Publications

Displaying 1 - 94 of 94
  • 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.
  • 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)
  • 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.

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    supporting information
  • 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.
  • 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.

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    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.
  • 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., 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
  • 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.

    Additional information

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

    Abstract

    A number of recent studies have hypothesized that monitoring in speech production may occur via domain-general mechanisms responsible for the detection of response conflict. Outside of language, two ERP components have consistently been elicited in conflict-inducing tasks (e.g., the flanker task): the stimulus-locked N2 on correct trials, and the response-locked error-related negativity (ERN). The present investigation used these electrophysiological markers to test whether a common response conflict monitor is responsible for monitoring in speech and non-speech tasks. Electroencephalography (EEG) was recorded while participants performed a tongue twister (TT) task and a manual version of the flanker task. In the TT task, people rapidly read sequences of four nonwords arranged in TT and non-TT patterns three times. In the flanker task, people responded with a left/right button press to a center-facing arrow, and conflict was manipulated by the congruency of the flanking arrows. Behavioral results showed typical effects of both tasks, with increased error rates and slower speech onset times for TT relative to non-TT trials and for incongruent relative to congruent flanker trials. In the flanker task, stimulus-locked EEG analyses replicated previous results, with a larger N2 for incongruent relative to congruent trials, and a response-locked ERN. In the TT task, stimulus-locked analyses revealed broad, frontally-distributed differences beginning around 50 ms and lasting until just before speech initiation, with TT trials more negative than non-TT trials; response-locked analyses revealed an ERN. Correlation across these measures showed some correlations within a task, but little evidence of systematic cross-task correlation. Although the present results do not speak against conflict signals from the production system serving as cues to self-monitoring, they are not consistent with signatures of response conflict being mediated by a single, domain-general conflict monitor
  • Araújo, S., Faísca, L., Bramão, I., Petersson, K. M., & Reis, A. (2014). Lexical and phonological processes in dyslexic readers: Evidences from a visual lexical decision task. Dyslexia, 20, 38-53. doi:10.1002/dys.1461.

    Abstract

    The aim of the present study was to investigate whether reading failure in the context of an orthography of intermediate consistency is linked to inefficient use of the lexical orthographic reading procedure. The performance of typically developing and dyslexic Portuguese-speaking children was examined in a lexical decision task, where the stimulus lexicality, word frequency and length were manipulated. Both lexicality and length effects were larger in the dyslexic group than in controls, although the interaction between group and frequency disappeared when the data were transformed to control for general performance factors. Children with dyslexia were influenced in lexical decision making by the stimulus length of words and pseudowords, whereas age-matched controls were influenced by the length of pseudowords only. These findings suggest that non-impaired readers rely mainly on lexical orthographic information, but children with dyslexia preferentially use the phonological decoding procedure—albeit poorly—most likely because they struggle to process orthographic inputs as a whole such as controls do. Accordingly, dyslexic children showed significantly poorer performance than controls for all types of stimuli, including words that could be considered over-learned, such as high-frequency words. This suggests that their orthographic lexical entries are less established in the orthographic lexicon
  • Basnakova, J., Weber, K., Petersson, K. M., Van Berkum, J. J. A., & Hagoort, P. (2014). Beyond the language given: The neural correlates of inferring speaker meaning. Cerebral Cortex, 24(10), 2572-2578. doi:10.1093/cercor/bht112.

    Abstract

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

    Abstract

    Heschl's gyrus (HG) is a core region of the auditory cortex whose morphology is highly variable across individuals. This variability has been linked to sound perception ability in both speech and music domains. Previous studies show that variations in morphological features of HG, such as cortical surface area and thickness, are heritable. To identify genetic variants that affect HG morphology, we conducted a genome-wide association scan (GWAS) meta-analysis in 3054 healthy individuals using HG surface area and thickness as quantitative traits. None of the single nucleotide polymorphisms (SNPs) showed association P values that would survive correction for multiple testing over the genome. The most significant association was found between right HG area and SNP rs72932726 close to gene DCBLD2 (3q12.1; P=2.77x10(-7)). This SNP was also associated with other regions involved in speech processing. The SNP rs333332 within gene KALRN (3q21.2; P=2.27x10(-6)) and rs143000161 near gene COBLL1 (2q24.3; P=2.40x10(-6)) were associated with the area and thickness of left HG, respectively. Both genes are involved in the development of the nervous system. The SNP rs7062395 close to the X-linked deafness gene POU3F4 was associated with right HG thickness (Xq21.1; P=2.38x10(-6)). This is the first molecular genetic analysis of variability in HG morphology
  • Capilla, A., Schoffelen, J.-M., Paterson, G., Thut, G., & Gross, J. (2014). Dissociated α-band modulations in the dorsal and ventral visual pathways in visuospatial attention and perception. Cerebral Cortex., 24(2), 550-561. doi:10.1093/cercor/bhs343.

    Abstract

    Modulations of occipito-parietal α-band (8–14 Hz) power that are opposite in direction (α-enhancement vs. α-suppression) and origin of generation (ipsilateral vs. contralateral to the locus of attention) are a robust correlate of anticipatory visuospatial attention. Yet, the neural generators of these α-band modulations, their interdependence across homotopic areas, and their respective contribution to subsequent perception remain unclear. To shed light on these questions, we employed magnetoencephalography, while human volunteers performed a spatially cued detection task. Replicating previous findings, we found α-power enhancement ipsilateral to the attended hemifield and contralateral α-suppression over occipitoparietal sensors. Source localization (beamforming) analysis showed that α-enhancement and suppression were generated in 2 distinct brain regions, located in the dorsal and ventral visual streams, respectively. Moreover, α-enhancement and suppression showed different dynamics and contribution to perception. In contrast to the initial and transient dorsal α-enhancement, α-suppression in ventro-lateral occipital cortex was sustained and influenced subsequent target detection. This anticipatory biasing of ventrolateral extrastriate α-activity probably reflects increased receptivity in the brain region specialized in processing upcoming target features. Our results add to current models on the role of α-oscillations in attention orienting by showing that α-enhancement and suppression can be dissociated in time, space, and perceptual relevance.

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    Capilla_Suppl_Data.pdf
  • Chu, M., Meyer, A. S., Foulkes, L., & Kita, S. (2014). Individual differences in frequency and saliency of speech-accompanying gestures: The role of cognitive abilities and empathy. Journal of Experimental Psychology: General, 143, 694-709. doi:10.1037/a0033861.

