally determine meaning. To this end, we recorded electroencephalography (EEG) of 38 native speakers who listened to naturally spoken Dutch stimuli in different conditions, which parametrically modulated the degree to which syntactic structure and lexical semantics determine sentence meaning. Tracking was quantified through mutual information between the EEG data and either the speech envelopes or abstract annotations of syntax, all of which were filtered in the frequency band corresponding to the presentation rate of phrases (1.1–2.1 Hz). Overall, these mutual information analyses showed stronger tracking of phrases in regular sentences than in stimuli whose lexical-syntactic content is reduced, but no consistent differences in tracking between sentences and stimuli that contain a combination of syntactic structure and lexical content. While there were no effects of compositional meaning on the degree of phrase-structure tracking, analyses of event-related potentials elicited by sentence-final words did reveal meaning-induced differences between conditions. Our findings suggest that cortical tracking of structure in sentences indexes the internal generation of this structure, a process that is modulated by the properties of its input, but not by the compositional interpretation of its output.

    Additional information

    supplementary information
  • Coopmans, C. W., & Cohn, N. (2022). An electrophysiological investigation of co-referential processes in visual narrative comprehension. Neuropsychologia, 172: 108253. doi:10.1016/j.neuropsychologia.2022.108253.

    Abstract

    Visual narratives make use of various means to convey referential and co-referential meaning, so comprehenders
    must recognize that different depictions across sequential images represent the same character(s). In this study,
    we investigated how the order in which different types of panels in visual sequences are presented affects how
    the unfolding narrative is comprehended. Participants viewed short comic strips while their electroencephalo-
    gram (EEG) was recorded. We analyzed evoked and induced EEG activity elicited by both full panels (showing a
    full character) and refiner panels (showing only a zoom of that full panel), and took into account whether they
    preceded or followed the panel to which they were co-referentially related (i.e., were cataphoric or anaphoric).
    We found that full panels elicited both larger N300 amplitude and increased gamma-band power compared to
    refiner panels. Anaphoric panels elicited a sustained negativity compared to cataphoric panels, which appeared
    to be sensitive to the referential status of the anaphoric panel. In the time-frequency domain, anaphoric panels
    elicited reduced 8–12 Hz alpha power and increased 45–65 Hz gamma-band power compared to cataphoric
    panels. These findings are consistent with models in which the processes involved in visual narrative compre-
    hension partially overlap with those in language comprehension.
  • Corps, R. E., Knudsen, B., & Meyer, A. S. (2022). Overrated gaps: Inter-speaker gaps provide limited information about the timing of turns in conversation. Cognition, 223: 105037. doi:10.1016/j.cognition.2022.105037.

    Abstract

    Corpus analyses have shown that turn-taking in conversation is much faster than laboratory studies of speech planning would predict. To explain fast turn-taking, Levinson and Torreira (2015) proposed that speakers are highly proactive: They begin to plan a response to their interlocutor's turn as soon as they have understood its gist, and launch this planned response when the turn-end is imminent. Thus, fast turn-taking is possible because speakers use the time while their partner is talking to plan their own utterance. In the present study, we asked how much time upcoming speakers actually have to plan their utterances. Following earlier psycholinguistic work, we used transcripts of spoken conversations in Dutch, German, and English. These transcripts consisted of segments, which are continuous stretches of speech by one speaker. In the psycholinguistic and phonetic literature, such segments have often been used as proxies for turns. We found that in all three corpora, large proportions of the segments comprised of only one or two words, which on our estimate does not give the next speaker enough time to fully plan a response. Further analyses showed that speakers indeed often did not respond to the immediately preceding segment of their partner, but continued an earlier segment of their own. More generally, our findings suggest that speech segments derived from transcribed corpora do not necessarily correspond to turns, and the gaps between speech segments therefore only provide limited information about the planning and timing of turns.
  • Dai, B., McQueen, J. M., Terporten, R., Hagoort, P., & Kösem, A. (2022). Distracting Linguistic Information Impairs Neural Tracking of Attended Speech. Current Research in Neurobiology, 3: 100043. doi:10.1016/j.crneur.2022.100043.

    Abstract

    Listening to speech is difficult in noisy environments, and is even harder when the interfering noise consists of intelligible speech as compared to unintelligible sounds. This suggests that the competing linguistic information interferes with the neural processing of target speech. Interference could either arise from a degradation of the neural representation of the target speech, or from increased representation of distracting speech that enters in competition with the target speech. We tested these alternative hypotheses using magnetoencephalography (MEG) while participants listened to a target clear speech in the presence of distracting noise-vocoded speech. Crucially, the distractors were initially unintelligible but became more intelligible after a short training session. Results showed that the comprehension of the target speech was poorer after training than before training. The neural tracking of target speech in the delta range (1–4 Hz) reduced in strength in the presence of a more intelligible distractor. In contrast, the neural tracking of distracting signals was not significantly modulated by intelligibility. These results suggest that the presence of distracting speech signals degrades the linguistic representation of target speech carried by delta oscillations.
  • Dijkstra, T., Peeters, D., Hieselaar, W., & van Geffen, A. (2022). Orthographic and semantic priming effects in neighbour cognates: Experiments and simulations. Bilingualism: Language and Cognition, 26(2), 371-383. doi:10.1017/S1366728922000591.

    Abstract

    To investigate how orthography and semantics interact during bilingual visual word recognition, Dutch–English bilinguals made lexical decisions in two masked priming experiments. Dutch primes and English targets were presented that were either neighbour cognates (boek – BOOK), noncognate translations (kooi – CAGE), orthographically related neighbours (neus – NEWS), or unrelated words (huid - COAT). Prime durations of 50 ms (Experiment 1) and 83 ms (Experiment 2) led to similar result patterns. Both experiments reported a large cognate facilitation effect, a smaller facilitatory noncognate translation effect, and the absence of inhibitory orthographic neighbour effects. These results indicate that cognate facilitation is in large part due to orthographic-semantic resonance. Priming results for each condition were simulated well (all r's >.50) by Multilink+, a recent computational model for word retrieval. Limitations to the role of lateral inhibition in bilingual word recognition are discussed.
  • Drijvers, L., & Holler, J. (2022). Face-to-face spatial orientation fine-tunes the brain for neurocognitive processing in conversation. iScience, 25(11): 105413. doi:10.1016/j.isci.2022.105413.

    Abstract

    We here demonstrate that face-to-face spatial orientation induces a special ‘social mode’ for neurocognitive processing during conversation, even in the absence of visibility. Participants conversed face-to-face, face-to-face but visually occluded, and back-to-back to tease apart effects caused by seeing visual communicative signals and by spatial orientation. Using dual-EEG, we found that 1) listeners’ brains engaged more strongly while conversing in face-to-face than back-to-back, irrespective of the visibility of communicative signals, 2) listeners attended to speech more strongly in a back-to-back compared to a face-to-face spatial orientation without visibility; visual signals further reduced the attention needed; 3) the brains of interlocutors were more in sync in a face-to-face compared to a back-to-back spatial orientation, even when they could not see each other; visual signals further enhanced this pattern. Communicating in face-to-face spatial orientation is thus sufficient to induce a special ‘social mode’ which fine-tunes the brain for neurocognitive processing in conversation.
  • Eekhof, L. S., Van Krieken, K., & Willems, R. M. (2022). Reading about minds: The social-cognitive potential of narratives. Psychonomic Bulletin & Review, 29, 1703-1718. doi:10.3758/s13423-022-02079-z.

