Publications

Displaying 1 - 100 of 167
  • Armeni, K., Willems, R. M., Van den Bosch, A., & Schoffelen, J.-M. (2019). Frequency-specific brain dynamics related to prediction during language comprehension. NeuroImage, 198, 283-295. doi:10.1016/j.neuroimage.2019.04.083.

    Abstract

    The brain's remarkable capacity to process spoken language virtually in real time requires fast and efficient information processing machinery. In this study, we investigated how frequency-specific brain dynamics relate to models of probabilistic language prediction during auditory narrative comprehension. We recorded MEG activity while participants were listening to auditory stories in Dutch. Using trigram statistical language models, we estimated for every word in a story its conditional probability of occurrence. On the basis of word probabilities, we computed how unexpected the current word is given its context (word perplexity) and how (un)predictable the current linguistic context is (word entropy). We then evaluated whether source-reconstructed MEG oscillations at different frequency bands are modulated as a function of these language processing metrics. We show that theta-band source dynamics are increased in high relative to low entropy states, likely reflecting lexical computations. Beta-band dynamics are increased in situations of low word entropy and perplexity possibly reflecting maintenance of ongoing cognitive context. These findings lend support to the idea that the brain engages in the active generation and evaluation of predicted language based on the statistical properties of the input signal.

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  • Bocanegra, B. R., Poletiek, F. H., Ftitache, B., & Clark, A. (2019). Intelligent problem-solvers externalize cognitive operations. Nature Human Behaviour, 3, 136-142. doi:10.1038/s41562-018-0509-y.

    Abstract

    Humans are nature’s most intelligent and prolific users of external props and aids (such as written texts, slide-rules and software packages). Here we introduce a method for investigating how people make active use of their task environment during problem-solving and apply this approach to the non-verbal Raven Advanced Progressive Matrices test for fluid intelligence. We designed a click-and-drag version of the Raven test in which participants could create different external spatial configurations while solving the puzzles. In our first study, we observed that the click-and-drag test was better than the conventional static test at predicting academic achievement of university students. This pattern of results was partially replicated in a novel sample. Importantly, environment-altering actions were clustered in between periods of apparent inactivity, suggesting that problem-solvers were delicately balancing the execution of internal and external cognitive operations. We observed a systematic relationship between this critical phasic temporal signature and improved test performance. Our approach is widely applicable and offers an opportunity to quantitatively assess a powerful, although understudied, feature of human intelligence: our ability to use external objects, props and aids to solve complex problems.
  • Bosker, H. R., Van Os, M., Does, R., & Van Bergen, G. (2019). Counting 'uhm's: how tracking the distribution of native and non-native disfluencies influences online language comprehension. Journal of Memory and Language, 106, 189-202. doi:10.1016/j.jml.2019.02.006.

    Abstract

    Disfluencies, like 'uh', have been shown to help listeners anticipate reference to low-frequency words. The associative account of this 'disfluency bias' proposes that listeners learn to associate disfluency with low-frequency referents based on prior exposure to non-arbitrary disfluency distributions (i.e., greater probability of low-frequency words after disfluencies). However, there is limited evidence for listeners actually tracking disfluency distributions online. The present experiments are the first to show that adult listeners, exposed to a typical or more atypical disfluency distribution (i.e., hearing a talker unexpectedly say uh before high-frequency words), flexibly adjust their predictive strategies to the disfluency distribution at hand (e.g., learn to predict high-frequency referents after disfluency). However, when listeners were presented with the same atypical disfluency distribution but produced by a non-native speaker, no adjustment was observed. This suggests pragmatic inferences can modulate distributional learning, revealing the flexibility of, and constraints on, distributional learning in incremental language comprehension.
  • Fields, E. C., Weber, K., Stillerman, B., Delaney-Busch, N., & Kuperberg, G. (2019). Functional MRI reveals evidence of a self-positivity bias in the medial prefrontal cortex during the comprehension of social vignettes. Social Cognitive and Affective Neuroscience, 14(6), 613-621. doi:10.1093/scan/nsz035.

    Abstract

    A large literature in social neuroscience has associated the medial prefrontal cortex (mPFC) with the processing of self-related information. However, only recently have social neuroscience studies begun to consider the large behavioral literature showing a strong self-positivity bias, and these studies have mostly focused on its correlates during self-related judgments and decision making. We carried out a functional MRI (fMRI) study to ask whether the mPFC would show effects of the self-positivity bias in a paradigm that probed participants’ self-concept without any requirement of explicit self-judgment. We presented social vignettes that were either self-relevant or non-self-relevant with a neutral, positive, or negative outcome described in the second sentence. In previous work using event-related potentials, this paradigm has shown evidence of a self-positivity bias that influences early stages of semantically processing incoming stimuli. In the present fMRI study, we found evidence for this bias within the mPFC: an interaction between self-relevance and valence, with only positive scenarios showing a self vs other effect within the mPFC. We suggest that the mPFC may play a role in maintaining a positively-biased self-concept and discuss the implications of these findings for the social neuroscience of the self and the role of the mPFC.

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  • Fitz, H., & Chang, F. (2019). Language ERPs reflect learning through prediction error propagation. Cognitive Psychology, 111, 15-52. doi:10.1016/j.cogpsych.2019.03.002.

    Abstract

    Event-related potentials (ERPs) provide a window into how the brain is processing language. Here, we propose a theory that argues that ERPs such as the N400 and P600 arise as side effects of an error-based learning mechanism that explains linguistic adaptation and language learning. We instantiated this theory in a connectionist model that can simulate data from three studies on the N400 (amplitude modulation by expectancy, contextual constraint, and sentence position), five studies on the P600 (agreement, tense, word category, subcategorization and garden-path sentences), and a study on the semantic P600 in role reversal anomalies. Since ERPs are learning signals, this account explains adaptation of ERP amplitude to within-experiment frequency manipulations and the way ERP effects are shaped by word predictability in earlier sentences. Moreover, it predicts that ERPs can change over language development. The model provides an account of the sensitivity of ERPs to expectation mismatch, the relative timing of the N400 and P600, the semantic nature of the N400, the syntactic nature of the P600, and the fact that ERPs can change with experience. This approach suggests that comprehension ERPs are related to sentence production and language acquisition mechanisms
  • Franken, M. K., Acheson, D. J., McQueen, J. M., Hagoort, P., & Eisner, F. (2019). Consistency influences altered auditory feedback processing. Quarterly Journal of Experimental Psychology, 72(10), 2371-2379. doi:10.1177/1747021819838939.

    Abstract

    Previous research on the effect of perturbed auditory feedback in speech production has focused on two types of responses. In the short term, speakers generate compensatory motor commands in response to unexpected perturbations. In the longer term, speakers adapt feedforward motor programmes in response to feedback perturbations, to avoid future errors. The current study investigated the relation between these two types of responses to altered auditory feedback. Specifically, it was hypothesised that consistency in previous feedback perturbations would influence whether speakers adapt their feedforward motor programmes. In an altered auditory feedback paradigm, formant perturbations were applied either across all trials (the consistent condition) or only to some trials, whereas the others remained unperturbed (the inconsistent condition). The results showed that speakers’ responses were affected by feedback consistency, with stronger speech changes in the consistent condition compared with the inconsistent condition. Current models of speech-motor control can explain this consistency effect. However, the data also suggest that compensation and adaptation are distinct processes, which are not in line with all current models.
  • Gao, Y., Zheng, L., Liu, X., Nichols, E. S., Zhang, M., Shang, L., Ding, G., Meng, Z., & Liu, L. (2019). First and second language reading difficulty among Chinese–English bilingual children: The prevalence and influences from demographic characteristics. Frontiers in Psychology, 10: 2544. doi:10.3389/fpsyg.2019.02544.

    Abstract

    Learning to read a second language (L2) can pose a great challenge for children who have already been struggling to read in their first language (L1). Moreover, it is not clear whether, to what extent, and under what circumstances L1 reading difficulty increases the risk of L2 reading difficulty. This study investigated Chinese (L1) and English (L2) reading skills in a large representative sample of 1,824 Chinese–English bilingual children in Grades 4 and 5 from both urban and rural schools in Beijing. We examined the prevalence of reading difficulty in Chinese only (poor Chinese readers, PC), English only (poor English readers, PE), and both Chinese and English (poor bilingual readers, PB) and calculated the co-occurrence, that is, the chances of becoming a poor reader in English given that the child was already a poor reader in Chinese. We then conducted a multinomial logistic regression analysis and compared the prevalence of PC, PE, and PB between children in Grade 4 versus Grade 5, in urban versus rural areas, and in boys versus girls. Results showed that compared to girls, boys demonstrated significantly higher risk of PC, PE, and PB. Meanwhile, compared to the 5th graders, the 4th graders demonstrated significantly higher risk of PC and PB. In addition, children enrolled in the urban schools were more likely to become better second language readers, thus leading to a concerning rural–urban gap in the prevalence of L2 reading difficulty. Finally, among these Chinese–English bilingual children, regardless of sex and school location, poor reading skill in Chinese significantly increased the risk of also being a poor English reader, with a considerable and stable co-occurrence of approximately 36%. In sum, this study suggests that despite striking differences between alphabetic and logographic writing systems, L1 reading difficulty still significantly increases the risk of L2 reading difficulty. This indicates the shared meta-linguistic skills in reading different writing systems and the importance of understanding the universality and the interdependent relationship of reading between different writing systems. Furthermore, the male disadvantage (in both L1 and L2) and the urban–rural gap (in L2) found in the prevalence of reading difficulty calls for special attention to disadvantaged populations in educational practice.
  • Gao, X., Dera, J., Nijhoff, A. D., & Willems, R. M. (2019). Is less readable liked better? The case of font readability in poetry appreciation. PLoS One, 14(12): e0225757. doi:10.1371/journal.pone.0225757.

    Abstract

    Previous research shows conflicting findings for the effect of font readability on comprehension and memory for language. It has been found that—perhaps counterintuitively–a hard to read font can be beneficial for language comprehension, especially for difficult language. Here we test how font readability influences the subjective experience of poetry reading. In three experiments we tested the influence of poem difficulty and font readability on the subjective experience of poems. We specifically predicted that font readability would have opposite effects on the subjective experience of easy versus difficult poems. Participants read poems which could be more or less difficult in terms of conceptual or structural aspects, and which were presented in a font that was either easy or more difficult to read. Participants read existing poems and subsequently rated their subjective experience (measured through four dependent variables: overall liking, perceived flow of the poem, perceived topic clarity, and perceived structure). In line with previous literature we observed a Poem Difficulty x Font Readability interaction effect for subjective measures of poetry reading. We found that participants rated easy poems as nicer when presented in an easy to read font, as compared to when presented in a hard to read font. Despite the presence of the interaction effect, we did not observe the predicted opposite effect for more difficult poems. We conclude that font readability can influence reading of easy and more difficult poems differentially, with strongest effects for easy poems.

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    https://osf.io/jwcqt/
  • Gehrig, J., Michalareas, G., Forster, M.-T., Lei, J., Hok, P., Laufs, H., Senft, C., Seifert, V., Schoffelen, J.-M., Hanslmayr, H., & Kell, C. A. (2019). Low-frequency oscillations code speech during verbal working memory. The Journal of Neuroscience, 39(33), 6498-6512. doi:10.1523/JNEUROSCI.0018-19.2019.

    Abstract

    The way the human brain represents speech in memory is still unknown. An obvious characteristic of speech is its evolvement over time.
    During speech processing, neural oscillations are modulated by the temporal properties of the acoustic speech signal, but also acquired
    knowledge on the temporal structure of language influences speech perception-related brain activity. This suggests that speech could be
    represented in the temporal domain, a form of representation that the brain also uses to encode autobiographic memories. Empirical
    evidence for such a memory code is lacking. We investigated the nature of speech memory representations using direct cortical recordings
    in the left perisylvian cortex during delayed sentence reproduction in female and male patients undergoing awake tumor surgery.
    Our results reveal that the brain endogenously represents speech in the temporal domain. Temporal pattern similarity analyses revealed
    that the phase of frontotemporal low-frequency oscillations, primarily in the beta range, represents sentence identity in working memory.
    The positive relationship between beta power during working memory and task performance suggests that working memory
    representations benefit from increased phase separation.
  • Hagoort, P. (2019). The meaning making mechanism(s) behind the eyes and between the ears. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 375: 20190301. doi:10.1098/rstb.2019.0301.