    Abstract

    The present study concerns individual differences in gesture production. We used correlational and multiple regression analyses to examine the relationship between individuals’ cognitive abilities and empathy levels and their gesture frequency and saliency. We chose predictor variables according to experimental evidence of the functions of gesture in speech production and communication. We examined 3 types of gestures: representational gestures, conduit gestures, and palm-revealing gestures. Higher frequency of representational gestures was related to poorer visual and spatial working memory, spatial transformation ability, and conceptualization ability; higher frequency of conduit gestures was related to poorer visual working memory, conceptualization ability, and higher levels of empathy; and higher frequency of palm-revealing gestures was related to higher levels of empathy. The saliency of all gestures was positively related to level of empathy. These results demonstrate that cognitive abilities and empathy levels are related to individual differences in gesture frequency and saliency
  • Chu, M., & Hagoort, P. (2014). Synchronization of speech and gesture: Evidence for interaction in action. Journal of Experimental Psychology: General, 143(4), 1726-1741. doi:10.1037/a0036281.

    Abstract

    Language and action systems are highly interlinked. A critical piece of evidence is that speech and its accompanying gestures are tightly synchronized. Five experiments were conducted to test 2 hypotheses about the synchronization of speech and gesture. According to the interactive view, there is continuous information exchange between the gesture and speech systems, during both their planning and execution phases. According to the ballistic view, information exchange occurs only during the planning phases of gesture and speech, but the 2 systems become independent once their execution has been initiated. In all experiments, participants were required to point to and/or name a light that had just lit up. Virtual reality and motion tracking technologies were used to disrupt their gesture or speech execution. Participants delayed their speech onset when their gesture was disrupted. They did so even when their gesture was disrupted at its late phase and even when they received only the kinesthetic feedback of their gesture. Also, participants prolonged their gestures when their speech was disrupted. These findings support the interactive view and add new constraints on models of speech and gesture production
  • Cristia, A., Minagawa-Kawai, Y., Egorova, N., Gervain, J., Filippin, L., Cabrol, D., & Dupoux, E. (2014). Neural correlates of infant accent discrimination: An fNIRS study. Developmental Science, 17(4), 628-635. doi:10.1111/desc.12160.

    Abstract

    The present study investigated the neural correlates of infant discrimination of very similar linguistic varieties (Quebecois and Parisian French) using functional Near InfraRed Spectroscopy. In line with previous behavioral and electrophysiological data, there was no evidence that 3-month-olds discriminated the two regional accents, whereas 5-month-olds did, with the locus of discrimination in left anterior perisylvian regions. These neuroimaging results suggest that a developing language network relying crucially on left perisylvian cortices sustains infants' discrimination of similar linguistic varieties within this early period of infancy.

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  • Cristia, A., Seidl, A., Junge, C., Soderstrom, M., & Hagoort, P. (2014). Predicting individual variation in language from infant speech perception measures. Child development, 85(4), 1330-1345. doi:10.1111/cdev.12193.

    Abstract

    There are increasing reports that individual variation in behavioral and neurophysiological measures of infant speech processing predicts later language outcomes, and specifically concurrent or subsequent vocabulary size. If such findings are held up under scrutiny, they could both illuminate theoretical models of language development and contribute to the prediction of communicative disorders. A qualitative, systematic review of this emergent literature illustrated the variety of approaches that have been used and highlighted some conceptual problems regarding the measurements. A quantitative analysis of the same data established that the bivariate relation was significant, with correlations of similar strength to those found for well-established nonlinguistic predictors of language. Further exploration of infant speech perception predictors, particularly from a methodological perspective, is recommended.
  • Cristia, A., & Seidl, A. (2014). The hyperarticulation hypothesis of infant-directed speech. Journal of Child Language, 41(4), 913-934. doi:10.1017/S0305000912000669.

    Abstract

    Typically, the point vowels [i,ɑ,u] are acoustically more peripheral in infant-directed speech (IDS) compared to adult-directed speech (ADS). If caregivers seek to highlight lexically relevant contrasts in IDS, then two sounds that are contrastive should become more distinct, whereas two sounds that are surface realizations of the same underlying sound category should not. To test this prediction, vowels that are phonemically contrastive ([i-ɪ] and [eɪ-ε]), vowels that map onto the same underlying category ([æ- ] and [ε- ]), and the point vowels [i,ɑ,u] were elicited in IDS and ADS by American English mothers of two age groups of infants (four- and eleven-month-olds). As in other work, point vowels were produced in more peripheral positions in IDS compared to ADS. However, there was little evidence of hyperarticulation per se (e.g. [i-ɪ] was hypoarticulated). We suggest that across-the-board lexically based hyperarticulation is not a necessary feature of IDS.

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    CORRIGENDUM
  • Dautriche, I., Cristia, A., Brusini, P., Yuan, S., Fisher, C., & Christophe, A. (2014). Toddlers default to canonical surface-to-meaning mapping when learning verbs. Child Development, 85(3), 1168-1180. doi:10.1111/cdev.12183.

    Abstract

    This work was supported by grants from the French Agence Nationale de la Recherche (ANR-2010-BLAN-1901) and from French Fondation de France to Anne Christophe, from the National Institute of Child Health and Human Development (HD054448) to Cynthia Fisher, Fondation Fyssen and Ecole de Neurosciences de Paris to Alex Cristia, and a PhD fellowship from the Direction Générale de l'Armement (DGA, France) supported by the PhD program FdV (Frontières du Vivant) to Isabelle Dautriche. We thank Isabelle Brunet for the recruitment, Michel Dutat for the technical support, and Hernan Anllo for his puppet mastery skill. We are grateful to the families that participated in this study. We also thank two anonymous reviewers for their comments on an earlier draft of this manuscript.
  • Dolscheid, S., Hunnius, S., Casasanto, D., & Majid, A. (2014). Prelinguistic infants are sensitive to space-pitch associations found across cultures. Psychological Science, 25(6), 1256-1261. doi:10.1177/0956797614528521.