    Abstract

    It is often argued that narratives improve social cognition, either by appealing to social-cognitive abilities as we engage with the story world and its characters, or by conveying social knowledge. Empirical studies have found support for both a correlational and a causal link between exposure to (literary, fictional) narratives and social cognition. However, a series of failed replications has cast doubt on the robustness of these claims. Here, we review the existing empirical literature and identify open questions and challenges. An important conclusion of the review is that previous research has given too little consideration to the diversity of narratives, readers, and social-cognitive processes involved in the social-cognitive potential of narratives. We therefore establish a research agenda, proposing that future research should focus on (1) the specific text characteristics that drive the social-cognitive potential of narratives, (2) the individual differences between readers with respect to their sensitivity to this potential, and (3) the various aspects of social cognition that are potentially affected by reading narratives. Our recommendations can guide the design of future studies that will help us understand how, for whom, and in what respect exposure to narratives can advantage social cognition.
  • Ferrari, A., Richter, D., & De Lange, F. (2022). Updating contextual sensory expectations for adaptive behaviour. The Journal of Neuroscience, 42(47), 8855-8869. doi:10.1523/JNEUROSCI.1107-22.2022.

    Abstract

    The brain has the extraordinary capacity to construct predictive models of the environment by internalizing statistical regularities in the sensory inputs. The resulting sensory expectations shape how we perceive and react to the world; at the neural level, this relates to decreased neural responses to expected than unexpected stimuli (‘expectation suppression’). Crucially, expectations may need revision as context changes. However, existing research has often neglected this issue. Further, it is unclear whether contextual revisions apply selectively to expectations relevant to the task at hand, hence serving adaptive behaviour. The present fMRI study examined how contextual visual expectations spread throughout the cortical hierarchy as participants update their beliefs. We created a volatile environment with two state spaces presented over separate contexts and controlled by an independent contextualizing signal. Participants attended a training session before scanning to learn contextual temporal associations among pairs of object images. The fMRI experiment then tested for the emergence of contextual expectation suppression in two separate tasks, respectively with task-relevant and task-irrelevant expectations. Behavioural and neural effects of contextual expectation emerged progressively across the cortical hierarchy as participants attuned themselves to the context: expectation suppression appeared first in the insula, inferior frontal gyrus and posterior parietal cortex, followed by the ventral visual stream, up to early visual cortex. This applied selectively to task-relevant expectations. Taken together, the present results suggest that an insular and frontoparietal executive control network may guide the flexible deployment of contextual sensory expectations for adaptive behaviour in our complex and dynamic world.
  • Gao, Y., Meng, X., Bai, Z., Liu, X., Zhang, M., Li, H., Ding, G., Liu, L., & Booth, J. R. (2022). Left and right arcuate fasciculi are uniquely related to word reading skills in Chinese-English bilingual children. Neurobiology of Language, 3(1), 109-131. doi:10.1162/nol_a_00051.

    Abstract

    Whether reading in different writing systems recruits language-unique or language-universal neural processes is a long-standing debate. Many studies have shown the left arcuate fasciculus (AF) to be involved in phonological and reading processes. In contrast, little is known about the role of the right AF in reading, but some have suggested that it may play a role in visual spatial aspects of reading or the prosodic components of language. The right AF may be more important for reading in Chinese due to its logographic and tonal properties, but this hypothesis has yet to be tested. We recruited a group of Chinese-English bilingual children (8.2 to 12.0 years old) to explore the common and unique relation of reading skill in English and Chinese to fractional anisotropy (FA) in the bilateral AF. We found that both English and Chinese reading skills were positively correlated with FA in the rostral part of the left AF-direct segment. Additionally, English reading skill was positively correlated with FA in the caudal part of the left AF-direct segment, which was also positively correlated with phonological awareness. In contrast, Chinese reading skill was positively correlated with FA in certain segments of the right AF, which was positively correlated with visual spatial ability, but not tone discrimination ability. Our results suggest that there are language universal substrates of reading across languages, but that certain left AF nodes support phonological mechanisms important for reading in English, whereas certain right AF nodes support visual spatial mechanisms important for reading in Chinese.

    Additional information

    supplementary materials
  • Giglio, L., Ostarek, M., Weber, K., & Hagoort, P. (2022). Commonalities and asymmetries in the neurobiological infrastructure for language production and comprehension. Cerebral Cortex, 32(7), 1405-1418. doi:10.1093/cercor/bhab287.

    Abstract

    The neurobiology of sentence production has been largely understudied compared to the neurobiology of sentence comprehension, due to difficulties with experimental control and motion-related artifacts in neuroimaging. We studied the neural response to constituents of increasing size and specifically focused on the similarities and differences in the production and comprehension of the same stimuli. Participants had to either produce or listen to stimuli in a gradient of constituent size based on a visual prompt. Larger constituent sizes engaged the left inferior frontal gyrus (LIFG) and middle temporal gyrus (LMTG) extending to inferior parietal areas in both production and comprehension, confirming that the neural resources for syntactic encoding and decoding are largely overlapping. An ROI analysis in LIFG and LMTG also showed that production elicited larger responses to constituent size than comprehension and that the LMTG was more engaged in comprehension than production, while the LIFG was more engaged in production than comprehension. Finally, increasing constituent size was characterized by later BOLD peaks in comprehension but earlier peaks in production. These results show that syntactic encoding and parsing engage overlapping areas, but there are asymmetries in the engagement of the language network due to the specific requirements of production and comprehension.

    Additional information

    supplementary material
  • Gussenhoven, C., Lu, Y.-A., Lee-Kim, S.-I., Liu, C., Rahmani, H., Riad, T., & Zora, H. (2022). The sequence recall task and lexicality of tone: Exploring tone “deafness”. Frontiers in Psychology, 13: 902569. doi:10.3389/fpsyg.2022.902569.

    Abstract

    Many perception and processing effects of the lexical status of tone have been found in behavioral, psycholinguistic, and neuroscientific research, often pitting varieties of tonal Chinese against non-tonal Germanic languages. While the linguistic and cognitive evidence for lexical tone is therefore beyond dispute, the word prosodic systems of many languages continue to escape the categorizations of typologists. One controversy concerns the existence of a typological class of “pitch accent languages,” another the underlying phonological nature of surface tone contrasts, which in some cases have been claimed to be metrical rather than tonal. We address the question whether the Sequence Recall Task (SRT), which has been shown to discriminate between languages with and without word stress, can distinguish languages with and without lexical tone. Using participants from non-tonal Indonesian, semi-tonal Swedish, and two varieties of tonal Mandarin, we ran SRTs with monosyllabic tonal contrasts to test the hypothesis that high performance in a tonal SRT indicates the lexical status of tone. An additional question concerned the extent to which accuracy scores depended on phonological and phonetic properties of a language’s tone system, like its complexity, the existence of an experimental contrast in a language’s phonology, and the phonetic salience of a contrast. The results suggest that a tonal SRT is not likely to discriminate between tonal and non-tonal languages within a typologically varied group, because of the effects of specific properties of their tone systems. Future research should therefore address the first hypothesis with participants from otherwise similar tonal and non-tonal varieties of the same language, where results from a tonal SRT may make a useful contribution to the typological debate on word prosody.