    Abstract

    In this contribution, the following four questions are discussed: (i) where is meaning?; (ii) what is meaning?; (iii) what is the meaning of mechanism?; (iv) what are the mechanisms of meaning? I will argue that meanings are in the head. Meanings have multiple facets, but minimally one needs to make a distinction between single word meanings (lexical meaning) and the meanings of multi-word utterances. The latter ones cannot be retrieved from memory, but need to be constructed on the fly. A mechanistic account of the meaning-making mind requires an analysis at both a functional and a neural level, the reason being that these levels are causally interdependent. I will show that an analysis exclusively focusing on patterns of brain activation lacks explanatory power. Finally, I shall present an initial sketch of how the dynamic interaction between temporo-parietal areas and inferior frontal cortex might instantiate the interpretation of linguistic utterances in the context of a multimodal setting and ongoing discourse information.
  • Hagoort, P. (2019). The neurobiology of language beyond single word processing. Science, 366(6461), 55-58. doi:10.1126/science.aax0289.

    Abstract

    In this Review, I propose a multiple-network view for the neurobiological basis of distinctly human language skills. A much more complex picture of interacting brain areas emerges than in the classical neurobiological model of language. This is because using language is more than single-word processing, and much goes on beyond the information given in the acoustic or orthographic tokens that enter primary sensory cortices. This requires the involvement of multiple networks with functionally nonoverlapping contributions

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  • Hervais-Adelman, A., Kumar, U., Mishra, R. K., Tripathi, V. N., Guleria, A., Singh, J. P., Eisner, F., & Huettig, F. (2019). Learning to read recycles visual cortical networks without destruction. Science Advances, 5(9): eaax0262. doi:10.1126/sciadv.aax0262.

    Abstract

    Learning to read is associated with the appearance of an orthographically sensitive brain region known as the visual word form area. It has been claimed that development of this area proceeds by impinging upon territory otherwise available for the processing of culturally relevant stimuli such as faces and houses. In a large-scale functional magnetic resonance imaging study of a group of individuals of varying degrees of literacy (from completely illiterate to highly literate), we examined cortical responses to orthographic and nonorthographic visual stimuli. We found that literacy enhances responses to other visual input in early visual areas and enhances representational similarity between text and faces, without reducing the extent of response to nonorthographic input. Thus, acquisition of literacy in childhood recycles existing object representation mechanisms but without destructive competition.

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  • Heyselaar, E., & Segaert, K. (2019). Memory encoding of syntactic information involves domain-general attentional resources. Evidence from dual-task studies. Quarterly Journal of Experimental Psychology, 72(6), 1285-1296. doi:10.1177/1747021818801249.

    Abstract

    We investigate the type of attention (domain-general or language-specific) used during
    syntactic processing. We focus on syntactic priming: In this task, participants listen to a
    sentence that describes a picture (prime sentence), followed by a picture the participants need
    to describe (target sentence). We measure the proportion of times participants use the
    syntactic structure they heard in the prime sentence to describe the current target sentence as a
    measure of syntactic processing. Participants simultaneously conducted a motion-object
    tracking (MOT) task, a task commonly used to tax domain-general attentional resources. We
    manipulated the number of objects the participant had to track; we thus measured
    participants’ ability to process syntax while their attention is not-, slightly-, or overly-taxed.
    Performance in the MOT task was significantly worse when conducted as a dual-task
    compared to as a single task. We observed an inverted U-shaped curve on priming magnitude
    when conducting the MOT task concurrently with prime sentences (i.e., memory encoding),
    but no effect when conducted with target sentences (i.e., memory retrieval). Our results
    illustrate how, during the encoding of syntactic information, domain-general attention
    differentially affects syntactic processing, whereas during the retrieval of syntactic
    information domain-general attention does not influence syntactic processing
  • Hubbard, R. J., Rommers, J., Jacobs, C. L., & Federmeier, K. D. (2019). Downstream behavioral and electrophysiological consequences of word prediction on recognition memory. Frontiers in Human Neuroscience, 13: 291. doi:10.3389/fnhum.2019.00291.

    Abstract

    When people process language, they can use context to predict upcoming information,
    influencing processing and comprehension as seen in both behavioral and neural
    measures. Although numerous studies have shown immediate facilitative effects
    of confirmed predictions, the downstream consequences of prediction have been
    less explored. In the current study, we examined those consequences by probing
    participants’ recognition memory for words after they read sets of sentences.
    Participants read strongly and weakly constraining sentences with expected or
    unexpected endings (“I added my name to the list/basket”), and later were tested on
    their memory for the sentence endings while EEG was recorded. Critically, the memory
    test contained words that were predictable (“list”) but were never read (participants
    saw “basket”). Behaviorally, participants showed successful discrimination between old
    and new items, but false alarmed to the expected-item lures more often than to new
    items, showing that predicted words or concepts can linger, even when predictions
    are disconfirmed. Although false alarm rates did not differ by constraint, event-related
    potentials (ERPs) differed between false alarms to strongly and weakly predictable words.
    Additionally, previously unexpected (compared to previously expected) endings that
    appeared on the memory test elicited larger N1 and LPC amplitudes, suggesting greater
    attention and episodic recollection. In contrast, highly predictable sentence endings that
    had been read elicited reduced LPC amplitudes during the memory test. Thus, prediction
    can facilitate processing in the moment, but can also lead to false memory and reduced
    recollection for predictable information.
  • Hulten, A., Schoffelen, J.-M., Udden, J., Lam, N. H. L., & Hagoort, P. (2019). How the brain makes sense beyond the processing of single words – An MEG study. NeuroImage, 186, 586-594. doi:10.1016/j.neuroimage.2018.11.035.

    Abstract

    Human language processing involves combinatorial operations that make human communication stand out in the animal kingdom. These operations rely on a dynamic interplay between the inferior frontal and the posterior temporal cortices. Using source reconstructed magnetoencephalography, we tracked language processing in the brain, in order to investigate how individual words are interpreted when part of sentence context. The large sample size in this study (n = 68) allowed us to assess how event-related activity is associated across distinct cortical areas, by means of inter-areal co-modulation within an individual. We showed that, within 500 ms of seeing a word, the word's lexical information has been retrieved and unified with the sentence context. This does not happen in a strictly feed-forward manner, but by means of co-modulation between the left posterior temporal cortex (LPTC) and left inferior frontal cortex (LIFC), for each individual word. The co-modulation of LIFC and LPTC occurs around 400 ms after the onset of each word, across the progression of a sentence. Moreover, these core language areas are supported early on by the attentional network. The results provide a detailed description of the temporal orchestration related to single word processing in the context of ongoing language.

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    1-s2.0-S1053811918321165-mmc1.pdf
  • De Kleijn, R., Wijnen, M., & Poletiek, F. H. (2019). The effect of context-dependent information and sentence constructions on perceived humanness of an agent in a Turing test. Knowledge-Based Systems, 163, 794-799. doi:10.1016/j.knosys.2018.10.006.

    Abstract

    In a Turing test, a judge decides whether their conversation partner is either a machine or human. What cues does the judge use to determine this? In particular, are presumably unique features of human language actually perceived as humanlike? Participants rated the humanness of a set of sentences that were manipulated for grammatical construction: linear right-branching or hierarchical center-embedded and their plausibility with regard to world knowledge.

    We found that center-embedded sentences are perceived as less humanlike than right-branching sentences and more plausible sentences are regarded as more humanlike. However, the effect of plausibility of the sentence on perceived humanness is smaller for center-embedded sentences than for right-branching sentences.

    Participants also rated a conversation with either correct or incorrect use of the context by the agent. No effect of context use was found. Also, participants rated a full transcript of either a real human or a real chatbot, and we found that chatbots were reliably perceived as less humanlike than real humans, in line with our expectation. We did, however, find individual differences between chatbots and humans.
  • Kochari, A., & Flecken, M. (2019). Lexical prediction in language comprehension: A replication study of grammatical gender effects in Dutch. Language, Cognition and Neuroscience, 34(2), 239-253. doi:10.1080/23273798.2018.1524500.

    Abstract

    An important question in predictive language processing is the extent to which prediction effects can reliably be measured on pre-nominal material (e.g. articles before nouns). Here, we present a large sample (N = 58) close replication of a study by Otten and van Berkum (2009). They report ERP modulations in relation to the predictability of nouns in sentences, measured on gender-marked Dutch articles. We used nearly identical materials, procedures, and data analysis steps. We fail to replicate the original effect, but do observe a pattern consistent with the original data. Methodological differences between our replication and the original study that could potentially have contributed to the diverging results are discussed. In addition, we discuss the suitability of Dutch gender-marked determiners as a test-case for future studies of pre-activation of lexical items.
  • Kuperberg, G., Weber, K., Delaney-Busch, N., Ustine, C., Stillerman, B., Hämäläinen, M., & Lau, E. (2019). Multimodal neuroimaging evidence for looser lexico-semantic connections in schizophrenia. Neuropsychologia, 124, 337-349. doi:10.1016/j.neuropsychologia.2018.10.024.

    Abstract

    It has been hypothesized that schizophrenia is characterized by overly broad automatic activity within lexico-semantic networks. We used two complementary neuroimaging techniques, Magnetoencephalography (MEG) and functional Magnetic Resonance Imaging (fMRI), in combination with a highly automatic indirect semantic priming paradigm, to spatiotemporally localize this abnormality in the brain.

    Eighteen people with schizophrenia and 20 demographically-matched control participants viewed target words (“bell”) preceded by directly related (“church”), indirectly related (“priest”), or unrelated (“pants”) prime words in MEG and fMRI sessions. To minimize top-down processing, the prime was masked, the target appeared only 140ms after prime onset, and participants simply monitored for words within a particular semantic category that appeared in filler trials.

    Both techniques revealed a significantly larger automatic indirect priming effect in people with schizophrenia than in control participants. MEG temporally localized this enhanced effect to the N400 time window (300-500ms) — the critical stage of accessing meaning from words. fMRI spatially localized the effect to the left temporal fusiform cortex, which plays a role in mapping of orthographic word-form on to meaning. There was no evidence of an enhanced automatic direct semantic priming effect in the schizophrenia group.

    These findings provide converging neural evidence for abnormally broad highly automatic lexico-semantic activity in schizophrenia. We argue that, rather than arising from an unconstrained spread of automatic activation across semantic memory, this broader automatic lexico-semantic activity stems from looser connections between the form and meaning of words.

    Additional information

    1-s2.0-S0028393218307310-mmc1.docx
  • Mak, M., & Willems, R. M. (2019). Mental simulation during literary reading: Individual differences revealed with eye-tracking. Language, Cognition and Neuroscience, 34(4), 511-535. doi:10.1080/23273798.2018.1552007.

    Abstract

    People engage in simulation when reading literary narratives. In this study, we tried to pinpoint how different kinds of simulation (perceptual and motor simulation, mentalising) affect reading behaviour. Eye-tracking (gaze durations, regression probability) and questionnaire data were collected from 102 participants, who read three literary short stories. In a pre-test, 90 additional participants indicated which parts of the stories were high in one of the three kinds of simulation-eliciting content. The results show that motor simulation reduces gaze duration (faster reading), whereas perceptual simulation and mentalising increase gaze duration (slower reading). Individual differences in the effect of simulation on gaze duration were found, which were related to individual differences in aspects of story world absorption and story appreciation. These findings suggest fundamental differences between different kinds of simulation and confirm the role of simulation in absorption and appreciation.
  • Mantegna, F., Hintz, F., Ostarek, M., Alday, P. M., & Huettig, F. (2019). Distinguishing integration and prediction accounts of ERP N400 modulations in language processing through experimental design. Neuropsychologia, 134: 107199. doi:10.1016/j.neuropsychologia.2019.107199.

    Abstract

    Prediction of upcoming input is thought to be a main characteristic of language processing (e.g. Altmann & Mirkovic, 2009; Dell & Chang, 2014; Federmeier, 2007; Ferreira & Chantavarin, 2018; Pickering & Gambi, 2018; Hale, 2001; Hickok, 2012; Huettig 2015; Kuperberg & Jaeger, 2016; Levy, 2008; Norris, McQueen, & Cutler, 2016; Pickering & Garrod, 2013; Van Petten & Luka, 2012). One of the main pillars of experimental support for this notion comes from studies that have attempted to measure electrophysiological markers of prediction when participants read or listened to sentences ending in highly predictable words. The N400, a negative-going and centro-parietally distributed component of the ERP occurring approximately 400ms after (target) word onset, has been frequently interpreted as indexing prediction of the word (or the semantic representations and/or the phonological form of the predicted word, see Kutas & Federmeier, 2011; Nieuwland, 2019; Van Petten & Luka, 2012; for review). A major difficulty for interpreting N400 effects in language processing however is that it has been difficult to establish whether N400 target word modulations conclusively reflect prediction rather than (at least partly) ease of integration. In the present exploratory study, we attempted to distinguish lexical prediction (i.e. ‘top-down’ activation) from lexical integration (i.e. ‘bottom-up’ activation) accounts of ERP N400 modulations in language processing.
  • Martinez-Conde, S., Alexander, R. G., Blum, D., Britton, N., Lipska, B. K., Quirk, G. J., Swiss, J. I., Willems, R. M., & Macknik, S. L. (2019). The storytelling brain: How neuroscience stories help bridge the gap between research and society. The Journal of Neuroscience, 39(42), 8285-8290. doi:10.1523/JNEUROSCI.1180-19.2019.