    Abstract

    People often talk about musical pitch using spatial metaphors. In English, for instance, pitches can be “high” or “low” (i.e., height-pitch association), whereas in other languages, pitches are described as “thin” or “thick” (i.e., thickness-pitch association). According to results from psychophysical studies, metaphors in language can shape people’s nonlinguistic space-pitch representations. But does language establish mappings between space and pitch in the first place, or does it only modify preexisting associations? To find out, we tested 4-month-old Dutch infants’ sensitivity to height-pitch and thickness-pitch mappings using a preferential-looking paradigm. The infants looked significantly longer at cross-modally congruent stimuli for both space-pitch mappings, which indicates that infants are sensitive to these associations before language acquisition. The early presence of space-pitch mappings means that these associations do not originate from language. Instead, language builds on preexisting mappings, changing them gradually via competitive associative learning. Space-pitch mappings that are language-specific in adults develop from mappings that may be universal in infants.
  • Folia, V., & Petersson, K. M. (2014). Implicit structured sequence learning: An fMRI study of the structural mere-exposure effect. Frontiers in Psychology, 5: 41. doi:10.3389/fpsyg.2014.00041.

    Abstract

    In this event-related FMRI study we investigated the effect of five days of implicit acquisition on preference classification by means of an artificial grammar learning (AGL) paradigm based on the structural mere-exposure effect and preference classification using a simple right-linear unification grammar. This allowed us to investigate implicit AGL in a proper learning design by including baseline measurements prior to grammar exposure. After 5 days of implicit acquisition, the FMRI results showed activations in a network of brain regions including the inferior frontal (centered on BA 44/45) and the medial prefrontal regions (centered on BA 8/32). Importantly, and central to this study, the inclusion of a naive preference FMRI baseline measurement allowed us to conclude that these FMRI findings were the intrinsic outcomes of the learning process itself and not a reflection of a preexisting functionality recruited during classification, independent of acquisition. Support for the implicit nature of the knowledge utilized during preference classification on day 5 come from the fact that the basal ganglia, associated with implicit procedural learning, were activated during classification, while the medial temporal lobe system, associated with explicit declarative memory, was consistently deactivated. Thus, preference classification in combination with structural mere-exposure can be used to investigate structural sequence processing (syntax) in unsupervised AGL paradigms with proper learning designs.
  • De Grauwe, S., Willems, R. M., Rüschemeyer, S.-A., Lemhöfer, K., & Schriefers, H. (2014). Embodied language in first- and second-language speakers: Neural correlates of processing motor verbs. Neuropsychologia, 56, 334-349. doi:10.1016/j.neuropsychologia.2014.02.003.

    Abstract

    The involvement of neural motor and sensory systems in the processing of language has so far mainly been studied in native (L1) speakers. In an fMRI experiment, we investigated whether non-native (L2) semantic representations are rich enough to allow for activation in motor and somatosensory brain areas. German learners of Dutch and a control group of Dutch native speakers made lexical decisions about visually presented Dutch motor and non-motor verbs. Region-of-interest (ROI) and whole-brain analyses indicated that L2 speakers, like L1 speakers, showed significantly increased activation for simple motor compared to non-motor verbs in motor and somatosensory regions. This effect was not restricted to Dutch-German cognate verbs, but was also present for non-cognate verbs. These results indicate that L2 semantic representations are rich enough for motor-related activations to develop in motor and somatosensory areas.
  • De Grauwe, S., Lemhöfer, K., Willems, R. M., & Schriefers, H. (2014). L2 speakers decompose morphologically complex verbs: fMRI evidence from priming of transparent derived verbs. Frontiers in Human Neuroscience, 8: 802. doi:10.3389/fnhum.2014.00802.

    Abstract

    In this functional magnetic resonance imaging (fMRI) long-lag priming study, we investigated the processing of Dutch semantically transparent, derived prefix verbs. In such words, the meaning of the word as a whole can be deduced from the meanings of its parts, e.g., wegleggen “put aside.” Many behavioral and some fMRI studies suggest that native (L1) speakers decompose transparent derived words. The brain region usually implicated in morphological decomposition is the left inferior frontal gyrus (LIFG). In non-native (L2) speakers, the processing of transparent derived words has hardly been investigated, especially in fMRI studies, and results are contradictory: some studies find more reliance on holistic (i.e., non-decompositional) processing by L2 speakers; some find no difference between L1 and L2 speakers. In this study, we wanted to find out whether Dutch transparent derived prefix verbs are decomposed or processed holistically by German L2 speakers of Dutch. Half of the derived verbs (e.g., omvallen “fall down”) were preceded by their stem (e.g., vallen “fall”) with a lag of 4–6 words (“primed”); the other half (e.g., inslapen “fall asleep”) were not (“unprimed”). L1 and L2 speakers of Dutch made lexical decisions on these visually presented verbs. Both region of interest analyses and whole-brain analyses showed that there was a significant repetition suppression effect for primed compared to unprimed derived verbs in the LIFG. This was true both for the analyses over L2 speakers only and for the analyses over the two language groups together. The latter did not reveal any interaction with language group (L1 vs. L2) in the LIFG. Thus, L2 speakers show a clear priming effect in the LIFG, an area that has been associated with morphological decomposition. Our findings are consistent with the idea that L2 speakers engage in decomposition of transparent derived verbs rather than processing them holistically

    Additional information

    Data Sheet 1.docx
  • Guadalupe, T., Willems, R. M., Zwiers, M., Arias Vasquez, A., Hoogman, M., Hagoort, P., Fernández, G., Buitelaar, J., Franke, B., Fisher, S. E., & Francks, C. (2014). Differences in cerebral cortical anatomy of left- and right-handers. Frontiers in Psychology, 5: 261. doi:10.3389/fpsyg.2014.00261.

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

    A hallmark of human language is that we combine lexical building blocks retrieved from memory in endless new ways. This combinatorial aspect of language is referred to as unification. Here we focus on the neurobiological infrastructure for syntactic and semantic unification. Unification is characterized by a high-speed temporal profile including both prediction and integration of retrieved lexical elements. A meta-analysis of numerous neuroimaging studies reveals a clear dorsal/ventral gradient in both left inferior frontal cortex and left posterior temporal cortex, with dorsal foci for syntactic processing and ventral foci for semantic processing. In addition to core areas for unification, further networks need to be recruited to realize language-driven communication to its full extent. One example is the theory of mind network, which allows listeners and readers to infer the intended message (speaker meaning) from the coded meaning of the linguistic utterance. This indicates that sensorimotor simulation cannot handle all of language processing.
  • Holler, J., Schubotz, L., Kelly, S., Hagoort, P., Schuetze, M., & Ozyurek, A. (2014). Social eye gaze modulates processing of speech and co-speech gesture. Cognition, 133, 692-697. doi:10.1016/j.cognition.2014.08.008.