    Additional information

    also published as book chapter (2023)
  • Hagoort, P. (2022). Reasoning and the brain. In M. Stokhof, & K. Stenning (Eds.), Rules, regularities, randomness. Festschrift for Michiel van Lambalgen (pp. 83-85). Amsterdam: Institute for Logic, Language and Computation.
  • Heilbron, M., Armeni, K., Schoffelen, J.-M., Hagoort, P., & De Lange, F. P. (2022). A hierarchy of linguistic predictions during natural language comprehension. Proceedings of the National Academy of Sciences of the United States of America, 119(32): e2201968119. doi:10.1073/pnas.2201968119.

    Abstract

    Understanding spoken language requires transforming ambiguous acoustic streams into a hierarchy of representations, from phonemes to meaning. It has been suggested that the brain uses prediction to guide the interpretation of incoming input. However, the role of prediction in language processing remains disputed, with disagreement about both the ubiquity and representational nature of predictions. Here, we address both issues by analyzing brain recordings of participants listening to audiobooks, and using a deep neural network (GPT-2) to precisely quantify contextual predictions. First, we establish that brain responses to words are modulated by ubiquitous predictions. Next, we disentangle model-based predictions into distinct dimensions, revealing dissociable neural signatures of predictions about syntactic category (parts of speech), phonemes, and semantics. Finally, we show that high-level (word) predictions inform low-level (phoneme) predictions, supporting hierarchical predictive processing. Together, these results underscore the ubiquity of prediction in language processing, showing that the brain spontaneously predicts upcoming language at multiple levels of abstraction.

    Additional information

    supporting information
  • Heilbron, M. (2022). Getting ahead: Prediction as a window into language, and language as a window into the predictive brain. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Hervais-Adelman, A., Kumar, U., Mishra, R., Tripathi, V., Guleria, A., Singh, J. P., & Huettig, F. (2022). How does literacy affect speech processing? Not by enhancing cortical responses to speech, but by promoting connectivity of acoustic-phonetic and graphomotor cortices. Journal of Neuroscience, 42(47), 8826-8841. doi:10.1523/JNEUROSCI.1125-21.2022.

    Abstract

    Previous research suggests that literacy, specifically learning alphabetic letter-to-phoneme mappings, modifies online speech processing, and enhances brain responses, as indexed by the blood-oxygenation level dependent signal (BOLD), to speech in auditory areas associated with phonological processing (Dehaene et al., 2010). However, alphabets are not the only orthographic systems in use in the world, and hundreds of millions of individuals speak languages that are not written using alphabets. In order to make claims that literacy per se has broad and general consequences for brain responses to speech, one must seek confirmatory evidence from non-alphabetic literacy. To this end, we conducted a longitudinal fMRI study in India probing the effect of literacy in Devanagari, an abugida, on functional connectivity and cerebral responses to speech in 91 variously literate Hindi-speaking male and female human participants. Twenty-two completely illiterate participants underwent six months of reading and writing training. Devanagari literacy increases functional connectivity between acoustic-phonetic and graphomotor brain areas, but we find no evidence that literacy changes brain responses to speech, either in cross-sectional or longitudinal analyses. These findings shows that a dramatic reconfiguration of the neurofunctional substrates of online speech processing may not be a universal result of learning to read, and suggest that the influence of writing on speech processing should also be investigated.
  • Hoeksema, N., Hagoort, P., & Vernes, S. C. (2022). Piecing together the building blocks of the vocal learning bat brain. In A. Ravignani, R. Asano, D. Valente, F. Ferretti, S. Hartmann, M. Hayashi, Y. Jadoul, M. Martins, Y. Oseki, E. D. Rodrigues, O. Vasileva, & S. Wacewicz (Eds.), The evolution of language: Proceedings of the Joint Conference on Language Evolution (JCoLE) (pp. 294-296). Nijmegen: Joint Conference on Language Evolution (JCoLE).
  • Holler, J., Drijvers, L., Rafiee, A., & Majid, A. (2022). Embodied space-pitch associations are shaped by language. Cognitive Science, 46(2): e13083. doi:10.1111/cogs.13083.

    Abstract

    Height-pitch associations are claimed to be universal and independent of language, but this claim remains controversial. The present study sheds new light on this debate with a multimodal analysis of individual sound and melody descriptions obtained in an interactive communication paradigm with speakers of Dutch and Farsi. The findings reveal that, in contrast to Dutch speakers, Farsi speakers do not use a height-pitch metaphor consistently in speech. Both Dutch and Farsi speakers’ co-speech gestures did reveal a mapping of higher pitches to higher space and lower pitches to lower space, and this gesture space-pitch mapping tended to co-occur with corresponding spatial words (high-low). However, this mapping was much weaker in Farsi speakers than Dutch speakers. This suggests that cross-linguistic differences shape the conceptualization of pitch and further calls into question the universality of height-pitch associations.

    Additional information

    supporting information
  • Huizeling, E., Arana, S., Hagoort, P., & Schoffelen, J.-M. (2022). Lexical frequency and sentence context influence the brain’s response to single words. Neurobiology of Language, 3(1), 149-179. doi:10.1162/nol_a_00054.

    Abstract

    Typical adults read remarkably quickly. Such fast reading is facilitated by brain processes that are sensitive to both word frequency and contextual constraints. It is debated as to whether these attributes have additive or interactive effects on language processing in the brain. We investigated this issue by analysing existing magnetoencephalography data from 99 participants reading intact and scrambled sentences. Using a cross-validated model comparison scheme, we found that lexical frequency predicted the word-by-word elicited MEG signal in a widespread cortical network, irrespective of sentential context. In contrast, index (ordinal word position) was more strongly encoded in sentence words, in left front-temporal areas. This confirms that frequency influences word processing independently of predictability, and that contextual constraints affect word-by-word brain responses. With a conservative multiple comparisons correction, only the interaction between lexical frequency and surprisal survived, in anterior temporal and frontal cortex, and not between lexical frequency and entropy, nor between lexical frequency and index. However, interestingly, the uncorrected index*frequency interaction revealed an effect in left frontal and temporal cortex that reversed in time and space for intact compared to scrambled sentences. Finally, we provide evidence to suggest that, in sentences, lexical frequency and predictability may independently influence early (<150ms) and late stages of word processing, but interact during later stages of word processing (>150-250ms), thus helping to converge previous contradictory eye-tracking and electrophysiological literature. Current neuro-cognitive models of reading would benefit from accounting for these differing effects of lexical frequency and predictability on different stages of word processing.
  • Huizeling, E., Peeters, D., & Hagoort, P. (2022). Prediction of upcoming speech under fluent and disfluent conditions: Eye tracking evidence from immersive virtual reality. Language, Cognition and Neuroscience, 37(4), 481-508. doi:10.1080/23273798.2021.1994621.