    Abstract

    Active communication between researchers and society is necessary for the scientific community’s involvement in developing sciencebased
    policies. This need is recognized by governmental and funding agencies that compel scientists to increase their public engagement
    and disseminate research findings in an accessible fashion. Storytelling techniques can help convey science by engaging people’s imagination
    and emotions. Yet, many researchers are uncertain about how to approach scientific storytelling, or feel they lack the tools to
    undertake it. Here we explore some of the techniques intrinsic to crafting scientific narratives, as well as the reasons why scientific
    storytellingmaybe an optimal way of communicating research to nonspecialists.Wealso point out current communication gaps between
    science and society, particularly in the context of neurodiverse audiences and those that include neurological and psychiatric patients.
    Present shortcomings may turn into areas of synergy with the potential to link neuroscience education, research, and advocacy
  • Misersky, J., Majid, A., & Snijders, T. M. (2019). Grammatical gender in German influences how role-nouns are interpreted: Evidence from ERPs. Discourse Processes, 56(8), 643-654. doi:10.1080/0163853X.2018.1541382.

    Abstract

    Grammatically masculine role-nouns (e.g., Studenten-masc.‘students’) can refer to men and women, but may favor an interpretation where only men are considered the referent. If true, this has implications for a society aiming to achieve equal representation in the workplace since, for example, job adverts use such role descriptions. To investigate the interpretation of role-nouns, the present ERP study assessed grammatical gender processing in German. Twenty participants read sentences where a role-noun (masculine or feminine) introduced a group of people, followed by a congruent (masculine–men, feminine–women) or incongruent (masculine–women, feminine–men) continuation. Both for feminine-men and masculine-women continuations a P600 (500 to 800 ms) was observed; another positivity was already present from 300 to 500 ms for feminine-men continuations, but critically not for masculine-women continuations. The results imply a male-biased rather than gender-neutral interpretation of the masculine—despite widespread usage of the masculine as a gender-neutral form—suggesting masculine forms are inadequate for representing genders equally.
  • Mongelli, V., Meijs, E. L., Van Gaal, S., & Hagoort, P. (2019). No language unification without neural feedback: How awareness affects sentence processing. Neuroimage, 202: 116063. doi:10.1016/j.neuroimage.2019.116063.

    Abstract

    How does the human brain combine a finite number of words to form an infinite variety of sentences? According to the Memory, Unification and Control (MUC) model, sentence processing requires long-range feedback from the left inferior frontal cortex (LIFC) to left posterior temporal cortex (LPTC). Single word processing however may only require feedforward propagation of semantic information from sensory regions to LPTC. Here we tested the claim that long-range feedback is required for sentence processing by reducing visual awareness of words using a masking technique. Masking disrupts feedback processing while leaving feedforward processing relatively intact. Previous studies have shown that masked single words still elicit an N400 ERP effect, a neural signature of semantic incongruency. However, whether multiple words can be combined to form a sentence under reduced levels of awareness is controversial. To investigate this issue, we performed two experiments in which we measured electroencephalography (EEG) while 40 subjects performed a masked priming task. Words were presented either successively or simultaneously, thereby forming a short sentence that could be congruent or incongruent with a target picture. This sentence condition was compared with a typical single word condition. In the masked condition we only found an N400 effect for single words, whereas in the unmasked condition we observed an N400 effect for both unmasked sentences and single words. Our findings suggest that long-range feedback processing is required for sentence processing, but not for single word processing.
  • Nieuwland, M. S., Coopmans, C. W., & Sommers, R. P. (2019). Distinguishing old from new referents during discourse comprehension: Evidence from ERPs and oscillations. Frontiers in Human Neuroscience, 13: 398. doi:10.3389/fnhum.2019.00398.

    Abstract

    In this EEG study, we used pre-registered and exploratory ERP and time-frequency analyses to investigate the resolution of anaphoric and non-anaphoric noun phrases during discourse comprehension. Participants listened to story contexts that described two antecedents, and subsequently read a target sentence with a critical noun phrase that lexically matched one antecedent (‘old’), matched two antecedents (‘ambiguous’), partially matched one antecedent in terms of semantic features (‘partial-match’), or introduced another referent (non-anaphoric, ‘new’). After each target sentence, participants judged whether the noun referred back to an antecedent (i.e., an ‘old/new’ judgment), which was easiest for ambiguous nouns and hardest for partially matching nouns. The noun-elicited N400 ERP component demonstrated initial sensitivity to repetition and semantic overlap, corresponding to repetition and semantic priming effects, respectively. New and partially matching nouns both elicited a subsequent frontal positivity, which suggested that partially matching anaphors may have been processed as new nouns temporarily. ERPs in an even later time window and ERPs time-locked to sentence-final words suggested that new and partially matching nouns had different effects on comprehension, with partially matching nouns incurring additional processing costs up to the end of the sentence. In contrast to the ERP results, the time-frequency results primarily demonstrated sensitivity to noun repetition, and did not differentiate partially matching anaphors from new nouns. In sum, our results show the ERP and time-frequency effects of referent repetition during discourse comprehension, and demonstrate the potentially demanding nature of establishing the anaphoric meaning of a novel noun.
  • Nieuwland, M. S. (2019). Do ‘early’ brain responses reveal word form prediction during language comprehension? A critical review. Neuroscience and Biobehavioral Reviews, 96, 367-400. doi:10.1016/j.neubiorev.2018.11.019.

    Abstract

    Current theories of language comprehension posit that readers and listeners routinely try to predict the meaning but also the visual or sound form of upcoming words. Whereas
    most neuroimaging studies on word rediction focus on the N400 ERP or its magnetic equivalent, various studies claim that word form prediction manifests itself in ‘early’, pre
    N400 brain responses (e.g., ELAN, M100, P130, N1, P2, N200/PMN, N250). Modulations of these components are often taken as evidence that word form prediction impacts early sensory processes (the sensory hypothesis) or, alternatively, the initial stages of word recognition before word meaning is integrated with sentence context (the recognition hypothesis). Here, I
    comprehensively review studies on sentence- or discourse-level language comprehension that report such effects of prediction on early brain responses. I conclude that the reported evidence for the sensory hypothesis or word recognition hypothesis is weak and inconsistent,
    and highlight the urgent need for replication of previous findings. I discuss the implications and challenges to current theories of linguistic prediction and suggest avenues for future research.
  • Ostarek, M., Van Paridon, J., & Montero-Melis, G. (2019). Sighted people’s language is not helpful for blind individuals’ acquisition of typical animal colors. Proceedings of the National Academy of Sciences of the United States of America, 116(44), 21972-21973. doi:10.1073/pnas.1912302116.
  • Peeters, D., Vanlangendonck, F., Rüschemeyer, S.-A., & Dijkstra, T. (2019). Activation of the language control network in bilingual visual word recognition. Cortex, 111, 63-73. doi:10.1016/j.cortex.2018.10.012.

    Abstract

    Research into bilingual language production has identified a language control network that subserves control operations when bilinguals produce speech. Here we explore which brain areas are recruited for control purposes in bilingual language comprehension. In two experimental fMRI sessions, Dutch-English unbalanced bilinguals read words that differed in cross-linguistic form and meaning overlap across their two languages. The need for control operations was further manipulated by varying stimulus list composition across the two experimental sessions. We observed activation of the language control network in bilingual language comprehension as a function of both cross-linguistic form and meaning overlap and stimulus list composition. These findings suggest that the language control network is shared across bilingual language production and comprehension. We argue that activation of the language control network in language comprehension allows bilinguals to quickly and efficiently grasp the context-relevant meaning of words.

    Additional information

    1-s2.0-S0010945218303459-mmc1.docx
  • Peeters, D. (2019). Virtual reality: A game-changing method for the language sciences. Psychonomic Bulletin & Review, 26(3), 894-900. doi:10.3758/s13423-019-01571-3.

    Abstract

    This paper introduces virtual reality as an experimental method for the language sciences and provides a review of recent studies using the method to answer fundamental, psycholinguistic research questions. It is argued that virtual reality demonstrates that ecological validity and
    experimental control should not be conceived of as two extremes on a continuum, but rather as two orthogonal factors. Benefits of using virtual reality as an experimental method include that in a virtual environment, as in the real world, there is no artificial spatial divide between participant and stimulus. Moreover, virtual reality experiments do not necessarily have to include a repetitive trial structure or an unnatural experimental task. Virtual agents outperform experimental confederates in terms of the consistency and replicability of their behaviour, allowing for reproducible science across participants and research labs. The main promise of virtual reality as a tool for the experimental language sciences, however, is that it shifts theoretical focus towards the interplay between different modalities (e.g., speech, gesture, eye gaze, facial expressions) in dynamic and communicative real-world environments, complementing studies that focus on one modality (e.g. speech) in isolation.
  • Preisig, B., Sjerps, M. J., Kösem, A., & Riecke, L. (2019). Dual-site high-density 4Hz transcranial alternating current stimulation applied over auditory and motor cortical speech areas does not influence auditory-motor mapping. Brain Stimulation, 12(3), 775-777. doi:10.1016/j.brs.2019.01.007.
  • Preisig, B., & Sjerps, M. J. (2019). Hemispheric specializations affect interhemispheric speech sound integration during duplex perception. The Journal of the Acoustical Society of America, 145, EL190-EL196. doi:10.1121/1.5092829.

    Abstract

    The present study investigated whether speech-related spectral information benefits from initially predominant right or left hemisphere processing. Normal hearing individuals categorized speech sounds composed of an ambiguous base (perceptually intermediate between /ga/ and /da/), presented to one ear, and a disambiguating low or high F3 chirp presented to the other ear. Shorter response times were found when the chirp was presented to the left ear than to the right ear (inducing initially right-hemisphere chirp processing), but no between-ear differences in strength of overall integration. The results are in line with the assumptions of a right hemispheric dominance for spectral processing.

    Additional information

    Supplementary material
  • Sakarias, M., & Flecken, M. (2019). Keeping the result in sight and mind: General cognitive principles and language-specific influences in the perception and memory of resultative events. Cognitive Science, 43(1), 1-30. doi:10.1111/cogs.12708.

    Abstract

    We study how people attend to and memorize endings of events that differ in the degree to which objects in them are affected by an action: Resultative events show objects that undergo a visually salient change in state during the course of the event (peeling a potato), and non‐resultative events involve objects that undergo no, or only partial state change (stirring in a pan). We investigate general cognitive principles, and potential language‐specific influences, in verbal and nonverbal event encoding and memory, across two experiments with Dutch and Estonian participants. Estonian marks a viewer's perspective on an event's result obligatorily via grammatical case on direct object nouns: Objects undergoing a partial/full change in state in an event are marked with partitive/accusative case, respectively. Therefore, we hypothesized increased saliency of object states and event results in Estonian speakers, as compared to speakers of Dutch. Findings show (a) a general cognitive principle of attending carefully to endings of resultative events, implying cognitive saliency of object states in event processing; (b) a language‐specific boost on attention and memory of event results under verbal task demands in Estonian speakers. Results are discussed in relation to theories of event cognition, linguistic relativity, and thinking for speaking.
  • Schoffelen, J.-M., Oostenveld, R., Lam, N. H. L., Udden, J., Hulten, A., & Hagoort, P. (2019). A 204-subject multimodal neuroimaging dataset to study language processing. Scientific Data, 6(1): 17. doi:10.1038/s41597-019-0020-y.

    Abstract

    This dataset, colloquially known as the Mother Of Unification Studies (MOUS) dataset, contains multimodal neuroimaging data that has been acquired from 204 healthy human subjects. The neuroimaging protocol consisted of magnetic resonance imaging (MRI) to derive information at high spatial resolution about brain anatomy and structural connections, and functional data during task, and at rest. In addition, magnetoencephalography (MEG) was used to obtain high temporal resolution electrophysiological measurements during task, and at rest. All subjects performed a language task, during which they processed linguistic utterances that either consisted of normal or scrambled sentences. Half of the subjects were reading the stimuli, the other half listened to the stimuli. The resting state measurements consisted of 5 minutes eyes-open for the MEG and 7 minutes eyes-closed for fMRI. The neuroimaging data, as well as the information about the experimental events are shared according to the Brain Imaging Data Structure (BIDS) format. This unprecedented neuroimaging language data collection allows for the investigation of various aspects of the neurobiological correlates of language.
  • Schoot, L., Hagoort, P., & Segaert, K. (2019). Stronger syntactic alignment in the presence of an interlocutor. Frontiers in Psychology, 10: 685. doi:10.3389/fpsyg.2019.00685.