    Abstract

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

    Abstract

    Most words that infants hear occur within fluent speech. To compile a vocabulary, infants therefore need to segment words from speech contexts. This study is the first to investigate whether infants (here: 10-month-olds) can recognize words when both initial exposure and test presentation are in continuous speech. Electrophysiological evidence attests that this indeed occurs: An increased extended negativity (word recognition effect) appears for familiarized target words relative to control words. This response proved constant at the individual level: Only infants who showed this negativity at test had shown such a response, within six repetitions after first occurrence, during familiarization.
  • Keller, K. L., Fritz, R. S., Zoubek, C. M., Kennedy, E. H., Cronin, K. A., Rothwell, E. S., & Serfass, T. L. (2014). Effects of transport on fecal glucocorticoid levels in captive-bred cotton-top tamarins (Saguinus oedipus). Journal of the Pennsylvania Academy of Science, 88(2), 84-88.

    Abstract

    The relocation of animals can induce stress when animals are placed in novel environmental conditions. The movement of captive animals among facilities is common, especially for non-human primates used in research. The stress response begins with the activation of the hypothalamic-pituitary-adrenal (HPA) axis which results in the release of glucocorticoid hormones (GC), which at chronic levels could lead to deleterious physiological effects. There is a substantial body of data concerning GC levels affecting reproduction, and rank and aggression in primates. However, the effect of transport has received much less attention. Fecal samples from eight (four male and four female) captive-bred cotton-top tamarins (Saguinus oedipus) were collected at four different time points (two pre-transport and two post-transport). The fecal samples were analyzed using an immunoassay to determine GC levels. A repeated measures analysis of variance (ANOVA) demonstrated that GC levels differed among transport times (p = 0.009), but not between sexes (p = 0.963). Five of the eight tamarins exhibited an increase in GC levels after transport. Seven of the eight tamarins exhibited a decrease in GC levels from three to six days post-transport to three weeks post-transport. Most values returned to pre-transport levels after three weeks. The results indicate that these tamarins experienced elevated GC levels following transport, but these increases were of short duration. This outcome would suggest that the negative effects of elevated GC levels were also of short duration.
  • Kunert, R., & Scheepers, C. (2014). Speed and accuracy of dyslexic versus typical word recognition: An eye-movement investigation. Frontiers in Psychology, 5: 1129. doi:10.3389/fpsyg.2014.01129.

    Abstract

    Developmental dyslexia is often characterized by a dual deficit in both word recognition accuracy and general processing speed. While previous research into dyslexic word recognition may have suffered from speed-accuracy trade-off, the present study employed a novel eye-tracking task that is less prone to such confounds. Participants (10 dyslexics and 12 controls) were asked to look at real word stimuli, and to ignore simultaneously presented non-word stimuli, while their eye-movements were recorded. Improvements in word recognition accuracy over time were modeled in terms of a continuous non-linear function. The words' rhyme consistency and the non-words' lexicality (unpronounceable, pronounceable, pseudohomophone) were manipulated within-subjects. Speed-related measures derived from the model fits confirmed generally slower processing in dyslexics, and showed a rhyme consistency effect in both dyslexics and controls. In terms of overall error rate, dyslexics (but not controls) performed less accurately on rhyme-inconsistent words, suggesting a representational deficit for such words in dyslexics. Interestingly, neither group showed a pseudohomophone effect in speed or accuracy, which might call the task-independent pervasiveness of this effect into question. The present results illustrate the importance of distinguishing between speed- vs. accuracy-related effects for our understanding of dyslexic word recognition

    Additional information

    Kunert_Data Sheet 1.DOCX
  • Lai, V. T., Garrido Rodriguez, G., & Narasimhan, B. (2014). Thinking-for-speaking in early and late bilinguals. Bilingualism: Language and Cognition, 17, 139-152. doi:10.1017/S1366728913000151.

    Abstract

    When speakers describe motion events using different languages, they subsequently classify those events in language-specific ways (Gennari, Sloman, Malt & Fitch, 2002). Here we ask if bilingual speakers flexibly shift their event classification preferences based on the language in which they verbally encode those events. English–Spanish bilinguals and monolingual controls described motion events in either Spanish or English. Subsequently they judged the similarity of the motion events in a triad task. Bilinguals tested in Spanish and Spanish monolinguals were more likely to make similarity judgments based on the path of motion versus bilinguals tested in English and English monolinguals. The effect is modulated in bilinguals by the age of acquisition of the second language. Late bilinguals based their judgments on path more often when Spanish was used to describe the motion events versus English. Early bilinguals had a path preference independent of the language in use. These findings support “thinking-for-speaking” (Slobin, 1996) in late bilinguals.
  • Lartseva, A., Dijkstra, T., Kan, C. C., & Buitelaar, J. K. (2014). Processing of emotion words by patients with Autism Spectrum Disorders: Evidence from reaction times and EEG. Journal of Autism and Developmental Disorders, 44, 2882-2894. doi:10.1007/s10803-014-2149-z.

    Abstract

    This study investigated processing of emotion words in autism spectrum disorders (ASD) using reaction times and event-related potentials (ERP). Adults with (n = 21) and without (n = 20) ASD performed a lexical decision task on emotion and neutral words while their brain activity was recorded. Both groups showed faster responses to emotion words compared to neutral, suggesting intact early processing of emotion in ASD. In the ERPs, the control group showed a typical late positive component (LPC) at 400-600 ms for emotion words compared to neutral, while the ASD group showed no LPC. The between-group difference in LPC amplitude was significant, suggesting that emotion words were processed differently by individuals with ASD, although their behavioral performance was similar to that of typical individuals
  • Levy, J., Hagoort, P., & Démonet, J.-F. (2014). A neuronal gamma oscillatory signature during morphological unification in the left occipitotemporal junction. Human Brain Mapping, 35, 5847-5860. doi:10.1002/hbm.22589.

    Abstract

    Morphology is the aspect of language concerned with the internal structure of words. In the past decades, a large body of masked priming (behavioral and neuroimaging) data has suggested that the visual word recognition system automatically decomposes any morphologically complex word into a stem and its constituent morphemes. Yet the reliance of morphology on other reading processes (e.g., orthography and semantics), as well as its underlying neuronal mechanisms are yet to be determined. In the current magnetoencephalography study, we addressed morphology from the perspective of the unification framework, that is, by applying the Hold/Release paradigm, morphological unification was simulated via the assembly of internal morphemic units into a whole word. Trials representing real words were divided into words with a transparent (true) or a nontransparent (pseudo) morphological relationship. Morphological unification of truly suffixed words was faster and more accurate and additionally enhanced induced oscillations in the narrow gamma band (60–85 Hz, 260–440 ms) in the left posterior occipitotemporal junction. This neural signature could not be explained by a mere automatic lexical processing (i.e., stem perception), but more likely it related to a semantic access step during the morphological unification process. By demonstrating the validity of unification at the morphological level, this study contributes to the vast empirical evidence on unification across other language processes. Furthermore, we point out that morphological unification relies on the retrieval of lexical semantic associations via induced gamma band oscillations in a cerebral hub region for visual word form processing.
  • Lüttjohann, A., Schoffelen, J.-M., & Van Luijtelaar, G. (2014). Termination of ongoing spike-wave discharges investigated by cortico-thalamic network analyses. Neurobiology of Disease, 70, 127-137. doi:10.1016/j.nbd.2014.06.007.