    Abstract

    Traditional experiments indicate that prediction is important for efficient speech processing. In three virtual reality visual world paradigm experiments, we tested whether such findings hold in naturalistic settings (Experiment 1) and provided novel insights into whether disfluencies in speech (repairs/hesitations) inform one’s predictions in rich environments (Experiments 2–3). Experiment 1 supports that listeners predict upcoming speech in naturalistic environments, with higher proportions of anticipatory target fixations in predictable compared to unpredictable trials. In Experiments 2–3, disfluencies reduced anticipatory fixations towards predicted referents, compared to conjunction (Experiment 2) and fluent (Experiment 3) sentences. Unexpectedly, Experiment 2 provided no evidence that participants made new predictions from a repaired verb. Experiment 3 provided novel findings that fixations towards the speaker increase upon hearing a hesitation, supporting current theories of how hesitations influence sentence processing. Together, these findings unpack listeners’ use of visual (objects/speaker) and auditory (speech/disfluencies) information when predicting upcoming words.
  • Lai, V. T., Van Berkum, J. J. A., & Hagoort, P. (2022). Negative affect increases reanalysis of conflicts between discourse context and world knowledge. Frontiers in Communication, 7: 910482. doi:10.3389/fcomm.2022.910482.

    Abstract

    Introduction: Mood is a constant in our daily life and can permeate all levels of cognition. We examined whether and how mood influences the processing of discourse content that is relatively neutral and not loaded with emotion. During discourse processing, readers have to constantly strike a balance between what they know in long term memory and what the current discourse is about. Our general hypothesis is that mood states would affect this balance. We hypothesized that readers in a positive mood would rely more on default world knowledge, whereas readers in a negative mood would be more inclined to analyze the details in the current discourse.

    Methods: Participants were put in a positive and a negative mood via film clips, one week apart. In each session, after mood manipulation, they were presented with sentences in discourse materials. We created sentences such as “With the lights on you can see...” that end with critical words (CWs) “more” or “less”, where general knowledge supports “more”, not “less”. We then embedded each of these sentences in a wider discourse that does/does not support the CWs (a story about driving in the night vs. stargazing). EEG was recorded throughout.

    Results: The results showed that first, mood manipulation was successful in that there was a significant mood difference between sessions. Second, mood did not modulate the N400 effects. Participants in both moods detected outright semantic violations and allowed world knowledge to be overridden by discourse context. Third, mood modulated the LPC (Late Positive Component) effects, distributed in the frontal region. In negative moods, the LPC was sensitive to one-level violation. That is, CWs that were supported by only world knowledge, only discourse, and neither, elicited larger frontal LPCs, in comparison to the condition where CWs were supported by both world knowledge and discourse.

    Discussion: These results suggest that mood does not influence all processes involved in discourse processing. Specifically, mood does not influence lexical-semantic retrieval (N400), but it does influence elaborative processes for sensemaking (P600) during discourse processing. These results advance our understanding of the impact and time course of mood on discourse.

    Additional information

    Table 1.XLSX
  • Levshina, N. (2022). Frequency, informativity and word length: Insights from typologically diverse corpora. Entropy, 24(2): 280. doi:10.3390/e24020280.

    Abstract

    Zipf’s law of abbreviation, which posits a negative correlation between word frequency and length, is one of the most famous and robust cross-linguistic generalizations. At the same time, it has been shown that contextual informativity (average surprisal given previous context) is more strongly correlated with word length, although this tendency is not observed consistently, depending on several methodological choices. The present study examines a more diverse sample of languages than the previous studies (Arabic, Finnish, Hungarian, Indonesian, Russian, Spanish and Turkish). I use large web-based corpora from the Leipzig Corpora Collection to estimate word lengths in UTF-8 characters and in phonemes (for some of the languages), as well as word frequency, informativity given previous word and informativity given next word, applying different methods of bigrams processing. The results show different correlations between word length and the corpus-based measure for different languages. I argue that these differences can be explained by the properties of noun phrases in a language, most importantly, by the order of heads and modifiers and their relative morphological complexity, as well as by orthographic conventions

    Additional information

    datasets
  • Levshina, N., & Hawkins, J. A. (2022). Verb-argument lability and its correlations with other typological parameters. A quantitative corpus-based study. Linguistic Typology at the Crossroads, 2(1), 94-120. doi:10.6092/issn.2785-0943/13861.

    Abstract

    We investigate the correlations between A- and P-lability for verbal arguments with other typological parameters using large, syntactically annotated corpora of online news in 28 languages. To estimate how much lability is observed in a language, we measure associations between Verbs or Verb + Noun combinations and the alternating constructions in which they occur. Our correlational analyses show that high P-lability scores correlate strongly with the following parameters: little or no case marking; weaker associations between lexemes and the grammatical roles A and P; rigid order of Subject and Object; and a high proportion of verb-medial clauses (SVO). Low P-lability correlates with the presence of case marking, stronger associations between nouns and grammatical roles, relatively flexible ordering of Subject and Object, and verb-final order. As for A-lability, it is not correlated with any other parameters. A possible reason is that A-lability is a result of more universal discourse processes, such as deprofiling of the object, and also exhibits numerous lexical and semantic idiosyncrasies. The fact that P-lability is strongly correlated with other parameters can be interpreted as evidence for a more general typology of languages, in which some tend to have highly informative morphosyntactic and lexical cues, whereas others rely predominantly on contextual environment, which is possibly due to fixed word order. We also find that P-lability is more strongly correlated with the other parameters than any of these parameters are with each other, which means that it can be a very useful typological variable.
  • Levshina, N., & Lorenz, D. (2022). Communicative efficiency and the Principle of No Synonymy: Predictability effects and the variation of want to and wanna. Language and Cognition, 14(2), 249-274. doi:10.1017/langcog.2022.7.

    Abstract

    There is ample psycholinguistic evidence that speakers behave efficiently, using shorter and less effortful constructions when the meaning is more predictable, and longer and more effortful ones when it is less predictable. However, the Principle of No Synonymy requires that all formally distinct variants should also be functionally different. The question is how much two related constructions should overlap semantically and pragmatically in order to be used for the purposes of efficient communication. The case study focuses on want to + Infinitive and its reduced variant with wanna, which have different stylistic and sociolinguistic connotations. Bayesian mixed-effects regression modelling based on the spoken part of the British National Corpus reveals a very limited effect of efficiency: predictability increases the chances of the reduced variant only in fast speech. We conclude that efficient use of more and less effortful variants is restricted when two variants are associated with different registers or styles. This paper also pursues a methodological goal regarding missing values in speech corpora. We impute missing data based on the existing values. A comparison of regression models with and without imputed values reveals similar tendencies. This means that imputation is useful for dealing with missing values in corpora.

    Additional information

    supplementary materials
  • Levshina, N. (2022). Semantic maps of causation: New hybrid approaches based on corpora and grammar descriptions. Zeitschrift für Sprachwissenschaft, 41(1), 179-205. doi:10.1515/zfs-2021-2043.