    Abstract

    Speakers are influenced by the linguistic context: hearing one syntactic alternative leads to an increased chance that the speaker will repeat this structure in the subsequent utterance (i.e., syntactic priming, or structural persistence). Top-down influences, such as whether a conversation partner (or, interlocutor) is present, may modulate the degree to which syntactic priming occurs. In the current study, we indeed show that the magnitude of syntactic alignment increases when speakers are interacting with an interlocutor as opposed to doing the experiment alone. The structural persistence effect for passive sentences is stronger in the presence of an interlocutor than when no interlocutor is present (i.e., when the participant is primed by a recording). We did not find evidence, however, that a speaker’s syntactic priming magnitude is influenced by the degree of their conversation partner’s priming magnitude. Together, these results support a mediated account of syntactic priming, in which syntactic choices are not only affected by preceding linguistic input, but also by top-down influences, such as the speakers’ communicative intent.
  • Schubotz, L., Ozyurek, A., & Holler, J. (2019). Age-related differences in multimodal recipient design: Younger, but not older adults, adapt speech and co-speech gestures to common ground. Language, Cognition and Neuroscience, 34(2), 254-271. doi:10.1080/23273798.2018.1527377.

    Abstract

    Speakers can adapt their speech and co-speech gestures based on knowledge shared with an addressee (common ground-based recipient design). Here, we investigate whether these adaptations are modulated by the speaker’s age and cognitive abilities. Younger and older participants narrated six short comic stories to a same-aged addressee. Half of each story was known to both participants, the other half only to the speaker. The two age groups did not differ in terms of the number of words and narrative events mentioned per narration, or in terms of gesture frequency, gesture rate, or percentage of events expressed multimodally. However, only the younger participants reduced the amount of verbal and gestural information when narrating mutually known as opposed to novel story content. Age-related differences in cognitive abilities did not predict these differences in common ground-based recipient design. The older participants’ communicative behaviour may therefore also reflect differences in social or pragmatic goals.

    Additional information

    plcp_a_1527377_sm4510.pdf
  • Seymour, R. A., Rippon, G., Goordin-Williams, G., Schoffelen, J.-M., & Kessler, K. (2019). Dysregulated oscillatory connectivity in thevisual system in autism spectrum disorder. Brain, 142(10), 3294-3305. doi:10.1093/brain/awz214.

    Abstract

    Autism spectrum disorder is increasingly associated with atypical perceptual and sensory symptoms. Here we explore the hypothesis
    that aberrant sensory processing in autism spectrum disorder could be linked to atypical intra- (local) and interregional (global)
    brain connectivity. To elucidate oscillatory dynamics and connectivity in the visual domain we used magnetoencephalography and
    a simple visual grating paradigm with a group of 18 adolescent autistic participants and 18 typically developing control subjects.
    Both groups showed similar increases in gamma (40–80 Hz) and decreases in alpha (8–13 Hz) frequency power in occipital cortex.
    However, systematic group differences emerged when analysing intra- and interregional connectivity in detail. First, directed
    connectivity was estimated using non-parametric Granger causality between visual areas V1 and V4. Feedforward V1-to-V4
    connectivity, mediated by gamma oscillations, was equivalent between autism spectrum disorder and control groups, but importantly,
    feedback V4-to-V1 connectivity, mediated by alpha (8–13 Hz) oscillations, was significantly reduced in the autism spectrum
    disorder group. This reduction was positively correlated with autistic quotient scores, consistent with an atypical visual hierarchy
    in autism, characterized by reduced top-down modulation of visual input via alpha-band oscillations. Second, at the local level in
    V1, coupling of alpha-phase to gamma amplitude (alpha-gamma phase amplitude coupling) was reduced in the autism spectrum
    disorder group. This implies dysregulated local visual processing, with gamma oscillations decoupled from patterns of wider alphaband
    phase synchrony (i.e. reduced phase amplitude coupling), possibly due to an excitation-inhibition imbalance. More generally,
    these results are in agreement with predictive coding accounts of neurotypical perception and indicate that visual processes in
    autism are less modulated by contextual feedback information.
  • Sharoh, D., Van Mourik, T., Bains, L. J., Segaert, K., Weber, K., Hagoort, P., & Norris, D. (2019). Laminar specific fMRI reveals directed interactions in distributed networks during language processing. Proceedings of the National Academy of Sciences of the United States of America, 116(42), 21185-21190. doi:10.1073/pnas.1907858116.

    Abstract

    Interactions between top-down and bottom-up information streams are integral to brain function but challenging to measure noninvasively. Laminar resolution, functional MRI (lfMRI) is sensitive to depth-dependent properties of the blood oxygen level-dependent (BOLD) response, which can be potentially related to top-down and bottom-up signal contributions. In this work, we used lfMRI to dissociate the top-down and bottom-up signal contributions to the left occipitotemporal sulcus (LOTS) during word reading. We further demonstrate that laminar resolution measurements could be used to identify condition-specific distributed networks on the basis of whole-brain connectivity patterns specific to the depth-dependent BOLD signal. The networks corresponded to top-down and bottom-up signal pathways targeting the LOTS during word reading. We show that reading increased the top-down BOLD signal observed in the deep layers of the LOTS and that this signal uniquely related to the BOLD response in other language-critical regions. These results demonstrate that lfMRI can reveal important patterns of activation that are obscured at standard resolution. In addition to differences in activation strength as a function of depth, we also show meaningful differences in the interaction between signals originating from different depths both within a region and with the rest of the brain. We thus show that lfMRI allows the noninvasive measurement of directed interaction between brain regions and is capable of resolving different connectivity patterns at submillimeter resolution, something previously considered to be exclusively in the domain of invasive recordings.
  • Sjerps, M. J., Fox, N. P., Johnson, K., & Chang, E. F. (2019). Speaker-normalized sound representations in the human auditory cortex. Nature Communications, 10: 2465. doi:10.1038/s41467-019-10365-z.

    Abstract

    The acoustic dimensions that distinguish speech sounds (like the vowel differences in “boot” and “boat”) also differentiate speakers’ voices. Therefore, listeners must normalize across speakers without losing linguistic information. Past behavioral work suggests an important role for auditory contrast enhancement in normalization: preceding context affects listeners’ perception of subsequent speech sounds. Here, using intracranial electrocorticography in humans, we investigate whether and how such context effects arise in auditory cortex. Participants identified speech sounds that were preceded by phrases from two different speakers whose voices differed along the same acoustic dimension as target words (the lowest resonance of the vocal tract). In every participant, target vowels evoke a speaker-dependent neural response that is consistent with the listener’s perception, and which follows from a contrast enhancement model. Auditory cortex processing thus displays a critical feature of normalization, allowing listeners to extract meaningful content from the voices of diverse speakers.

    Additional information

    41467_2019_10365_MOESM1_ESM.pdf
  • De Swart, P., & Van Bergen, G. (2019). How animacy and verbal information influence V2 sentence processing: Evidence from eye movements. Open Linguistics, 5(1), 630-649. doi:10.1515/opli-2019-0035.

    Abstract

    There exists a clear association between animacy and the grammatical function of transitive subject. The grammar of some languages require the transitive subject to be high in animacy, or at least higher than the object. A similar animacy preference has been observed in processing studies in languages without such a categorical animacy effect. This animacy preference has been mainly established in structures in which either one or both arguments are provided before the verb. Our goal was to establish (i) whether this preference can already be observed before any argument is provided, and (ii) whether this preference is mediated by verbal information. To this end we exploited the V2 property of Dutch which allows the verb to precede its arguments. Using a visual-world eye-tracking paradigm we presented participants with V2 structures with either an auxiliary (e.g. Gisteren heeft X … ‘Yesterday, X has …’) or a lexical main verb (e.g. Gisteren motiveerde X … ‘Yesterday, X motivated …’) and we measured looks to the animate referent. The results indicate that the animacy preference can already be observed before arguments are presented and that the selectional restrictions of the verb mediate this bias, but do not override it completely.
  • Takashima, A., Bakker-Marshall, I., Van Hell, J. G., McQueen, J. M., & Janzen, G. (2019). Neural correlates of word learning in children. Developmental Cognitive Neuroscience, 37: 100647. doi:10.1016/j.dcn.2019.100649.

    Abstract

    Memory representations of words are thought to undergo changes with consolidation: Episodic memories of novel words are transformed into lexical representations that interact with other words in the mental dictionary. Behavioral studies have shown that this lexical integration process is enhanced when there is more time for consolidation. Neuroimaging studies have further revealed that novel word representations are initially represented in a hippocampally-centered system, whereas left posterior middle temporal cortex activation increases with lexicalization. In this study, we measured behavioral and brain responses to newly-learned words in children. Two groups of Dutch children, aged between 8-10 and 14-16 years, were trained on 30 novel Japanese words depicting novel concepts. Children were tested on word-forms, word-meanings, and the novel words’ influence on existing word processing immediately after training, and again after a week. In line with the adult findings, hippocampal involvement decreased with time. Lexical integration, however, was not observed immediately or after a week, neither behaviorally nor neurally. It appears that time alone is not always sufficient for lexical integration to occur. We suggest that other factors (e.g., the novelty of the concepts and familiarity with the language the words are derived from) might also influence the integration process.

    Additional information

    Supplementary data
  • Takashima, A., & Verhoeven, L. (2019). Radical repetition effects in beginning learners of Chinese as a foreign language reading. Journal of Neurolinguistics, 50, 71-81. doi:10.1016/j.jneuroling.2018.03.001.

    Abstract

    The aim of the present study was to examine whether repetition of radicals during training of Chinese characters leads to better word acquisition performance in beginning learners of Chinese as a foreign language. Thirty Dutch university students were trained on 36 Chinese one-character words for their pronunciations and meanings. They were also exposed to the specifics of the radicals, that is, for phonetic radicals, the associated pronunciation was explained, and for semantic radicals the associated categorical meanings were explained. Results showed that repeated exposure to phonetic and semantic radicals through character pronunciation and meaning trainings indeed induced better understanding of those radicals that were shared among different characters. Furthermore, characters in the training set that shared phonetic radicals were pronounced better than those that did not. Repetition of semantic radicals across different characters, however, hindered the learning of exact meanings. Students generally confused the meanings of other characters that shared the semantic radical. The study shows that in the initial stage of learning, overlapping information of the shared radicals are effectively learned. Acquisition of the specifics of individual characters, however, requires more training.

    Additional information

    Supplementary data
  • Udden, J., Hulten, A., Bendt, K., Mineroff, Z., Kucera, K. S., Vino, A., Fedorenko, E., Hagoort, P., & Fisher, S. E. (2019). Towards robust functional neuroimaging genetics of cognition. Journal of Neuroscience, 39(44), 8778-8787. doi:10.1523/JNEUROSCI.0888-19.2019.

    Abstract

    A commonly held assumption in cognitive neuroscience is that, because measures of human brain function are closer to underlying biology than distal indices of behavior/cognition, they hold more promise for uncovering genetic pathways. Supporting this view is an influential fMRI-based study of sentence reading/listening by Pinel et al. (2012), who reported that common DNA variants in specific candidate genes were associated with altered neural activation in language-related regions of healthy individuals that carried them. In particular, different single-nucleotide polymorphisms (SNPs) of FOXP2 correlated with variation in task-based activation in left inferior frontal and precentral gyri, whereas a SNP at the KIAA0319/TTRAP/THEM2 locus was associated with variable functional asymmetry of the superior temporal sulcus. Here, we directly test each claim using a closely matched neuroimaging genetics approach in independent cohorts comprising 427 participants, four times larger than the original study of 94 participants. Despite demonstrating power to detect associations with substantially smaller effect sizes than those of the original report, we do not replicate any of the reported associations. Moreover, formal Bayesian analyses reveal substantial to strong evidence in support of the null hypothesis (no effect). We highlight key aspects of the original investigation, common to functional neuroimaging genetics studies, which could have yielded elevated false-positive rates. Genetic accounts of individual differences in cognitive functional neuroimaging are likely to be as complex as behavioral/cognitive tests, involving many common genetic variants, each of tiny effect. Reliable identification of true biological signals requires large sample sizes, power calculations, and validation in independent cohorts with equivalent paradigms.

    SIGNIFICANCE STATEMENT A pervasive idea in neuroscience is that neuroimaging-based measures of brain function, being closer to underlying neurobiology, are more amenable for uncovering links to genetics. This is a core assumption of prominent studies that associate common DNA variants with altered activations in task-based fMRI, despite using samples (10–100 people) that lack power for detecting the tiny effect sizes typical of genetically complex traits. Here, we test central findings from one of the most influential prior studies. Using matching paradigms and substantially larger samples, coupled to power calculations and formal Bayesian statistics, our data strongly refute the original findings. We demonstrate that neuroimaging genetics with task-based fMRI should be subject to the same rigorous standards as studies of other complex traits.
  • Van den Broek, G. S. E., Segers, E., Van Rijn, H., Takashima, A., & Verhoeven, L. (2019). Effects of elaborate feedback during practice tests: Costs and benefits of retrieval prompts. Journal of Experimental Psychology: Applied, 25(4), 588-601. doi:10.1037/xap0000212.