    Abstract

    Purpose While decades of research were devoted to study generation mechanisms of spontaneous spike and wave discharges (SWD), little attention has been paid to network mechanisms associated with the spontaneous termination of SWD. In the current study coupling-dynamics at the onset and termination of SWD were studied in an extended part of the cortico-thalamo-cortical system of freely moving, genetic absence epileptic WAG/Rij rats. Methods Local-field potential recordings of 16 male WAG/Rij rats, equipped with multiple electrodes targeting layer 4 to 6 of the somatosensory-cortex (ctx4, ctx5, ctx6), rostral and caudal reticular thalamic nucleus (rRTN & cRTN), Ventral Postero Medial (VPM), anterior- (ATN) and posterior (Po) thalamic nucleus, were obtained. Six seconds lasting pre-SWD->SWD, SWD->post SWD and control periods were analyzed with time-frequency methods and between-region interactions were quantified with frequencyresolved Granger Causality (GC) analysis. Results Most channel-pairs showed increases in GC lasting from onset to offset of the SWD. While for most thalamo-thalamic pairs a dominant coupling direction was found during the complete SWD, most cortico-thalamic pairs only showed a dominant directional drive (always from cortex to thalamus) during the first 500ms of SWD. Channel-pair ctx4-rRTN showed a longer lasting dominant cortical drive, which stopped 1.5 sec prior to SWD offset. This early decrease in directional coupling was followed by an increase in directional coupling from cRTN to rRTN 1 sec prior to SWD offset. For channel pairs ctx5-Po and ctx6-Po the heightened cortex->thalamus coupling remained until 1.5 sec following SWD offset, while the thalamus->cortex coupling for these pairs stopped at SWD offset. Conclusion The high directional coupling from somatosensory cortex to the thalamus at SWD onset is in good agreement with the idea of a cortical epileptic focus that initiates and entrains other brain structures into seizure activity. The decrease of cortex to rRTN coupling as well as the increased coupling from cRTN to rRTN preceding SWD termination demonstrate that SWD termination is a gradual process that involves both cortico-thalamic as well as intrathalamic processes. The rostral RTN seems to be an important resonator for SWD and relevant for maintenance, while the cRTN might inhibit this oscillation. The somatosensory cortex seems to attempt to reinitiate SWD following its offset via its strong coupling to the posterior thalamus.
  • Magyari, L., Bastiaansen, M. C. M., De Ruiter, J. P., & Levinson, S. C. (2014). Early anticipation lies behind the speed of response in conversation. Journal of Cognitive Neuroscience, 26(11), 2530-2539. doi:10.1162/jocn_a_00673.

    Abstract

    RTs in conversation, with average gaps of 200 msec and often less, beat standard RTs, despite the complexity of response and the lag in speech production (600 msec or more). This can only be achieved by anticipation of timing and content of turns in conversation, about which little is known. Using EEG and an experimental task with conversational stimuli, we show that estimation of turn durations are based on anticipating the way the turn would be completed. We found a neuronal correlate of turn-end anticipation localized in ACC and inferior parietal lobule, namely a beta-frequency desynchronization as early as 1250 msec, before the end of the turn. We suggest that anticipation of the other's utterance leads to accurately timed transitions in everyday conversations.
  • Pacheco, A., Araújo, S., Faísca, L., de Castro, S. L., Petersson, K. M., & Reis, A. (2014). Dyslexia's heterogeneity: Cognitive profiling of Portuguese children with dyslexia. Reading and Writing, 27(9), 1529-1545. doi:10.1007/s11145-014-9504-5.

    Abstract

    Recent studies have emphasized that developmental dyslexia is a multiple-deficit disorder, in contrast to the traditional single-deficit view. In this context, cognitive profiling of children with dyslexia may be a relevant contribution to this unresolved discussion. The aim of this study was to profile 36 Portuguese children with dyslexia from the 2nd to 5th grade. Hierarchical cluster analysis was used to group participants according to their phonological awareness, rapid automatized naming, verbal short-term memory, vocabulary, and nonverbal intelligence abilities. The results suggested a two-cluster solution: a group with poorer performance on phoneme deletion and rapid automatized naming compared with the remaining variables (Cluster 1) and a group characterized by underperforming on the variables most related to phonological processing (phoneme deletion and digit span), but not on rapid automatized naming (Cluster 2). Overall, the results seem more consistent with a hybrid perspective, such as that proposed by Pennington and colleagues (2012), for understanding the heterogeneity of dyslexia. The importance of characterizing the profiles of individuals with dyslexia becomes clear within the context of constructing remediation programs that are specifically targeted and are more effective in terms of intervention outcome.

    Additional information

    11145_2014_9504_MOESM1_ESM.doc
  • Peeters, D., Runnqvist, E., Bertrand, D., & Grainger, J. (2014). Asymmetrical switch costs in bilingual language production induced by reading words. Journal of Experimental Psychology: Learning, Memory, and Cognition, 40(1), 284-292. doi:10.1037/a0034060.

    Abstract

    We examined language-switching effects in French–English bilinguals using a paradigm where pictures are always named in the same language (either French or English) within a block of trials, and on each trial, the picture is preceded by a printed word from the same language or from the other language. Participants had to either make a language decision on the word or categorize it as an animal name or not. Picture-naming latencies in French (Language 1 [L1]) were slower when pictures were preceded by an English word than by a French word, independently of the task performed on the word. There were no language-switching effects when pictures were named in English (L2). This pattern replicates asymmetrical switch costs found with the cued picture-naming paradigm and shows that the asymmetrical pattern can be obtained (a) in the absence of artificial (nonlinguistic) language cues, (b) when the switch involves a shift from comprehension in 1 language to production in another, and (c) when the naming language is blocked (univalent response). We concluded that language switch costs in bilinguals cannot be reduced to effects driven by task control or response-selection mechanisms.
  • Peeters, D., & Dresler, M. (2014). The scientific significance of sleep-talking. Frontiers for Young Minds, 2(9). Retrieved from http://kids.frontiersin.org/articles/24/the_scientific_significance_of_sleep_talking/.