    Abstract

    The present paper discusses connectivity and proximity maps of causative constructions and combines them with different types of typological data. In the first case study, I show how one can create a connectivity map based on a parallel corpus. This allows us to solve many problems, such as incomplete descriptions, inconsistent terminology and the problem of determining the semantic nodes. The second part focuses on proximity maps based on Multidimensional Scaling and compares the most important semantic distinctions, which are inferred from a parallel corpus of film subtitles and from grammar descriptions. The results suggest that corpus-based maps of tokens are more sensitive to cultural and genre-related differences in the prominence of specific causation scenarios than maps based on constructional types, which are described in reference grammars. The grammar-based maps also reveal a less clear structure, which can be due to incomplete semantic descriptions in grammars. Therefore, each approach has its shortcomings, which researchers need to be aware of.
  • Levshina, N. (2022). Corpus-based typology: Applications, challenges and some solutions. Linguistic Typology, 26(1), 129-160. doi:10.1515/lingty-2020-0118.

    Abstract

    Over the last few years, the number of corpora that can be used for language comparison has dramatically increased. The corpora are so diverse in their structure, size and annotation style, that a novice might not know where to start. The present paper charts this new and changing territory, providing a few landmarks, warning signs and safe paths. Although no corpora corpus at present can replace the traditional type of typological data based on language description in reference grammars, they corpora can help with diverse tasks, being particularly well suited for investigating probabilistic and gradient properties of languages and for discovering and interpreting cross-linguistic generalizations based on processing and communicative mechanisms. At the same time, the use of corpora for typological purposes has not only advantages and opportunities, but also numerous challenges. This paper also contains an empirical case study addressing two pertinent problems: the role of text types in language comparison and the problem of the word as a comparative concept.
  • Levshina, N. (2022). Comparing Bayesian and frequentist models of language variation: The case of help + (to) Infinitive. In O. Schützler, & J. Schlüter (Eds.), Data and methods in corpus linguistics – Comparative Approaches (pp. 224-258). Cambridge: Cambridge University Press.
  • Mak, M., Faber, M., & Willems, R. M. (2022). Different routes to liking: How readers arrive at narrative evaluations. Cognitive Research: Principles and implications, 7: 72. doi:10.1186/s41235-022-00419-0.

    Abstract

    When two people read the same story, they might both end up liking it very much. However, this does not necessarily mean that their reasons for liking it were identical. We therefore ask what factors contribute to “liking” a story, and—most importantly—how people vary in this respect. We found that readers like stories because they find them interesting, amusing, suspenseful and/or beautiful. However, the degree to which these components of appreciation were related to how much readers liked stories differed between individuals. Interestingly, the individual slopes of the relationships between many of the components and liking were (positively or negatively) correlated. This indicated, for instance, that individuals displaying a relatively strong relationship between interest and liking, generally display a relatively weak relationship between sadness and liking. The individual differences in the strengths of the relationships between the components and liking were not related to individual differences in expertize, a characteristic strongly associated with aesthetic appreciation of visual art. Our work illustrates that it is important to take into consideration the fact that individuals differ in how they arrive at their evaluation of literary stories, and that it is possible to quantify these differences in empirical experiments. Our work suggests that future research should be careful about “overfitting” theories of aesthetic appreciation to an “idealized reader,” but rather take into consideration variations across individuals in the reason for liking a particular story.
  • Misersky, J. (2022). About time: Exploring the role of grammatical aspect in event cognition. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Misersky, J., Peeters, D., & Flecken, M. (2022). The potential of immersive virtual reality for the study of event perception. Frontiers in Virtual Reality, 3: 697934. doi:10.3389/frvir.2022.697934.

    Abstract

    In everyday life, we actively engage in different activities from a first-person perspective. However, experimental psychological research in the field of event perception is often limited to relatively passive, third-person computer-based paradigms. In the present study, we tested the feasibility of using immersive virtual reality in combination with eye tracking with participants in active motion. Behavioral research has shown that speakers of aspectual and non-aspectual languages attend to goals (endpoints) in motion events differently, with speakers of non-aspectual languages showing relatively more attention to goals (endpoint bias). In the current study, native speakers of German (non-aspectual) and English (aspectual) walked on a treadmill across 3-D terrains in VR, while their eye gaze was continuously tracked. Participants encountered landmark objects on the side of the road, and potential endpoint objects at the end of it. Using growth curve analysis to analyze fixation patterns over time, we found no differences in eye gaze behavior between German and English speakers. This absence of cross-linguistic differences was also observed in behavioral tasks with the same participants. Methodologically, based on the quality of the data, we conclude that our dynamic eye-tracking setup can be reliably used to study what people look at while moving through rich and dynamic environments that resemble the real world.
  • Montero-Melis, G., Van Paridon, J., Ostarek, M., & Bylund, E. (2022). No evidence for embodiment: The motor system is not needed to keep action words in working memory. Cortex, 150, 108-125. doi:10.1016/j.cortex.2022.02.006.

    Abstract

    Increasing evidence implicates the sensorimotor systems with high-level cognition, but the extent to which these systems play a functional role remains debated. Using an elegant design, Shebani and Pulvermüller (2013) reported that carrying out a demanding rhythmic task with the hands led to selective impairment of working memory for hand-related words (e.g., clap), while carrying out the same task with the feet led to selective memory impairment for foot-related words (e.g., kick). Such a striking double dissociation is acknowledged even by critics to constitute strong evidence for an embodied account of working memory. Here, we report on an attempt at a direct replication of this important finding. We followed a sequential sampling design and stopped data collection at N=77 (more than five times the original sample size), at which point the evidence for the lack of the critical selective interference effect was very strong (BF01 = 91). This finding constitutes strong evidence against a functional contribution of the motor system to keeping action words in working memory. Our finding fits into the larger emerging picture in the field of embodied cognition that sensorimotor simulations are neither required nor automatic in high-level cognitive processes, but that they may play a role depending on the task. Importantly, we urge researchers to engage in transparent, high-powered, and fully pre-registered experiments like the present one to ensure the field advances on a solid basis.
  • Morey, R. D., Kaschak, M. P., Díez-Álamo, A. M., Glenberg, A. M., Zwaan, R. A., Lakens, D., Ibáñez, A., García, A., Gianelli, C., Jones, J. L., Madden, J., Alifano, F., Bergen, B., Bloxsom, N. G., Bub, D. N., Cai, Z. G., Chartier, C. R., Chatterjee, A., Conwell, E., Cook, S. W. and 25 moreMorey, R. D., Kaschak, M. P., Díez-Álamo, A. M., Glenberg, A. M., Zwaan, R. A., Lakens, D., Ibáñez, A., García, A., Gianelli, C., Jones, J. L., Madden, J., Alifano, F., Bergen, B., Bloxsom, N. G., Bub, D. N., Cai, Z. G., Chartier, C. R., Chatterjee, A., Conwell, E., Cook, S. W., Davis, J. D., Evers, E., Girard, S., Harter, D., Hartung, F., Herrera, E., Huettig, F., Humphries, S., Juanchich, M., Kühne, K., Lu, S., Lynes, T., Masson, M. E. J., Ostarek, M., Pessers, S., Reglin, R., Steegen, S., Thiessen, E. D., Thomas, L. E., Trott, S., Vandekerckhove, J., Vanpaemel, W., Vlachou, M., Williams, K., & Ziv-Crispel, N. (2022). A pre-registered, multi-lab non-replication of the Action-sentence Compatibility Effect (ACE). Psychonomic Bulletin & Review, 29, 613-626. doi:10.3758/s13423-021-01927-8.