    Abstract

    This study explores the effect of feedback with hints on students’ recall of words. In three classroom experiments, high school students individually practiced vocabulary words through computerized retrieval practice with either standard show-answer feedback (display of answer) or hints feedback after incorrect responses. Hints feedback gave students a second chance to find the correct response using orthographic (Experiment 1), mnemonic (Experiment 2), or cross-language hints (Experiment 3). During practice, hints led to a shift of practice time from further repetitions to longer feedback processing but did not reduce (repeated) errors. There was no effect of feedback on later recall except when the hints from practice were also available on the test, indicating limited transfer of practice with hints to later recall without hints (in Experiments 1 and 2). Overall, hints feedback was not preferable over show-answer feedback. The common notion that hints are beneficial may not hold when the total practice time is limited.
  • Van Bergen, G., Flecken, M., & Wu, R. (2019). Rapid target selection of object categories based on verbs: Implications for language-categorization interactions. Psychophysiology, 56(9): e13395. doi:10.1111/psyp.13395.

    Abstract

    Although much is known about how nouns facilitate object categorization, very little is known about how verbs (e.g., posture verbs such as stand or lie) facilitate object categorization. Native Dutch speakers are a unique population to investigate this issue with because the configurational categories distinguished by staan (to stand) and liggen (to lie) are inherent in everyday Dutch language. Using an ERP component (N2pc), four experiments demonstrate that selection of posture verb categories is rapid (between 220–320 ms). The effect was attenuated, though present, when removing the perceptual distinction between categories. A similar attenuated effect was obtained in native English speakers, where the category distinction is less familiar, and when category labels were implicit for native Dutch speakers. Our results are among the first to demonstrate that category search based on verbs can be rapid, although extensive linguistic experience and explicit labels may not be necessary to facilitate categorization in this case.

    Additional information

    psyp13395-sup-0001-appendixs1.pdf
  • Van Es, M. W. J., & Schoffelen, J.-M. (2019). Stimulus-induced gamma power predicts the amplitude of the subsequent visual evoked response. NeuroImage, 186, 703-712. doi:10.1016/j.neuroimage.2018.11.029.

    Abstract

    The efficiency of neuronal information transfer in activated brain networks may affect behavioral performance.
    Gamma-band synchronization has been proposed to be a mechanism that facilitates neuronal processing of
    behaviorally relevant stimuli. In line with this, it has been shown that strong gamma-band activity in visual
    cortical areas leads to faster responses to a visual go cue. We investigated whether there are directly observable
    consequences of trial-by-trial fluctuations in non-invasively observed gamma-band activity on the neuronal
    response. Specifically, we hypothesized that the amplitude of the visual evoked response to a go cue can be
    predicted by gamma power in the visual system, in the window preceding the evoked response. Thirty-three
    human subjects (22 female) performed a visual speeded response task while their magnetoencephalogram
    (MEG) was recorded. The participants had to respond to a pattern reversal of a concentric moving grating. We
    estimated single trial stimulus-induced visual cortical gamma power, and correlated this with the estimated single
    trial amplitude of the most prominent event-related field (ERF) peak within the first 100 ms after the pattern
    reversal. In parieto-occipital cortical areas, the amplitude of the ERF correlated positively with gamma power, and
    correlated negatively with reaction times. No effects were observed for the alpha and beta frequency bands,
    despite clear stimulus onset induced modulation at those frequencies. These results support a mechanistic model,
    in which gamma-band synchronization enhances the neuronal gain to relevant visual input, thus leading to more
    efficient downstream processing and to faster responses.
  • Varma, S., Takashima, A., Fu, L., & Kessels, R. P. C. (2019). Mindwandering propensity modulates episodic memory consolidation. Aging Clinical and Experimental Research, 31(11), 1601-1607. doi:10.1007/s40520-019-01251-1.

    Abstract

    Research into strategies that can combat episodic memory decline in healthy older adults has gained widespread attention over the years. Evidence suggests that a short period of rest immediately after learning can enhance memory consolidation, as compared to engaging in cognitive tasks. However, a recent study in younger adults has shown that post-encoding engagement in a working memory task leads to the same degree of memory consolidation as from post-encoding rest. Here, we tested whether this finding can be extended to older adults. Using a delayed recognition test, we compared the memory consolidation of word–picture pairs learned prior to 9 min of rest or a 2-Back working memory task, and examined its relationship with executive functioning and mindwandering propensity. Our results show that (1) similar to younger adults, memory for the word–picture associations did not differ when encoding was followed by post-encoding rest or 2-Back task and (2) older adults with higher mindwandering propensity retained more word–picture associations encoded prior to rest relative to those encoded prior to the 2-Back task, whereas participants with lower mindwandering propensity had better memory performance for the pairs encoded prior to the 2-Back task. Overall, our results indicate that the degree of episodic memory consolidation during both active and passive post-encoding periods depends on individual mindwandering tendency.

    Additional information

    Supplementary material
  • Warren, C. M., Tona, K. D., Ouwekerk, L., Van Paridon, J., Poletiek, F. H., Bosch, J. A., & Nieuwenhuis, S. (2019). The neuromodulatory and hormonal effects of transcutaneous vagus nerve stimulation as evidenced by salivary alpha amylase, salivary cortisol, pupil diameter, and the P3 event-related potential. Brain Stimulation, 12(3), 635-642. doi:10.1016/j.brs.2018.12.224.

    Abstract

    Background

    Transcutaneous vagus nerve stimulation (tVNS) is a new, non-invasive technique being investigated as an intervention for a variety of clinical disorders, including epilepsy and depression. It is thought to exert its therapeutic effect by increasing central norepinephrine (NE) activity, but the evidence supporting this notion is limited.
    Objective

    In order to test for an impact of tVNS on psychophysiological and hormonal indices of noradrenergic function, we applied tVNS in concert with assessment of salivary alpha amylase (SAA) and cortisol, pupil size, and electroencephalograph (EEG) recordings.
    Methods

    Across three experiments, we applied real and sham tVNS to 61 healthy participants while they performed a set of simple stimulus-discrimination tasks. Before and after the task, as well as during one break, participants provided saliva samples and had their pupil size recorded. EEG was recorded throughout the task. The target for tVNS was the cymba conchae, which is heavily innervated by the auricular branch of the vagus nerve. Sham stimulation was applied to the ear lobe.
    Results

    P3 amplitude was not affected by tVNS (Experiment 1A: N=24; Experiment 1B: N=20; Bayes factor supporting null model=4.53), nor was pupil size (Experiment 2: N=16; interaction of treatment and time: p=0.79). However, tVNS increased SAA (Experiments 1A and 2: N=25) and attenuated the decline of salivary cortisol compared to sham (Experiment 2: N=17), as indicated by significant interactions involving treatment and time (p=.023 and p=.040, respectively).
    Conclusion

    These findings suggest that tVNS modulates hormonal indices but not psychophysiological indices of noradrenergic function.
  • Weber, K., Christiansen, M., Indefrey, P., & Hagoort, P. (2019). Primed from the start: Syntactic priming during the first days of language learning. Language Learning, 69(1), 198-221. doi:10.1111/lang.12327.

    Abstract

    New linguistic information must be integrated into our existing language system. Using a novel experimental task that incorporates a syntactic priming paradigm into artificial language learning, we investigated how new grammatical regularities and words are learned. This innovation allowed us to control the language input the learner received, while the syntactic priming paradigm provided insight into the nature of the underlying syntactic processing machinery. The results of the present study pointed to facilitatory syntactic processing effects within the first days of learning: Syntactic and lexical priming effects revealed participants’ sensitivity to both novel words and word orders. This suggested that novel syntactic structures and their meaning (form–function mapping) can be acquired rapidly through incidental learning. More generally, our study indicated similar mechanisms for learning and processing in both artificial and natural languages, with implications for the relationship between first and second language learning.
  • Weber, K., Micheli, C., Ruigendijk, E., & Rieger, J. (2019). Sentence processing is modulated by the current linguistic environment and a priori information: An fMRI study. Brain and Behavior, 9(7): e01308. doi:10.1002/brb3.1308.

    Abstract

    Introduction
    Words are not processed in isolation but in rich contexts that are used to modulate and facilitate language comprehension. Here, we investigate distinct neural networks underlying two types of contexts, the current linguistic environment and verb‐based syntactic preferences.

    Methods
    We had two main manipulations. The first was the current linguistic environment, where the relative frequencies of two syntactic structures (prepositional object [PO] and double‐object [DO]) would either follow everyday linguistic experience or not. The second concerned the preference toward one or the other structure depending on the verb; learned in everyday language use and stored in memory. German participants were reading PO and DO sentences in German while brain activity was measured with functional magnetic resonance imaging.

    Results
    First, the anterior cingulate cortex (ACC) showed a pattern of activation that integrated the current linguistic environment with everyday linguistic experience. When the input did not match everyday experience, the unexpected frequent structure showed higher activation in the ACC than the other conditions and more connectivity from the ACC to posterior parts of the language network. Second, verb‐based surprisal of seeing a structure given a verb (PO verb preference but DO structure presentation) resulted, within the language network (left inferior frontal and left middle/superior temporal gyrus) and the precuneus, in increased activation compared to a predictable verb‐structure pairing.

    Conclusion
    In conclusion, (1) beyond the canonical language network, brain areas engaged in prediction and error signaling, such as the ACC, might use the statistics of syntactic structures to modulate language processing, (2) the language network is directly engaged in processing verb preferences. These two networks show distinct influences on sentence processing.

    Additional information

    Supporting information
  • Zhu, Z., Bastiaansen, M. C. M., Hakun, J. G., Petersson, K. M., Wang, S., & Hagoort, P. (2019). Semantic unification modulates N400 and BOLD signal change in the brain: A simultaneous EEG-fMRI study. Journal of Neurolinguistics, 52: 100855. doi:10.1016/j.jneuroling.2019.100855.

    Abstract

    Semantic unification during sentence comprehension has been associated with amplitude change of the N400 in event-related potential (ERP) studies, and activation in the left inferior frontal gyrus (IFG) in functional magnetic resonance imaging (fMRI) studies. However, the specificity of this activation to semantic unification remains unknown. To more closely examine the brain processes involved in semantic unification, we employed simultaneous EEG-fMRI to time-lock the semantic unification related N400 change, and integrated trial-by-trial variation in both N400 and BOLD change beyond the condition-level BOLD change difference measured in traditional fMRI analyses. Participants read sentences in which semantic unification load was parametrically manipulated by varying cloze probability. Separately, ERP and fMRI results replicated previous findings, in that semantic unification load parametrically modulated the amplitude of N400 and cortical activation. Integrated EEG-fMRI analyses revealed a different pattern in which functional activity in the left IFG and bilateral supramarginal gyrus (SMG) was associated with N400 amplitude, with the left IFG activation and bilateral SMG activation being selective to the condition-level and trial-level of semantic unification load, respectively. By employing the EEG-fMRI integrated analyses, this study among the first sheds light on how to integrate trial-level variation in language comprehension.
  • Araújo, S., Faísca, L., Bramão, I., Reis, A., & Petersson, K. M. (2015). Lexical and sublexical orthographic processing: An ERP study with skilled and dyslexic adult readers. Brain and Language, 141, 16-27. doi:10.1016/j.bandl.2014.11.007.

    Abstract

    This ERP study investigated the cognitive nature of the P1–N1 components during orthographic processing. We used an implicit reading task with various types of stimuli involving different amounts of sublexical or lexical orthographic processing (words, pseudohomophones, pseudowords, nonwords, and symbols), and tested average and dyslexic readers. An orthographic regularity effect (pseudowords– nonwords contrast) was observed in the average but not in the dyslexic group. This suggests an early sensitivity to the dependencies among letters in word-forms that reflect orthographic structure, while the dyslexic brain apparently fails to be appropriately sensitive to these complex features. Moreover, in the adults the N1-response may already reflect lexical access: (i) the N1 was sensitive to the familiar vs. less familiar orthographic sequence contrast; (ii) and early effects of the phonological form (words-pseudohomophones contrast) were also found. Finally, the later N320 component was attenuated in the dyslexics, suggesting suboptimal processing in later stages of phonological analysis.
  • Araújo, S., Reis, A., Petersson, K. M., & Faísca, L. (2015). Rapid automatized naming and reading performance: A meta-analysis. Journal of Educational Psychology, 107(3), 868-883. doi:10.1037/edu0000006.