    Abstract

    Did one of your parents, siblings, or friends ever tell you that you were talking in your sleep? Nothing to be ashamed of! A recent study found that more than half of all people have had the experience of speaking out loud while being asleep [1]. This might even be underestimated, because often people do not notice that they are sleep-talking, unless somebody wakes them up or tells them the next day. Most neuroscientists, linguists, and psychologists studying language are interested in our language production and language comprehension skills during the day. In the present article, we will explore what is known about the production of overt speech during the night. We suggest that the study of sleep-talking may be just as interesting and informative as the study of wakeful speech.
  • Piai, V., Roelofs, A., Jensen, O., Schoffelen, J.-M., & Bonnefond, M. (2014). Distinct patterns of brain activity characterise lexical activation and competition in spoken word production. PLoS One, 9(2): e88674. doi:10.1371/journal.pone.0088674.

    Abstract

    According to a prominent theory of language production, concepts activate multiple associated words in memory, which enter into competition for selection. However, only a few electrophysiological studies have identified brain responses reflecting competition. Here, we report a magnetoencephalography study in which the activation of competing words was manipulated by presenting pictures (e.g., dog) with distractor words. The distractor and picture name were semantically related (cat), unrelated (pin), or identical (dog). Related distractors are stronger competitors to the picture name because they receive additional activation from the picture relative to other distractors. Picture naming times were longer with related than unrelated and identical distractors. Phase-locked and non-phase-locked activity were distinct but temporally related. Phase-locked activity in left temporal cortex, peaking at 400 ms, was larger on unrelated than related and identical trials, suggesting differential activation of alternative words by the picture-word stimuli. Non-phase-locked activity between roughly 350–650 ms (4–10 Hz) in left superior frontal gyrus was larger on related than unrelated and identical trials, suggesting differential resolution of the competition among the alternatives, as reflected in the naming times. These findings characterise distinct patterns of activity associated with lexical activation and competition, supporting the theory that words are selected by competition.
  • Schoot, L., Menenti, L., Hagoort, P., & Segaert, K. (2014). A little more conversation - The influence of communicative context on syntactic priming in brain and behavior. Frontiers in Psychology, 5: 208. doi:10.3389/fpsyg.2014.00208.

    Abstract

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

    Abstract

    Speakers sometimes repeat syntactic structures across sentences, a phenomenon called syntactic priming. We investigated the influence of verb-bound syntactic preferences on syntactic priming effects in response choices and response latencies for German ditransitive sentences. In the response choices we found inverse preference effects: There were stronger syntactic priming effects for primes in the less preferred structure, given the syntactic preference of the prime verb. In the response latencies we found positive preference effects: There were stronger syntactic priming effects for primes in the more preferred structure, given the syntactic preference of the prime verb. These findings provide further support for the idea that syntactic processing is lexically guided.
  • Shao, Z., Roelofs, A., Acheson, D. J., & Meyer, A. S. (2014). Electrophysiological evidence that inhibition supports lexical selection in picture naming. Brain Research, 1586, 130-142. doi:10.1016/j.brainres.2014.07.009.

    Abstract

    We investigated the neural basis of inhibitory control during lexical selection. Participants overtly named pictures while response times (RTs) and event-related brain potentials (ERPs) were recorded. The difficulty of lexical selection was manipulated by using object and action pictures with high name agreement (few response candidates) versus low name agreement (many response candidates). To assess the involvement of inhibition, we conducted delta plot analyses of naming RTs and examined the N2 component of the ERP. We found longer mean naming RTs and a larger N2 amplitude in the low relative to the high name agreement condition. For action naming we found a negative correlation between the slopes of the slowest delta segment and the difference in N2 amplitude between the low and high name agreement conditions. The converging behavioral and electrophysiological evidence suggests that selective inhibition is engaged to reduce competition during lexical selection in picture naming.
  • Silva, S., Branco, P., Barbosa, F., Marques-Teixeira, J., Petersson, K. M., & Castro, S. L. (2014). Musical phrase boundaries, wrap-up and the closure positive shift. Brain Research, 1585, 99-107. doi:10.1016/j.brainres.2014.08.025.

    Abstract

    We investigated global integration (wrap-up) processes at the boundaries of musical phrases by comparing the effects of well and non-well formed phrases on event-related potentials time-locked to two boundary points: the onset and the offset of the boundary pause. The Closure Positive Shift, which is elicited at the boundary offset, was not modulated by the quality of phrase structure (well vs. non-well formed). In contrast, the boundary onset potentials showed different patterns for well and non-well formed phrases. Our results contribute to specify the functional meaning of the Closure Positive Shift in music, shed light on the large-scale structural integration of musical input, and raise new hypotheses concerning shared resources between music and language.
  • Silva, S., Barbosa, F., Marques-Teixeira, J., Petersson, K. M., & Castro, S. L. (2014). You know when: Event-related potentials and theta/beat power indicate boundary prediction in music. Journal of Integrative Neuroscience, 13(1), 19-34. doi:10.1142/S0219635214500022.

    Abstract

    Neuroscientific and musicological approaches to music cognition indicate that listeners familiarized in the Western tonal tradition expect a musical phrase boundary at predictable time intervals. However, phrase boundary prediction processes in music remain untested. We analyzed event-related potentials (ERPs) and event-related induced power changes at the onset and offset of a boundary pause. We made comparisons with modified melodies, where the pause was omitted and filled by tones. The offset of the pause elicited a closure positive shift (CPS), indexing phrase boundary detection. The onset of the filling tones elicited significant increases in theta and beta powers. In addition, the P2 component was larger when the filling tones started than when they ended. The responses to boundary omission suggest that listeners expected to hear a boundary pause. Therefore, boundary prediction seems to coexist with boundary detection in music segmentation.
  • Simanova, I., Hagoort, P., Oostenveld, R., & Van Gerven, M. A. J. (2014). Modality-independent decoding of semantic information from the human brain. Cerebral Cortex, 24, 426-434. doi:10.1093/cercor/bhs324.