    Abstract

    The Action-sentence Compatibility Effect (ACE) is a well-known demonstration of the role of motor activity in the comprehension of language. Participants are asked to make sensibility judgments on sentences by producing movements toward the body or away from the body. The ACE is the finding that movements are faster when the direction of the movement (e.g., toward) matches the direction of the action in the to-be-judged sentence (e.g., Art gave you the pen describes action toward you). We report on a pre- registered, multi-lab replication of one version of the ACE. The results show that none of the 18 labs involved in the study observed a reliable ACE, and that the meta-analytic estimate of the size of the ACE was essentially zero.
  • Murphy, E., Woolnough, O., Rollo, P. S., Roccaforte, Z., Segaert, K., Hagoort, P., & Tandon, N. (2022). Minimal phrase composition revealed by intracranial recordings. The Journal of Neuroscience, 42(15), 3216-3227. doi:10.1523/JNEUROSCI.1575-21.2022.

    Abstract

    The ability to comprehend phrases is an essential integrative property of the brain. Here we evaluate the neural processes that enable the transition from single word processing to a minimal compositional scheme. Previous research has reported conflicting timing effects of composition, and disagreement persists with respect to inferior frontal and posterior temporal contributions. To address these issues, 19 patients (10 male, 19 female) implanted with penetrating depth or surface subdural intracranial electrodes heard auditory recordings of adjective-noun, pseudoword-noun and adjective-pseudoword phrases and judged whether the phrase matched a picture. Stimulus-dependent alterations in broadband gamma activity, low frequency power and phase-locking values across the language-dominant left hemisphere were derived. This revealed a mosaic located on the lower bank of the posterior superior temporal sulcus (pSTS), in which closely neighboring cortical sites displayed exclusive sensitivity to either lexicality or phrase structure, but not both. Distinct timings were found for effects of phrase composition (210–300 ms) and pseudoword processing (approximately 300–700 ms), and these were localized to neighboring electrodes in pSTS. The pars triangularis and temporal pole encoded anticipation of composition in broadband low frequencies, and both regions exhibited greater functional connectivity with pSTS during phrase composition. Our results suggest that the pSTS is a highly specialized region comprised of sparsely interwoven heterogeneous constituents that encodes both lower and higher level linguistic features. This hub in pSTS for minimal phrase processing may form the neural basis for the human-specific computational capacity for forming hierarchically organized linguistic structures.
  • Poort, E. D., & Rodd, J. M. (2022). Cross-lingual priming of cognates and interlingual homographs from L2 to L1. Glossa Psycholinguistics, 1(1): 11. doi:10.5070/G601147.

    Abstract

    Many word forms exist in multiple languages, and can have either the same meaning (cognates) or a different meaning (interlingual homographs). Previous experiments have shown that processing of interlingual homographs in a bilingual’s second language is slowed down by recent experience with these words in the bilingual’s native language, while processing of cognates can be speeded up (Poort et al., 2016; Poort & Rodd, 2019a). The current experiment replicated Poort and Rodd’s (2019a) Experiment 2 but switched the direction of priming: Dutch–English bilinguals (n = 106) made Dutch semantic relatedness judgements to probes related to cognates (n = 50), interlingual homographs (n = 50) and translation equivalents (n = 50) they had seen 15 minutes previously embedded in English sentences. The current experiment is the first to show that a single encounter with an interlingual homograph in one’s second language can also affect subsequent processing in one’s native language. Cross-lingual priming did not affect the cognates. The experiment also extended Poort and Rodd (2019a)’s finding of a large interlingual homograph inhibition effect in a semantic relatedness task in the participants’ L2 to their L1, but again found no evidence for a cognate facilitation effect in a semantic relatedness task. These findings extend the growing literature that emphasises the high level of interaction in a bilingual’s mental lexicon, by demonstrating the influence of L2 experience on the processing of L1 words. Data, scripts, materials and pre-registration available via https://osf.io/2swyg/?view_only=b2ba2e627f6f4eaeac87edab2b59b236.
  • Poulton, V. R., & Nieuwland, M. S. (2022). Can you hear what’s coming? Failure to replicate ERP evidence for phonological prediction. Neurobiology of Language, 3(4), 556 -574. doi:10.1162/nol_a_00078.

    Abstract

    Prediction-based theories of language comprehension assume that listeners predict both the meaning and phonological form of likely upcoming words. In alleged event-related potential (ERP) demonstrations of phonological prediction, prediction-mismatching words elicit a phonological mismatch negativity (PMN), a frontocentral negativity that precedes the centroparietal N400 component. However, classification and replicability of the PMN has proven controversial, with ongoing debate on whether the PMN is a distinct component or merely an early part of the N400. In this electroencephalography (EEG) study, we therefore attempted to replicate the PMN effect and its separability from the N400, using a participant sample size (N = 48) that was more than double that of previous studies. Participants listened to sentences containing either a predictable word or an unpredictable word with/without phonological overlap with the predictable word. Preregistered analyses revealed a widely distributed negative-going ERP in response to unpredictable words in both the early (150–250 ms) and the N400 (300–500 ms) time windows. Bayes factor analysis yielded moderate evidence against a different scalp distribution of the effects in the two time windows. Although our findings do not speak against phonological prediction during sentence comprehension, they do speak against the PMN effect specifically as a marker of phonological prediction mismatch. Instead of an PMN effect, our results demonstrate the early onset of the auditory N400 effect associated with unpredictable words. Our failure to replicate further highlights the risk associated with commonly employed data-contingent analyses (e.g., analyses involving time windows or electrodes that were selected based on visual inspection) and small sample sizes in the cognitive neuroscience of language.
  • Preisig, B., & Hervais-Adelman, A. (2022). The predictive value of individual electric field modeling for transcranial alternating current stimulation induced brain modulation. Frontiers in Cellular Neuroscience, 16: 818703. doi:10.3389/fncel.2022.818703.

    Abstract

    There is considerable individual variability in the reported effectiveness of non-invasive brain stimulation. This variability has often been ascribed to differences in the neuroanatomy and resulting differences in the induced electric field inside the brain. In this study, we addressed the question whether individual differences in the induced electric field can predict the neurophysiological and behavioral consequences of gamma band tACS. In a within-subject experiment, bi-hemispheric gamma band tACS and sham stimulation was applied in alternating blocks to the participants’ superior temporal lobe, while task-evoked auditory brain activity was measured with concurrent functional magnetic resonance imaging (fMRI) and a dichotic listening task. Gamma tACS was applied with different interhemispheric phase lags. In a recent study, we could show that anti-phase tACS (180° interhemispheric phase lag), but not in-phase tACS (0° interhemispheric phase lag), selectively modulates interhemispheric brain connectivity. Using a T1 structural image of each participant’s brain, an individual simulation of the induced electric field was computed. From these simulations, we derived two predictor variables: maximal strength (average of the 10,000 voxels with largest electric field values) and precision of the electric field (spatial correlation between the electric field and the task evoked brain activity during sham stimulation). We found considerable variability in the individual strength and precision of the electric fields. Importantly, the strength of the electric field over the right hemisphere predicted individual differences of tACS induced brain connectivity changes. Moreover, we found in both hemispheres a statistical trend for the effect of electric field strength on tACS induced BOLD signal changes. In contrast, the precision of the electric field did not predict any neurophysiological measure. Further, neither strength, nor precision predicted interhemispheric integration. In conclusion, we found evidence for the dose-response relationship between individual differences in electric fields and tACS induced activity and connectivity changes in concurrent fMRI. However, the fact that this relationship was stronger in the right hemisphere suggests that the relationship between the electric field parameters, neurophysiology, and behavior may be more complex for bi-hemispheric tACS.
  • Preisig, B., Riecke, L., & Hervais-Adelman, A. (2022). Speech sound categorization: The contribution of non-auditory and auditory cortical regions. NeuroImage, 258: 119375. doi:10.1016/j.neuroimage.2022.119375.