    Abstract

    Evidence that rapid naming skill is associated with reading ability has become increasingly prevalent in recent years. However, there is considerable variation in the literature concerning the magnitude of this relationship. The objective of the present study was to provide a comprehensive analysis of the evidence on the relationship between rapid automatized naming (RAN) and reading performance. To this end, we conducted a meta-analysis of the correlational relationship between these 2 constructs to (a) determine the overall strength of the RAN–reading association and (b) identify variables that systematically moderate this relationship. A random-effects model analysis of data from 137 studies (857 effect sizes; 28,826 participants) indicated a moderate-to-strong relationship between RAN and reading performance (r = .43, I2 = 68.40). Further analyses revealed that RAN contributes to the 4 measures of reading (word reading, text reading, non-word reading, and reading comprehension), but higher coefficients emerged in favor of real word reading and text reading. RAN stimulus type and type of reading score were the factors with the greatest moderator effect on the magnitude of the RAN–reading relationship. The consistency of orthography and the subjects’ grade level were also found to impact this relationship, although the effect was contingent on reading outcome. It was less evident whether the subjects’ reading proficiency played a role in the relationship. Implications for future studies are discussed.
  • Asaridou, S. S., Hagoort, P., & McQueen, J. M. (2015). Effects of early bilingual experience with a tone and a non-tone language on speech-music. PLoS One, 10(12): e0144225. doi:10.1371/journal.pone.0144225.

    Abstract

    We investigated music and language processing in a group of early bilinguals who spoke a tone language and a non-tone language (Cantonese and Dutch). We assessed online speech-music processing interactions, that is, interactions that occur when speech and music are processed simultaneously in songs, with a speeded classification task. In this task, participants judged sung pseudowords either musically (based on the direction of the musical interval) or phonologically (based on the identity of the sung vowel). We also assessed longer-term effects of linguistic experience on musical ability, that is, the influence of extensive prior experience with language when processing music. These effects were assessed with a task in which participants had to learn to identify musical intervals and with four pitch-perception tasks. Our hypothesis was that due to their experience in two different languages using lexical versus intonational tone, the early Cantonese-Dutch bilinguals would outperform the Dutch control participants. In online processing, the Cantonese-Dutch bilinguals processed speech and music more holistically than controls. This effect seems to be driven by experience with a tone language, in which integration of segmental and pitch information is fundamental. Regarding longer-term effects of linguistic experience, we found no evidence for a bilingual advantage in either the music-interval learning task or the pitch-perception tasks. Together, these results suggest that being a Cantonese-Dutch bilingual does not have any measurable longer-term effects on pitch and music processing, but does have consequences for how speech and music are processed jointly.

    Additional information

    Data Availability
  • Baggio, G., van Lambalgen, M., & Hagoort, P. (2015). Logic as Marr's computational level: Four case studies. Topics in Cognitive Science, 7, 287-298. doi:10.1111/tops.12125.

    Abstract

    We sketch four applications of Marr's levels-of-analysis methodology to the relations between logic and experimental data in the cognitive neuroscience of language and reasoning. The first part of the paper illustrates the explanatory power of computational level theories based on logic. We show that a Bayesian treatment of the suppression task in reasoning with conditionals is ruled out by EEG data, supporting instead an analysis based on defeasible logic. Further, we describe how results from an EEG study on temporal prepositions can be reanalyzed using formal semantics, addressing a potential confound. The second part of the article demonstrates the predictive power of logical theories drawing on EEG data on processing progressive constructions and on behavioral data on conditional reasoning in people with autism. Logical theories can constrain processing hypotheses all the way down to neurophysiology, and conversely neuroscience data can guide the selection of alternative computational level models of cognition.
  • Bakker, I., Takashima, A., van Hell, J. G., Janzen, G., & McQueen, J. M. (2015). Tracking lexical consolidation with ERPs: Lexical and semantic-priming effects on N400 and LPC responses to newly-learned words. Neuropsychologia, 79, 33-41. doi:10.1016/j.neuropsychologia.2015.10.020.
  • Bašnákova, J., Van Berkum, J. J. A., Weber, K., & Hagoort, P. (2015). A job interview in the MRI scanner: How does indirectness affect addressees and overhearers? Neuropsychologia, 76, 79-91. doi:10.1016/j.neuropsychologia.2015.03.030.

    Abstract

    In using language, people not only exchange information, but also navigate their social world – for example, they can express themselves indirectly to avoid losing face. In this functional magnetic resonance imaging study, we investigated the neural correlates of interpreting face-saving indirect replies, in a situation where participants only overheard the replies as part of a conversation between two other people, as well as in a situation where the participants were directly addressed themselves. We created a fictional job interview context where indirect replies serve as a natural communicative strategy to attenuate one’s shortcomings, and asked fMRI participants to either pose scripted questions and receive answers from three putative job candidates (addressee condition) or to listen to someone else interview the same candidates (overhearer condition). In both cases, the need to evaluate the candidate ensured that participants had an active interest in comprehending the replies. Relative to direct replies, face-saving indirect replies increased activation in medial prefrontal cortex, bilateral temporo-parietal junction (TPJ), bilateral inferior frontal gyrus and bilateral middle temporal gyrus, in active overhearers and active addressees alike, with similar effect size, and comparable to findings obtained in an earlier passive listening study (Bašnáková et al., 2013). In contrast, indirectness effects in bilateral anterior insula and pregenual ACC, two regions implicated in emotional salience and empathy, were reliably stronger in addressees than in active overhearers. Our findings indicate that understanding face-saving indirect language requires additional cognitive perspective-taking and other discourse-relevant cognitive processing, to a comparable extent in active overhearers and addressees. Furthermore, they indicate that face-saving indirect language draws upon affective systems more in addressees than in overhearers, presumably because the addressee is the one being managed by a face-saving reply. In all, face-saving indirectness provides a window on the cognitive as well as affect-related neural systems involved in human communication.
  • Bastiaansen, M. C. M., & Hagoort, P. (2015). Frequency-based segregation of syntactic and semantic unification during online sentence level language comprehension. Journal of Cognitive Neuroscience, 27(11), 2095-2107. doi:10.1162/jocn_a_00829.

    Abstract

    During sentence level language comprehension, semantic and syntactic unification are functionally distinct operations. Nevertheless, both recruit roughly the same brain areas (spatially overlapping networks in the left frontotemporal cortex) and happen at the same time (in the first few hundred milliseconds after word onset). We tested the hypothesis that semantic and syntactic unification are segregated by means of neuronal synchronization of the functionally relevant networks in different frequency ranges: gamma (40 Hz and up) for semantic unification and lower beta (10–20 Hz) for syntactic unification. EEG power changes were quantified as participants read either correct sentences, syntactically correct though meaningless sentences (syntactic prose), or sentences that did not contain any syntactic structure (random word lists). Other sentences contained either a semantic anomaly or a syntactic violation at a critical word in the sentence. Larger EEG gamma-band power was observed for semantically coherent than for semantically anomalous sentences. Similarly, beta-band power was larger for syntactically correct sentences than for incorrect ones. These results confirm the existence of a functional dissociation in EEG oscillatory dynamics during sentence level language comprehension that is compatible with the notion of a frequency-based segregation of syntactic and semantic unification.
  • Bastos, A. M., Vezoli, J., Bosman, C. A., Schoffelen, J.-M., Oostenveld, R., Dowdall, J. R., De Weerd, P., Kennedy, H., & Fries, P. (2015). Visual areas exert feedforward and feedback influences through distinct frequency channels. Neuron, 85(2), 390-401. doi:10.1016/j.neuron.2014.12.018.

    Abstract

    Visual cortical areas subserve cognitive functions by interacting in both feedforward and feedback directions. While feedforward influences convey sensory signals, feedback influences modulate feedforward signaling according to the current behavioral context. We investigated whether these interareal influences are subserved differentially by rhythmic synchronization. We correlated frequency-specific directed influences among 28 pairs of visual areas with anatomical metrics of the feedforward or feedback character of the respective interareal projections. This revealed that in the primate visual system, feedforward influences are carried by theta-band ( approximately 4 Hz) and gamma-band ( approximately 60-80 Hz) synchronization, and feedback influences by beta-band ( approximately 14-18 Hz) synchronization. The functional directed influences constrain a functional hierarchy similar to the anatomical hierarchy, but exhibiting task-dependent dynamic changes in particular with regard to the hierarchical positions of frontal areas. Our results demonstrate that feedforward and feedback signaling use distinct frequency channels, suggesting that they subserve differential communication requirements.
  • Chang, F., Bauman, M., Pappert, S., & Fitz, H. (2015). Do lemmas speak German?: A verb position effect in German structural priming. Cognitive Science, 39(5), 1113-1130. doi:10.1111/cogs.12184.

    Abstract

    Lexicalized theories of syntax often assume that verb-structure regularities are mediated by lemmas, which abstract over variation in verb tense and aspect. German syntax seems to challenge this assumption, because verb position depends on tense and aspect. To examine how German speakers link these elements, a structural priming study was performed which varied syntactic structure, verb position (encoded by tense and aspect), and verb overlap. Abstract structural priming was found, both within and across verb position, but priming was larger when the verb position was the same between prime and target. Priming was boosted by verb overlap, but there was no interaction with verb position. The results can be explained by a lemma model where tense and aspect are linked to structural choices in German. Since the architecture of this lemma model is not consistent with results from English, a connectionist model was developed which could explain the cross-linguistic variation in the production system. Together, these findings support the view that language learning plays an important role in determining the nature of structural priming in different languages
  • Cronin, K. A., Acheson, D. J., Hernández, P., & Sánchez, A. (2015). Hierarchy is Detrimental for Human Cooperation. Scientific Reports, 5: 18634. doi:10.1038/srep18634.

    Abstract

    Studies of animal behavior consistently demonstrate that the social environment impacts cooperation, yet the effect of social dynamics has been largely excluded from studies of human cooperation. Here, we introduce a novel approach inspired by nonhuman primate research to address how social hierarchies impact human cooperation. Participants competed to earn hierarchy positions and then could cooperate with another individual in the hierarchy by investing in a common effort. Cooperation was achieved if the combined investments exceeded a threshold, and the higher ranked individual distributed the spoils unless control was contested by the partner. Compared to a condition lacking hierarchy, cooperation declined in the presence of a hierarchy due to a decrease in investment by lower ranked individuals. Furthermore, hierarchy was detrimental to cooperation regardless of whether it was earned or arbitrary. These findings mirror results from nonhuman primates and demonstrate that hierarchies are detrimental to cooperation. However, these results deviate from nonhuman primate findings by demonstrating that human behavior is responsive to changing hierarchical structures and suggests partnership dynamics that may improve cooperation. This work introduces a controlled way to investigate the social influences on human behavior, and demonstrates the evolutionary continuity of human behavior with other primate species.
  • Flecken, M., Carroll, M., Weimar, K., & Von Stutterheim, C. (2015). Driving along the road or heading for the village? Conceptual differences underlying motion event encoding in French, German, and French-German L2 users. Modern Language Journal, 99(S1), 100-122. doi:10.1111/j.1540-4781.2015.12181.x.

    Abstract

    The typological contrast between verb- and satellite-framed languages (Talmy, 1985) has set the basis for many empirical studies on L2 acquisition. The current analysis goes beyond this typology by looking in detail at the conceptualization of the path of motion in a motion event. We take as a starting point the cognitive salience of specific elements of motion events that are relevant when conceptualizing space. When expressing direction in French, specific spatial relations involving the entity in motion (its alignment and its distance toward a [potential] endpoint) are relevant, given a variety of path verbs in the lexicon expressing this information (e.g., se diriger vers, avancer to direct oneself toward,' to advance'). This is not the case in German (manner verbs in the lexicon mainly). In German, spatial information is packaged in adjuncts and particles and the path of motion is typically structured via features of the ground (entlanglaufen/fahren to walk/drive along') or endpoints (to walk/drive to/toward'). We investigate those fundamental differences in spatial conceptualization in French and German, as reflected in pre-articulatory patterns of attention allocation (measured with eye tracking) to moving entities and endpoints in motion scenes in an event description task. Our focus is on spatial conceptualization in an L2 (French L2 users of German), analyzing the extent to which these L2 users display target-like patterns or traces of L1 conceptualization transfer. Findings show that, in line with directional concepts expressed in verbs, L1 French speakers allocate more attention to entities in motion and endpoints, before utterance onset, than L1 German speakers do. The L2 German speakers pattern with L1 German speakers in the use of manner verbs, but they have not fully acquired the spatial concepts and means that structure the path of motion in the L2. This is reflected in pre-articulatory attention allocation patterns, according to which the L2 speakers pattern with native speakers of their L1 (French). The findings show a continued deep entrenchment of L1-based processing patterns and spatial frames of reference when speakers prepare for speech in an L2
  • Flecken, M., Walbert, K., & Dijkstra, T. (2015). ‘Right now, Sophie ∗swims in the pool?!’: Brain potentials of grammatical aspect processing. Frontiers in Psychology, 6: 1764. doi:10.3389/fpsyg.2015.01764.