    Abstract

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

    Abstract

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

    Abstract

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

    Abstract

    To investigate the neural underpinnings of word decoding, and how it changes as a function of repeated exposure, we trained Dutch participants repeatedly over the course of a month of training to articulate a set of novel disyllabic input strings written in Greek script to avoid the use of familiar orthographic representations. The syllables in the input were phonotactically legal combinations but non-existent in the Dutch language, allowing us to assess their role in novel word decoding. Not only trained disyllabic pseudowords were tested but also pseudowords with recombined patterns of syllables to uncover the emergence of syllabic representations. We showed that with extensive training, articulation became faster and more accurate for the trained pseudowords. On the neural level, the initial stage of decoding was reflected by increased activity in visual attention areas of occipito-temporal and occipito-parietal cortices, and in motor coordination areas of the precentral gyrus and the inferior frontal gyrus. After one month of training, memory representations for holistic information (whole word unit) were established in areas encompassing the angular gyrus, the precuneus and the middle temporal gyrus. Syllabic representations also emerged through repeated training of disyllabic pseudowords, such that reading recombined syllables of the trained pseudowords showed similar brain activation to trained pseudowords and were articulated faster than novel combinations of letter strings used in the trained pseudowords.
  • Tsuji, S., & Cristia, A. (2014). Perceptual attunement in vowels: A meta-analysis. Developmental Psychobiology, 56(2), 179-191. doi:10.1002/dev.21179.

    Abstract

    Although the majority of evidence on perceptual narrowing in speech sounds is based on consonants, most models of infant speech perception generalize these findings to vowels, assuming that vowel perception improves for vowel sounds that are present in the infant's native language within the first year of life, and deteriorates for non-native vowel sounds over the same period of time. The present meta-analysis contributes to assessing to what extent these descriptions are accurate in the first comprehensive quantitative meta-analysis of perceptual narrowing in infant vowel discrimination, including results from behavioral, electrophysiological, and neuroimaging methods applied to infants 0–14 months of age. An analysis of effect sizes for native and non-native vowel discrimination over the first year of life revealed that they changed with age in opposite directions, being significant by about 6 months of age
  • Van Leeuwen, T. M., Lamers, M. J. A., Petersson, K. M., Gussenhoven, C., Poser, B., & Hagoort, P. (2014). Phonological markers of information structure: An fMRI study. Neuropsychologia, 58(1), 64-74. doi:10.1016/j.neuropsychologia.2014.03.017.

    Abstract

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

    Additional information

    mmc1.doc
  • Veenstra, A., Acheson, D. J., Bock, K., & Meyer, A. S. (2014). Effects of semantic integration on subject–verb agreement: Evidence from Dutch. Language, Cognition and Neuroscience, 29(3), 355-380. doi:10.1080/01690965.2013.862284.

    Abstract

    The generation of subject–verb agreement is a central component of grammatical encoding. It is sensitive to conceptual and grammatical influences, but the interplay between these factors is still not fully understood. We investigate how semantic integration of the subject noun phrase (‘the secretary of/with the governor’) and the Local Noun Number (‘the secretary with the governor/governors’) affect the ease of selecting the verb form. Two hypotheses are assessed: according to the notional hypothesis, integration encourages the assignment of the singular notional number to the noun phrase and facilitates the choice of the singular verb form. According to the lexical interference hypothesis, integration strengthens the competition between nouns within the subject phrase, making it harder to select the verb form when the nouns mismatch in number. In two experiments, adult speakers of Dutch completed spoken preambles (Experiment 1) or selected appropriate verb forms (Experiment 2). Results showed facilitatory effects of semantic integration (fewer errors and faster responses with increasing integration). These effects did not interact with the effects of the Local Noun Number (slower response times and higher error rates for mismatching than for matching noun numbers). The findings thus support the notional hypothesis and a model of agreement where conceptual and lexical factors independently contribute to the determination of the number of the subject noun phrase and, ultimately, the verb.
  • Veenstra, A., Acheson, D. J., & Meyer, A. S. (2014). Keeping it simple: Studying grammatical encoding with lexically-reduced item sets. Frontiers in Psychology, 5: 783. doi:10.3389/fpsyg.2014.00783.

    Abstract

    Compared to the large body of work on lexical access, little research has been done on grammatical encoding in language production. An exception is the generation of subject-verb agreement. Here, two key findings have been reported: (1) Speakers make more agreement errors when the head and local noun of a phrase mismatch in number than when they match (e.g., the key to the cabinet(s)); and (2) this attraction effect is asymmetric, with stronger attraction for singular than for plural head nouns. Although these findings are robust, the cognitive processes leading to agreement errors and their significance for the generation of correct agreement are not fully understood. We propose that future studies of agreement, and grammatical encoding in general, may benefit from using paradigms that tightly control the variability of the lexical content of the material. We report two experiments illustrating this approach. In both of them, the experimental items featured combinations of four nouns, four color adjectives, and two prepositions. In Experiment 1, native speakers of Dutch described pictures in sentences such as the circle next to the stars is blue. In Experiment 2, they carried out a forced-choice task, where they read subject noun phrases (e.g., the circle next to the stars) and selected the correct verb-phrase (is blue or are blue) with a button press. Both experiments showed an attraction effect, with more errors after subject phrases with mismatching, compared to matching head and local nouns. This effect was stronger for singular than plural heads, replicating the attraction asymmetry. In contrast, the response times recorded in Experiment 2 showed similar attraction effects for singular and plural head nouns. These results demonstrate that critical agreement phenomena can be elicited reliably in lexically-reduced contexts. We discuss the theoretical implications of the findings and the potential and limitations of studies using lexically simple materials.
  • Wegman, J., Fonteijn, H. M., van Ekert, J., Tyborowska, A., Jansen, C., & Janzen, G. (2014). Gray and white matter correlates of navigational ability in humans. Human Brain Mapping, 35(6), 2561-2572. doi:10.1002/hbm.22349.