    Abstract

    Which processes in the human brain lead to the categorical perception of speech sounds? Investigation of this question is hampered by the fact that categorical speech perception is normally confounded by acoustic differences in the stimulus. By using ambiguous sounds, however, it is possible to dissociate acoustic from perceptual stimulus representations. Twenty-seven normally hearing individuals took part in an fMRI study in which they were presented with an ambiguous syllable (intermediate between /da/ and /ga/) in one ear and with disambiguating acoustic feature (third formant, F3) in the other ear. Multi-voxel pattern searchlight analysis was used to identify brain areas that consistently differentiated between response patterns associated with different syllable reports. By comparing responses to different stimuli with identical syllable reports and identical stimuli with different syllable reports, we disambiguated whether these regions primarily differentiated the acoustics of the stimuli or the syllable report. We found that BOLD activity patterns in left perisylvian regions (STG, SMG), left inferior frontal regions (vMC, IFG, AI), left supplementary motor cortex (SMA/pre-SMA), and right motor and somatosensory regions (M1/S1) represent listeners’ syllable report irrespective of stimulus acoustics. Most of these regions are outside of what is traditionally regarded as auditory or phonological processing areas. Our results indicate that the process of speech sound categorization implicates decision-making mechanisms and auditory-motor transformations.

    Additional information

    figures and table
  • Slivac, K. (2022). The enlanguaged brain: Cognitive and neural mechanisms of linguistic influence on perception. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Udden, J., Hulten, A., Schoffelen, J.-M., Lam, N. H. L., Harbusch, K., Van den Bosch, A., Kempen, G., Petersson, K. M., & Hagoort, P. (2022). Supramodal sentence processing in the human brain: fMRI evidence for the influence of syntactic complexity in more than 200 participants. Neurobiology of Language, 3(4), 575-598. doi:10.1162/nol_a_00076.

    Abstract

    This study investigated two questions. One is: To what degree is sentence processing beyond single words independent of the input modality (speech vs. reading)? The second question is: Which parts of the network recruited by both modalities is sensitive to syntactic complexity? These questions were investigated by having more than 200 participants read or listen to well-formed sentences or series of unconnected words. A largely left-hemisphere frontotemporoparietal network was found to be supramodal in nature, i.e., independent of input modality. In addition, the left inferior frontal gyrus (LIFG) and the left posterior middle temporal gyrus (LpMTG) were most clearly associated with left-branching complexity. The left anterior temporal lobe (LaTL) showed the greatest sensitivity to sentences that differed in right-branching complexity. Moreover, activity in LIFG and LpMTG increased from sentence onset to end, in parallel with an increase of the left-branching complexity. While LIFG, bilateral anterior temporal lobe, posterior MTG, and left inferior parietal lobe (LIPL) all contribute to the supramodal unification processes, the results suggest that these regions differ in their respective contributions to syntactic complexity related processing. The consequences of these findings for neurobiological models of language processing are discussed.

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    supporting information
  • Verdonschot, R. G., Phu'o'ng, H. T. L., & Tamaoka, K. (2022). Phonological encoding in Vietnamese: An experimental investigation. Quarterly Journal of Experimental Psychology, 75(7), 1355-1366. doi:10.1177/17470218211053244.

    Abstract

    In English, Dutch, and other Germanic languages the initial phonological unit used in word production has been shown to be the phoneme; conversely, others have revealed that in Chinese this is the atonal syllable and in Japanese the mora. The current paper is, to our knowledge, the first to report chronometric data on Vietnamese phonological encoding. Vietnamese, a tonal language, is of interest as, despite its Austroasiatic roots, it has clear similarities with Chinese through extended contact over a prolonged period. Four experiments (i.e., masked priming, phonological Stroop, picture naming with written distractors, picture naming with auditory distractors) have been conducted to investigate Vietnamese phonological encoding. Results show that in all four experiments both onset effects as well as whole syllable effects emerge. This indicates that the fundamental phonological encoding unit during Vietnamese language production is the phoneme despite its apparent similarities to Chinese. This result might have emerged due to tone assignment being a qualitatively different process in Vietnamese compared to Chinese.
  • Vernes, S. C., Devanna, P., Hörpel, S. G., Alvarez van Tussenbroek, I., Firzlaff, U., Hagoort, P., Hiller, M., Hoeksema, N., Hughes, G. M., Lavrichenko, K., Mengede, J., Morales, A. E., & Wiesmann, M. (2022). The pale spear‐nosed bat: A neuromolecular and transgenic model for vocal learning. Annals of the New York Academy of Sciences, 1517, 125-142. doi:10.1111/nyas.14884.

    Abstract

    Vocal learning, the ability to produce modified vocalizations via learning from acoustic signals, is a key trait in the evolution of speech. While extensively studied in songbirds, mammalian models for vocal learning are rare. Bats present a promising study system given their gregarious natures, small size, and the ability of some species to be maintained in captive colonies. We utilize the pale spear-nosed bat (Phyllostomus discolor) and report advances in establishing this species as a tractable model for understanding vocal learning. We have taken an interdisciplinary approach, aiming to provide an integrated understanding across genomics (Part I), neurobiology (Part II), and transgenics (Part III). In Part I, we generated new, high-quality genome annotations of coding genes and noncoding microRNAs to facilitate functional and evolutionary studies. In Part II, we traced connections between auditory-related brain regions and reported neuroimaging to explore the structure of the brain and gene expression patterns to highlight brain regions. In Part III, we created the first successful transgenic bats by manipulating the expression of FoxP2, a speech-related gene. These interdisciplinary approaches are facilitating a mechanistic and evolutionary understanding of mammalian vocal learning and can also contribute to other areas of investigation that utilize P. discolor or bats as study species.

    Additional information

    supplementary materials
  • Wanner-Kawahara, J., Yoshihara, M., Lupker, S. J., Verdonschot, R. G., & Nakayama, M. (2022). Morphological priming effects in L2 English verbs for Japanese-English bilinguals. Frontiers in Psychology, 13: 742965. doi:10.3389/fpsyg.2022.742965.