    Abstract

    We investigated whether brain potentials of grammatical aspect processing resemble semantic or morpho-syntactic processing, or whether they instead are characterized by an entirely distinct pattern in the same individuals. We studied aspect from the perspective of agreement between the temporal information in the context (temporal adverbials, e.g., Right now) and a morpho-syntactic marker of grammatical aspect (e.g., progressive is swimming). Participants read questions providing a temporal context that was progressive (What is Sophie doing in the pool right now?) or habitual (What does Sophie do in the pool every Monday?). Following a lead-in sentence context such as Right now, Sophie…, we measured event-related brain potentials (ERPs) time-locked to verb phrases in four different conditions, e.g., (a) is swimming (control); (b) ∗is cooking (semantic violation); (c) ∗are swimming (morpho-syntactic violation); or (d)?swims (aspect mismatch); …in the pool.” The collected ERPs show typical N400 and P600 effects for semantics and morpho-syntax, while aspect processing elicited an Early Negativity (250–350 ms). The aspect-related Negativity was short-lived and had a central scalp distribution with an anterior onset. This differentiates it not only from the semantic N400 effect, but also from the typical LAN (Left Anterior Negativity), that is frequently reported for various types of agreement processing. Moreover, aspect processing did not show a clear P600 modulation. We argue that the specific context for each item in this experiment provided a trigger for agreement checking with temporal information encoded on the verb, i.e., morphological aspect marking. The aspect-related Negativity obtained for aspect agreement mismatches reflects a violated expectation concerning verbal inflection (in the example above, the expected verb phrase was Sophie is X-ing rather than Sophie X-s in condition d). The absence of an additional P600 for aspect processing suggests that the mismatch did not require additional reintegration or processing costs. This is consistent with participants’ post hoc grammaticality judgements of the same sentences, which overall show a high acceptability of aspect mismatch sentences.

    Additional information

    data sheet 1.docx
  • Flecken, M., Athanasopoulos, P., Kuipers, J. R., & Thierry, G. (2015). On the road to somewhere: Brain potentials reflect language effects on motion event perception. Cognition, 141, 41-51. doi:10.1016/j.cognition.2015.04.006.

    Abstract

    Recent studies have identified neural correlates of language effects on perception in static domains of experience such as colour and objects. The generalization of such effects to dynamic domains like motion events remains elusive. Here, we focus on grammatical differences between languages relevant for the description of motion events and their impact on visual scene perception. Two groups of native speakers of German or English were presented with animated videos featuring a dot travelling along a trajectory towards a geometrical shape (endpoint). English is a language with grammatical aspect in which attention is drawn to trajectory and endpoint of motion events equally. German, in contrast, is a non-aspect language which highlights endpoints. We tested the comparative perceptual saliency of trajectory and endpoint of motion events by presenting motion event animations (primes) followed by a picture symbolising the event (target): In 75% of trials, the animation was followed by a mismatching picture (both trajectory and endpoint were different); in 10% of trials, only the trajectory depicted in the picture matched the prime; in 10% of trials, only the endpoint matched the prime; and in 5% of trials both trajectory and endpoint were matching, which was the condition requiring a response from the participant. In Experiment 1 we recorded event-related brain potentials elicited by the picture in native speakers of German and native speakers of English. German participants exhibited a larger P3 wave in the endpoint match than the trajectory match condition, whereas English speakers showed no P3 amplitude difference between conditions. In Experiment 2 participants performed a behavioural motion matching task using the same stimuli as those used in Experiment 1. German and English participants did not differ in response times showing that motion event verbalisation cannot readily account for the difference in P3 amplitude found in the first experiment. We argue that, even in a non-verbal context, the grammatical properties of the native language and associated sentence-level patterns of event encoding influence motion event perception, such that attention is automatically drawn towards aspects highlighted by the grammar.
  • Francken, J. C., Meijs, E. L., Ridderinkhof, O. M., Hagoort, P., de Lange, F. P., & van Gaal, S. (2015). Manipulating word awareness dissociates feed-forward from feedback models of language-perception interactions. Neuroscience of consciousness, 1. doi:10.1093/nc/niv003.

    Abstract

    Previous studies suggest that linguistic material can modulate visual perception, but it is unclear at which level of processing these interactions occur. Here we aim to dissociate between two competing models of language–perception interactions: a feed-forward and a feedback model. We capitalized on the fact that the models make different predictions on the role of feedback. We presented unmasked (aware) or masked (unaware) words implying motion (e.g. “rise,” “fall”), directly preceding an upward or downward visual motion stimulus. Crucially, masking leaves intact feed-forward information processing from low- to high-level regions, whereas it abolishes subsequent feedback. Under this condition, participants remained faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. This suggests that language–perception interactions are driven by the feed-forward convergence of linguistic and perceptual information at higher-level conceptual and decision stages.
  • Francken, J. C., Meijs, E. L., Hagoort, P., van Gaal, S., & de Lange, F. P. (2015). Exploring the automaticity of language-perception interactions: Effects of attention and awareness. Scientific Reports, 5: 17725. doi:10.1038/srep17725.

    Abstract

    Previous studies have shown that language can modulate visual perception, by biasing and/
    or enhancing perceptual performance. However, it is still debated where in the brain visual and
    linguistic information are integrated, and whether the effects of language on perception are
    automatic and persist even in the absence of awareness of the linguistic material. Here, we aimed
    to explore the automaticity of language-perception interactions and the neural loci of these
    interactions in an fMRI study. Participants engaged in a visual motion discrimination task (upward
    or downward moving dots). Before each trial, a word prime was briefly presented that implied
    upward or downward motion (e.g., “rise”, “fall”). These word primes strongly influenced behavior:
    congruent motion words sped up reaction times and improved performance relative to incongruent
    motion words. Neural congruency effects were only observed in the left middle temporal gyrus,
    showing higher activity for congruent compared to incongruent conditions. This suggests that higherlevel
    conceptual areas rather than sensory areas are the locus of language-perception interactions.
    When motion words were rendered unaware by means of masking, they still affected visual motion
    perception, suggesting that language-perception interactions may rely on automatic feed-forward
    integration of perceptual and semantic material in language areas of the brain.
  • Francken, J. C., Kok, P., Hagoort, P., & De Lange, F. P. (2015). The behavioral and neural effects of language on motion perception. Journal of Cognitive Neuroscience, 27(1), 175-184. doi:10.1162/jocn_a_00682.

    Abstract

    Perception does not function as an isolated module but is tightly linked with other cognitive functions. Several studies have demonstrated an influence of language on motion perception, but it remains debated at which level of processing this modulation takes place. Some studies argue for an interaction in perceptual areas, but it is also possible that the interaction is mediated by "language areas" that integrate linguistic and visual information. Here, we investigated whether language-perception interactions were specific to the language-dominant left hemisphere by comparing the effects of language on visual material presented in the right (RVF) and left visual fields (LVF). Furthermore, we determined the neural locus of the interaction using fMRI. Participants performed a visual motion detection task. On each trial, the visual motion stimulus was presented in either the LVF or in the RVF, preceded by a centrally presented word (e.g., "rise"). The word could be congruent, incongruent, or neutral with regard to the direction of the visual motion stimulus that was presented subsequently. Participants were faster and more accurate when the direction implied by the motion word was congruent with the direction of the visual motion stimulus. Interestingly, the speed benefit was present only for motion stimuli that were presented in the RVF. We observed a neural counterpart of the behavioral facilitation effects in the left middle temporal gyrus, an area involved in semantic processing of verbal material. Together, our results suggest that semantic information about motion retrieved in language regions may automatically modulate perceptual decisions about motion.
  • Franken, M. K., Hagoort, P., & Acheson, D. J. (2015). Modulations of the auditory M100 in an Imitation Task. Brain and Language, 142, 18-23. doi:10.1016/j.bandl.2015.01.001.

    Abstract

    Models of speech production explain event-related suppression of the auditory cortical
    response as reflecting a comparison between auditory predictions and feedback. The present MEG
    study was designed to test two predictions from this framework: 1) whether the reduced auditory
    response varies as a function of the mismatch between prediction and feedback; 2) whether individual
    variation in this response is predictive of speech-motor adaptation.
    Participants alternated between online imitation and listening tasks. In the imitation task, participants
    began each trial producing the same vowel (/e/) and subsequently listened to and imitated auditorilypresented
    vowels varying in acoustic distance from /e/.
    Results replicated suppression, with a smaller M100 during speaking than listening. Although we did
    not find unequivocal support for the first prediction, participants with less M100 suppression were
    better at the imitation task. These results are consistent with the enhancement of M100 serving as an
    error signal to drive subsequent speech-motor adaptation.
  • Guadalupe, T., Zwiers, M. P., Wittfeld, K., Teumer, A., Vasquez, A. A., Hoogman, M., Hagoort, P., Fernandez, G., Buitelaar, J., van Bokhoven, H., Hegenscheid, K., Völzke, H., Franke, B., Fisher, S. E., Grabe, H. J., & Francks, C. (2015). Asymmetry within and around the human planum temporale is sexually dimorphic and influenced by genes involved in steroid hormone receptor activity. Cortex, 62, 41-55. doi:10.1016/j.cortex.2014.07.015.

    Abstract

    The genetic determinants of cerebral asymmetries are unknown. Sex differences in asymmetry of the planum temporale, that overlaps Wernicke’s classical language area, have been inconsistently reported. Meta-analysis of previous studies has suggested that publication bias established this sex difference in the literature. Using probabilistic definitions of cortical regions we screened over the cerebral cortex for sexual dimorphisms of asymmetry in 2337 healthy subjects, and found the planum temporale to show the strongest sex-linked asymmetry of all regions, which was supported by two further datasets, and also by analysis with the Freesurfer package that performs automated parcellation of cerebral cortical regions. We performed a genome-wide association scan meta-analysis of planum temporale asymmetry in a pooled sample of 3095 subjects, followed by a candidate-driven approach which measured a significant enrichment of association in genes of the ´steroid hormone receptor activity´ and 'steroid metabolic process' pathways. Variants in the genes and pathways identified may affect the role of the planum temporale in language cognition.
  • Holler, J., Kokal, I., Toni, I., Hagoort, P., Kelly, S. D., & Ozyurek, A. (2015). Eye’m talking to you: Speakers’ gaze direction modulates co-speech gesture processing in the right MTG. Social Cognitive & Affective Neuroscience, 10, 255-261. doi:10.1093/scan/nsu047.

    Abstract

    Recipients process information from speech and co-speech gestures, but it is currently unknown how this processing is influenced by the presence of other important social cues, especially gaze direction, a marker of communicative intent. Such cues may modulate neural activity in regions associated either with the processing of ostensive cues, such as eye gaze, or with the processing of semantic information, provided by speech and gesture.
    Participants were scanned (fMRI) while taking part in triadic communication involving two recipients and a speaker. The speaker uttered sentences that
    were and were not accompanied by complementary iconic gestures. Crucially, the speaker alternated her gaze direction, thus creating two recipient roles: addressed (direct gaze) vs unaddressed (averted gaze) recipient. The comprehension of Speech&Gesture relative to SpeechOnly utterances recruited middle occipital, middle temporal and inferior frontal gyri, bilaterally. The calcarine sulcus and posterior cingulate cortex were sensitive to differences between direct and averted gaze. Most importantly, Speech&Gesture utterances, but not SpeechOnly utterances, produced additional activity in the right middle temporal gyrus when participants were addressed. Marking communicative intent with gaze direction modulates the processing of speech–gesture utterances in cerebral areas typically associated with the semantic processing of multi-modal communicative acts.
  • Horschig, J. M., Smolders, R., Bonnefond, M., Schoffelen, J.-M., Van den Munckhof, P., Schuurman, P. R., Cools, R., Denys, D., & Jensen, O. (2015). Directed communication between nucleus accumbens and neocortex in humans is differentially supported by synchronization in the theta and alpha band. PLoS One, 10(9): e0138685. doi:10.1371/journal.pone.0138685.

    Abstract

    Here, we report evidence for oscillatory bi-directional interactions between the nucleus accumbens and the neocortex in humans. Six patients performed a demanding covert visual attention task while we simultaneously recorded brain activity from deep-brain electrodes implanted in the nucleus accumbens and the surface electroencephalogram (EEG). Both theta and alpha oscillations were strongly coherent with the frontal and parietal EEG during the task. Theta-band coherence increased during processing of the visual stimuli. Granger causality analysis revealed that the nucleus accumbens was communicating with the neocortex primarily in the theta-band, while the cortex was communicating the nucleus accumbens in the alpha-band. These data are consistent with a model, in which theta- and alpha-band oscillations serve dissociable roles: Prior to stimulus processing, the cortex might suppress ongoing processing in the nucleus accumbens by modulating alpha-band activity. Subsequently, upon stimulus presentation, theta oscillations might facilitate the active exchange of stimulus information from the nucleus accumbens to the cortex.
  • Jiang, J., Chen, C., Dai, B., Shi, G., Liu, L., & Lu, C. (2015). Leader emergence through interpersonal neural synchronization. Proceedings of the National Academy of Sciences of the United States of America, 112(14), 4274-4279. doi:10.1073/pnas.1422930112.