    Abstract

    Humans differ widely in their navigational abilities. Studies have shown that self-reports on navigational abilities are good predictors of performance on navigation tasks in real and virtual environments. The caudate nucleus and medial temporal lobe regions have been suggested to subserve different navigational strategies. The ability to use different strategies might underlie navigational ability differences. This study examines the anatomical correlates of self-reported navigational ability in both gray and white matter. Local gray matter volume was compared between a group (N = 134) of good and bad navigators using voxel-based morphometry (VBM), as well as regional volumes. To compare between good and bad navigators, we also measured white matter anatomy using diffusion tensor imaging (DTI) and looked at fractional anisotropy (FA) values. We observed a trend toward higher local GM volume in right anterior parahippocampal/rhinal cortex for good versus bad navigators. Good male navigators showed significantly higher local GM volume in right hippocampus than bad male navigators. Conversely, bad navigators showed increased FA values in the internal capsule, the white matter bundle closest to the caudate nucleus and a trend toward higher local GM volume in the caudate nucleus. Furthermore, caudate nucleus regional volume correlated negatively with navigational ability. These convergent findings across imaging modalities are in line with findings showing that the caudate nucleus and the medial temporal lobes are involved in different wayfinding strategies. Our study is the first to show a link between self-reported large-scale navigational abilities and different measures of brain anatomy.
  • Whitmarsh, S., Barendregt, H., Schoffelen, J.-M., & Jensen, O. (2014). Metacognitive awareness of covert somatosensory attention corresponds to contralateral alpha power. NeuroImage, 85(2), 803-809. doi:10.1016/j.neuroimage.2013.07.031.

    Abstract

    Studies on metacognition have shown that participants can report on their performance on a wide range of perceptual, memory and behavioral tasks. We know little, however, about the ability to report on one's attentional focus. The degree and direction of somatosensory attention can, however, be readily discerned through suppression of alpha band frequencies in EEG/MEG produced by the somatosensory cortex. Such top-down attentional modulations of cortical excitability have been shown to result in better discrimination performance and decreased response times. In this study we asked whether the degree of attentional focus is also accessible for subjective report, and whether such evaluations correspond to the amount of somatosensory alpha activity. In response to auditory cues participants maintained somatosensory attention to either their left or right hand for intervals varying randomly between 5 and 32seconds, while their brain activity was recorded with MEG. Trials were terminated by a probe sound, to which they reported their level of attention on the cued hand right before probe-onset. Using a beamformer approach, we quantified the alpha activity in left and right somatosensory regions, one second before the probe. Alpha activity from contra- and ipsilateral somatosensory cortices for high versus low attention trials were compared. As predicted, the contralateral somatosensory alpha depression correlated with higher reported attentional focus. Finally, alpha activity two to three seconds before the probe-onset was correlated with attentional focus. We conclude that somatosensory attention is indeed accessible to metacognitive awareness.
  • Willems, R. M., Van der Haegen, L., Fisher, S. E., & Francks, C. (2014). On the other hand: Including left-handers in cognitive neuroscience and neurogenetics. Nature Reviews Neuroscience, 15, 193-201. doi:10.1038/nrn3679.

    Abstract

    Left-handers are often excluded from study cohorts in neuroscience and neurogenetics in order to reduce variance in the data. However, recent investigations have shown that the inclusion or targeted recruitment of left-handers can be informative in studies on a range of topics, such as cerebral lateralization and the genetic underpinning of asymmetrical brain development. Left-handed individuals represent a substantial portion of the human population and therefore left-handedness falls within the normal range of human diversity; thus, it is important to account for this variation in our understanding of brain functioning. We call for neuroscientists and neurogeneticists to recognize the potential of studying this often-discarded group of research subjects.
  • Willems, R. M., & Francks, C. (2014). Your left-handed brain. Frontiers for Young Minds, 2: 13. doi:10.3389/frym.2014.00013.

    Abstract

    While most people prefer to use their right hand to brush their teeth, throw a ball, or hold a tennis racket, left-handers prefer to use their left hand. This is the case for around 10 per cent of all people. There was a time (not so long ago) when left-handers were stigmatized in Western (and other) communities: it was considered a bad sign if you were left-handed, and left-handed children were often forced to write with their right hand. This is nonsensical: there is nothing wrong with being left-handed, and trying to write with the non-preferred hand is frustrating for almost everybody. As a matter of fact, science can learn from left-handers, and in this paper, we discuss how this may be the case. We review why some people are left-handed and others are not, how left-handers' brains differ from right-handers’, and why scientists study left-handedness in the first place
  • De Zubicaray, G. I., Hartsuiker, R. J., & Acheson, D. J. (2014). Mind what you say—general and specific mechanisms for monitoring in speech production. Frontiers in Human Neuroscience, 8: 514. doi:10.3389%2Ffnhum.2014.00514.

    Abstract

    For most people, speech production is relatively effortless and error-free. Yet it has long been recognized that we need some type of control over what we are currently saying and what we plan to say. Precisely how we monitor our internal and external speech has been a topic of research interest for several decades. The predominant approach in psycholinguistics has assumed monitoring of both is accomplished via systems responsible for comprehending others' speech.

    This special topic aimed to broaden the field, firstly by examining proposals that speech production might also engage more general systems, such as those involved in action monitoring. A second aim was to examine proposals for a production-specific, internal monitor. Both aims require that we also specify the nature of the representations subject to monitoring.
  • Zumer, J. M., Scheeringa, R., Schoffelen, J.-M., Norris, D. G., & Jensen, O. (2014). Occipital alpha activity during stimulus processing gates the information flow to object-selective cortex. PLoS Biology, 12(10): e1001965. doi:10.1371/journal.pbio.1001965.

    Abstract

    Given the limited processing capabilities of the sensory system, it is essential that attended information is gated to downstream areas, whereas unattended information is blocked. While it has been proposed that alpha band (8–13 Hz) activity serves to route information to downstream regions by inhibiting neuronal processing in task-irrelevant regions, this hypothesis remains untested. Here we investigate how neuronal oscillations detected by electroencephalography in visual areas during working memory encoding serve to gate information reflected in the simultaneously recorded blood-oxygenation-level-dependent (BOLD) signals recorded by functional magnetic resonance imaging in downstream ventral regions. We used a paradigm in which 16 participants were presented with faces and landscapes in the right and left hemifields; one hemifield was attended and the other unattended. We observed that decreased alpha power contralateral to the attended object predicted the BOLD signal representing the attended object in ventral object-selective regions. Furthermore, increased alpha power ipsilateral to the attended object predicted a decrease in the BOLD signal representing the unattended object. We also found that the BOLD signal in the dorsal attention network inversely correlated with visual alpha power. This is the first demonstration, to our knowledge, that oscillations in the alpha band are implicated in the gating of information from the visual cortex to the ventral stream, as reflected in the representationally specific BOLD signal. This link of sensory alpha to downstream activity provides a neurophysiological substrate for the mechanism of selective attention during stimulus processing, which not only boosts the attended information but also suppresses distraction. Although previous studies have shown a relation between the BOLD signal from the dorsal attention network and the alpha band at rest, we demonstrate such a relation during a visuospatial task, indicating that the dorsal attention network exercises top-down control of visual alpha activity.

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