    Abstract

    For native (L1) English readers, masked presentations of past-tense verb primes (e.g., fell and looked) produce faster lexical decision latencies to their present-tense targets (e.g., FALL and LOOK) than orthographically related (e.g., fill and loose) or unrelated (e.g., master and bank) primes. This facilitation observed with morphologically related prime-target pairs (morphological priming) is generally taken as evidence for strong connections based on morphological relationships in the L1 lexicon. It is unclear, however, if similar, morphologically based, connections develop in non-native (L2) lexicons. Several earlier studies with L2 English readers have reported mixed results. The present experiments examine whether past-tense verb primes (both regular and irregular verbs) significantly facilitate target lexical decisions for Japanese-English bilinguals beyond any facilitation provided by prime-target orthographic similarity. Overall, past-tense verb primes facilitated lexical decisions to their present-tense targets relative to both orthographically related and unrelated primes. Replicating previous masked priming experiments with L2 readers, orthographically related primes also facilitated target recognition relative to unrelated primes, confirming that orthographic similarity facilitates L2 target recognition. The additional facilitation from past-tense verb primes beyond that provided by orthographic primes suggests that, in the L2 English lexicon, connections based on morphological relationships develop in a way that is similar to how they develop in the L1 English lexicon even though the connections and processing of lower level, lexical/orthographic information may differ. Further analyses involving L2 proficiency revealed that as L2 proficiency increased, orthographic facilitation was reduced, indicating that there is a decrease in the fuzziness in orthographic representations in the L2 lexicon with increased proficiency.

    Additional information

    supplementary material
  • Wilms, V., Drijvers, L., & Brouwer, S. (2022). The Effects of Iconic Gestures and Babble Language on Word Intelligibility in Sentence Context. Journal of Speech, Language, and Hearing Research, 65, 1822-1838. doi:10.1044/2022\_JSLHR-21-00387.

    Abstract

    Purpose:This study investigated to what extent iconic co-speech gestures helpword intelligibility in sentence context in two different linguistic maskers (nativevs. foreign). It was hypothesized that sentence recognition improves with thepresence of iconic co-speech gestures and with foreign compared to nativebabble.Method:Thirty-two native Dutch participants performed a Dutch word recogni-tion task in context in which they were presented with videos in which anactress uttered short Dutch sentences (e.g.,Ze begint te openen,“She starts toopen”). Participants were presented with a total of six audiovisual conditions: nobackground noise (i.e., clear condition) without gesture, no background noise withgesture, French babble without gesture, French babble with gesture, Dutch bab-ble without gesture, and Dutch babble with gesture; and they were asked to typedown what was said by the Dutch actress. The accurate identification of theaction verbs at the end of the target sentences was measured.Results:The results demonstrated that performance on the task was better inthe gesture compared to the nongesture conditions (i.e., gesture enhancementeffect). In addition, performance was better in French babble than in Dutchbabble.Conclusions:Listeners benefit from iconic co-speech gestures during commu-nication and from foreign background speech compared to native. Theseinsights into multimodal communication may be valuable to everyone whoengages in multimodal communication and especially to a public who oftenworks in public places where competing speech is present in the background.
  • Yang, J. (2022). Discovering the units in language cognition: From empirical evidence to a computational model. PhD Thesis, Radboud University Nijmegen, Nijmegen.
  • Yang, J., Van den Bosch, A., & Frank, S. L. (2022). Unsupervised text segmentation predicts eye fixations during reading. Frontiers in Artificial Intelligence, 5: 731615. doi:10.3389/frai.2022.731615.

    Abstract

    Words typically form the basis of psycholinguistic and computational linguistic studies about sentence processing. However, recent evidence shows the basic units during reading, i.e., the items in the mental lexicon, are not always words, but could also be sub-word and supra-word units. To recognize these units, human readers require a cognitive mechanism to learn and detect them. In this paper, we assume eye fixations during reading reveal the locations of the cognitive units, and that the cognitive units are analogous with the text units discovered by unsupervised segmentation models. We predict eye fixations by model-segmented units on both English and Dutch text. The results show the model-segmented units predict eye fixations better than word units. This finding suggests that the predictive performance of model-segmented units indicates their plausibility as cognitive units. The Less-is-Better (LiB) model, which finds the units that minimize both long-term and working memory load, offers advantages both in terms of prediction score and efficiency among alternative models. Our results also suggest that modeling the least-effort principle for the management of long-term and working memory can lead to inferring cognitive units. Overall, the study supports the theory that the mental lexicon stores not only words but also smaller and larger units, suggests that fixation locations during reading depend on these units, and shows that unsupervised segmentation models can discover these units.
  • Zora, H., Gussenhoven, C., Tremblay, A., & Liu, F. (2022). Editorial: Crosstalk between intonation and lexical tones: Linguistic, cognitive and neuroscience perspectives. Frontiers in Psychology, 13: 1101499. doi:10.3389/fpsyg.2022.1101499.

    Abstract

    The interplay between categorical and continuous aspects of the speech signal remains central and yet controversial in the fields of phonetics and phonology. The division between phonological abstractions and phonetic variations has been particularly relevant to the unraveling of diverse communicative functions of pitch in the domain of prosody. Pitch influences vocal communication in two major but fundamentally different ways, and lexical and intonational tones exquisitely capture these functions. Lexical tone contrasts convey lexical meanings as well as derivational meanings at the word level and are grammatically encoded as discrete structures. Intonational tones, on the other hand, signal post-lexical meanings at the phrasal level and typically allow gradient pragmatic variations. Since categorical and gradient uses of pitch are ubiquitous and closely intertwined in their physiological and psychological processes, further research is warranted for a more detailed understanding of their structural and functional characterisations. This Research Topic addresses this matter from a wide range of perspectives, including first and second language acquisition, speech production and perception, structural and functional diversity, and working with distinct languages and experimental measures. In the following, we provide a short overview of the contributions submitted to this topic

    Additional information

    also published as book chapter (2023)
  • 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. (2007). The memory, unification, and control (MUC) model of language. In T. Sakamoto (Ed.), Communicating skills of intention (pp. 259-291). Tokyo: Hituzi Syobo.
  • Hagoort, P. (2007). The memory, unification, and control (MUC) model of language. In A. S. Meyer, L. Wheeldon, & A. Krott (Eds.), Automaticity and control in language processing (pp. 243-270). Hove: Psychology Press.
  • Hagoort, P., & Van Berkum, J. J. A. (2007). Beyond the sentence given. Philosophical Transactions of the Royal Society. Series B: Biological Sciences, 362, 801-811.

    Abstract

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

    Abstract

    In an ERP experiment we investigated how the recruitment and integration of world knowledge information relate to the integration of information within a current discourse context. Participants were presented with short discourse contexts which were followed by a sentence that contained a critical word that was correct or incorrect based on general world knowledge and the supporting discourse context, or was more or less acceptable based on the combination of general world knowledge and the specific local discourse context. Relative to the critical word in the correct world knowledge sentences following a neutral discourse, all other critical words elicited an N400 effect that began at about 300 ms after word onset. However, the magnitude of the N400 effect varied in a way that suggests an interaction between world knowledge and discourse context. The results indicate that both world knowledge and discourse context have an effect on sentence interpretation, but neither overrides the other.
  • 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., 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
  • 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.
  • 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., & 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.
  • 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.
  • 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.
  • 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., & Kelly, S. D. (2007). Gesture, language, and brain. Brain and Language, 101(3), 181-185. doi:10.1016/j.bandl.2007.03.006.
  • 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 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
  • 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.
  • 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., 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.
  • 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.

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