    Abstract

    The neural mechanism of leader emergence is not well understood. This study investigated (i) whether interpersonal neural synchronization (INS) plays an important role in leader emergence, and (ii) whether INS and leader emergence are associated with the frequency or the quality of communications. Eleven three-member groups were asked to perform a leaderless group discussion (LGD) task, and their brain activities were recorded via functional near infrared spectroscopy (fNIRS)-based hyperscanning. Video recordings of the discussions were coded for leadership and communication. Results showed that the INS for the leader–follower (LF) pairs was higher than that for the follower–follower (FF) pairs in the left temporo-parietal junction (TPJ), an area important for social mentalizing. Although communication frequency was higher for the LF pairs than for the FF pairs, the frequency of leader-initiated and follower-initiated communication did not differ significantly. Moreover, INS for the LF pairs was significantly higher during leader-initiated communication than during follower-initiated communications. In addition, INS for the LF pairs during leader-initiated communication was significantly correlated with the leaders’ communication skills and competence, but not their communication frequency. Finally, leadership could be successfully predicted based on INS as well as communication frequency early during the LGD (before half a minute into the task). In sum, this study found that leader emergence was characterized by high-level neural synchronization between the leader and followers and that the quality, rather than the frequency, of communications was associated with synchronization. These results suggest that leaders emerge because they are able to say the right things at the right time.
  • Kunert, R., & Slevc, L. R. (2015). A commentary on: “Neural overlap in processing music and speech”. Frontiers in Human Neuroscience, 9: 330. doi:10.3389/fnhum.2015.00330.
  • Kunert, R., Willems, R. M., Casasanto, D., Patel, A. D., & Hagoort, P. (2015). Music and language syntax interact in Broca’s Area: An fMRI study. PLoS One, 10(11): e0141069. doi:10.1371/journal.pone.0141069.

    Abstract

    Instrumental music and language are both syntactic systems, employing complex, hierarchically-structured sequences built using implicit structural norms. This organization allows listeners to understand the role of individual words or tones in the context of an unfolding sentence or melody. Previous studies suggest that the brain mechanisms of syntactic processing may be partly shared between music and language. However, functional neuroimaging evidence for anatomical overlap of brain activity involved in linguistic and musical syntactic processing has been lacking. In the present study we used functional magnetic resonance imaging (fMRI) in conjunction with an interference paradigm based on sung sentences. We show that the processing demands of musical syntax (harmony) and language syntax interact in Broca’s area in the left inferior frontal gyrus (without leading to music and language main effects). A language main effect in Broca’s area only emerged in the complex music harmony condition, suggesting that (with our stimuli and tasks) a language effect only becomes visible under conditions of increased demands on shared neural resources. In contrast to previous studies, our design allows us to rule out that the observed neural interaction is due to: (1) general attention mechanisms, as a psychoacoustic auditory anomaly behaved unlike the harmonic manipulation, (2) error processing, as the language and the music stimuli contained no structural errors. The current results thus suggest that two different cognitive domains—music and language—might draw on the same high level syntactic integration resources in Broca’s area.
  • Lai, V. T., & Curran, T. (2015). Erratum to “ERP evidence for conceptual mappings and comparison processes during the comprehension of conventional and novel metaphors” [Brain Lang. 127 (3) (2013) 484–496]. Brain and Language, 149, 148-150. doi:10.1016/j.bandl.2014.11.001.
  • Lai, V. T., Willems, R. M., & Hagoort, P. (2015). Feel between the Lines: Implied emotion from combinatorial semantics. Journal of Cognitive Neuroscience, 27(8), 1528-1541. doi:10.1162/jocn_a_00798.

    Abstract

    This study investigated the brain regions for the comprehension of implied emotion in sentences. Participants read negative sentences without negative words, for example, “The boy fell asleep and never woke up again,” and their neutral counterparts “The boy stood up and grabbed his bag.” This kind of negative sentence allows us to examine implied emotion derived at the sentence level, without associative emotion coming from word retrieval. We found that implied emotion in sentences, relative to neutral sentences, led to activation in some emotion-related areas, including the medial prefrontal cortex, the amygdala, and the insula, as well as certain language-related areas, including the inferior frontal gyrus, which has been implicated in combinatorial processing. These results suggest that the emotional network involved in implied emotion is intricately related to the network for combinatorial processing in language, supporting the view that sentence meaning is more than simply concatenating the meanings of its lexical building blocks.
  • Lartseva, A., Dijkstra, T., & Buitelaar, J. (2015). Emotional language processing in Autism Spectrum Disorders: A systematic review. Frontiers in Human Neuroscience, 8: 991. doi:10.3389/fnhum.2014.00991.

    Abstract

    In his first description of Autism Spectrum Disorders (ASD), Kanner emphasized emotional impairments by characterizing children with ASD as indifferent to other people, self-absorbed, emotionally cold, distanced, and retracted. Thereafter, emotional impairments became regarded as part of the social impairments of ASD, and research mostly focused on understanding how individuals with ASD recognize visual expressions of emotions from faces and body postures. However, it still remains unclear how emotions are processed outside of the visual domain. This systematic review aims to fill this gap by focusing on impairments of emotional language processing in ASD.
    We systematically searched PubMed for papers published between 1990 and 2013 using standardized search terms. Studies show that people with ASD are able to correctly classify emotional language stimuli as emotionally positive or negative. However, processing of emotional language stimuli in ASD is associated with atypical patterns of attention and memory performance, as well as abnormal physiological and neural activity. Particularly, younger children with ASD have difficulties in acquiring and developing emotional concepts, and avoid using these in discourse. These emotional language impairments were not consistently associated with age, IQ, or level of development of language skills.
    We discuss how emotional language impairments fit with existing cognitive theories of ASD, such as central coherence, executive dysfunction, and weak Theory of Mind. We conclude that emotional impairments in ASD may be broader than just a mere consequence of social impairments, and should receive more attention in future research
  • Lewis, A. G., & Bastiaansen, M. C. M. (2015). A predictive coding framework for rapid neural dynamics during sentence-level language comprehension. Cortex, 68, 155-168. doi:10.1016/j.cortex.2015.02.014.

    Abstract

    There is a growing literature investigating the relationship between oscillatory neural dynamics measured using EEG and/or MEG, and sentence-level language comprehension. Recent proposals have suggested a strong link between predictive coding accounts of the hierarchical flow of information in the brain, and oscillatory neural dynamics in the beta and gamma frequency ranges. We propose that findings relating beta and gamma oscillations to sentence-level language comprehension might be unified under such a predictive coding account. Our suggestion is that oscillatory activity in the beta frequency range may reflect both the active maintenance of the current network configuration responsible for representing the sentence-level meaning under construction, and the top-down propagation of predictions to hierarchically lower processing levels based on that representation. In addition, we suggest that oscillatory activity in the low and middle gamma range reflect the matching of top-down predictions with bottom-up linguistic input, while evoked high gamma might reflect the propagation of bottom-up prediction errors to higher levels of the processing hierarchy. We also discuss some of the implications of this predictive coding framework, and we outline ideas for how these might be tested experimentally
  • Lewis, A. G., Wang, L., & Bastiaansen, M. C. M. (2015). Fast oscillatory dynamics during language comprehension: Unification versus maintenance and prediction? Brain and Language, 148, 51-63. doi:10.1016/j.bandl.2015.01.003.

    Abstract

    The role of neuronal oscillations during language comprehension is not yet well understood. In this paper we review and reinterpret the functional roles of beta- and gamma-band oscillatory activity during language comprehension at the sentence and discourse level. We discuss the evidence in favor of a role for beta and gamma in unification (the unification hypothesis), and in light of mounting evidence that cannot be accounted for under this hypothesis, we explore an alternative proposal linking beta and gamma oscillations to maintenance and prediction (respectively) during language comprehension. Our maintenance/prediction hypothesis is able to account for most of the findings that are currently available relating beta and gamma oscillations to language comprehension, and is in good agreement with other proposals about the roles of beta and gamma in domain-general cognitive processing. In conclusion we discuss proposals for further testing and comparing the prediction and unification hypotheses.
  • Lockwood, G., & Dingemanse, M. (2015). Iconicity in the lab: A review of behavioural, developmental, and neuroimaging research into sound-symbolism. Frontiers in Psychology, 6: 1246. doi:10.3389/fpsyg.2015.01246.

    Abstract

    This review covers experimental approaches to sound-symbolism—from infants to adults, and from Sapir’s foundational studies to twenty-first century product naming. It synthesizes recent behavioral, developmental, and neuroimaging work into a systematic overview of the cross-modal correspondences that underpin iconic links between form and meaning. It also identifies open questions and opportunities, showing how the future course of experimental iconicity research can benefit from an integrated interdisciplinary perspective. Combining insights from psychology and neuroscience with evidence from natural languages provides us with opportunities for the experimental investigation of the role of sound-symbolism in language learning, language processing, and communication. The review finishes by describing how hypothesis-testing and model-building will help contribute to a cumulative science of sound-symbolism in human language.
  • Lockwood, G., & Tuomainen, J. (2015). Ideophones in Japanese modulate the P2 and late positive complex responses. Frontiers in Psychology, 6: 933. doi:10.3389/fpsyg.2015.00933.

    Abstract

    Sound-symbolism, or the direct link between sound and meaning, is typologically and behaviorally attested across languages. However, neuroimaging research has mostly focused on artificial non-words or individual segments, which do not represent sound-symbolism in natural language. We used EEG to compare Japanese ideophones, which are phonologically distinctive sound-symbolic lexical words, and arbitrary adverbs during a sentence reading task. Ideophones elicit a larger visual P2 response and a sustained late positive complex in comparison to arbitrary adverbs. These results and previous literature suggest that the larger P2 may indicate the integration of sound and sensory information by association in response to the distinctive phonology of ideophones. The late positive complex may reflect the facilitated lexical retrieval of ideophones in comparison to arbitrary words. This account provides new evidence that ideophones exhibit similar cross-modal correspondences to those which have been proposed for non-words and individual sounds, and that these effects are detectable in natural language.
  • Moreno, I., De Vega, M., León, I., Bastiaansen, M. C. M., Lewis, A. G., & Magyari, L. (2015). Brain dynamics in the comprehension of action-related language. A time-frequency analysis of mu rhythms. Neuroimage, 109, 50-62. doi:10.1016/j.neuroimage.2015.01.018.

    Abstract

    EEG mu rhythms (8-13Hz) recorded at fronto-central electrodes are generally considered as markers of motor cortical activity in humans, because they are modulated when participants perform an action, when they observe another’s action or even when they imagine performing an action. In this study, we analyzed the time-frequency (TF) modulation of mu rhythms while participants read action language (“You will cut the strawberry cake”), abstract language (“You will doubt the patient´s argument”), and perceptive language (“You will notice the bright day”). The results indicated that mu suppression at fronto-central sites is associated with action language rather than with abstract or perceptive language. Also, the largest difference between conditions occurred quite late in the sentence, while reading the first noun, (contrast Action vs. Abstract), or the second noun following the action verb (contrast Action vs. Perceptive). This suggests that motor activation is associated with the integration of words across the sentence beyond the lexical processing of the action verb. Source reconstruction localized mu suppression associated with action sentences in premotor cortex (BA 6). The present study suggests (1) that the understanding of action language activates motor networks in the human brain, and (2) that this activation occurs online based on semantic integration across multiple words in the sentence.
  • Nijhoff, A. D., & Willems, R. M. (2015). Simulating fiction: Individual differences in literature comprehension revealed with fMRI. PLoS One, 10(2): e0116492. doi:10.1371/journal.pone.0116492.

    Abstract

    When we read literary fiction, we are transported to fictional places, and we feel and think along with the characters. Despite the importance of narrative in adult life and during development, the neurocognitive mechanisms underlying fiction comprehension are unclear. We used functional magnetic resonance imaging (fMRI) to investigate how individuals differently employ neural networks important for understanding others’ beliefs and intentions (mentalizing), and for sensori-motor simulation while listening to excerpts from literary novels. Localizer tasks were used to localize both the cortical motor network and the mentalizing network in participants after they listened to excerpts from literary novels. Results show that participants who had high activation in anterior medial prefrontal cortex (aMPFC; part of the mentalizing network) when listening to mentalizing content of literary fiction, had lower motor cortex activity when they listened to action-related content of the story, and vice versa. This qualifies how people differ in their engagement with fiction: some people are mostly drawn into a story by mentalizing about the thoughts and beliefs of others, whereas others engage in literature by simulating more concrete events such as actions. This study provides on-line neural evidence for the existence of qualitatively different styles of moving into literary worlds, and adds to a growing body of literature showing the potential to study narrative comprehension with neuroimaging methods.
  • Peeters, D., Chu, M., Holler, J., Hagoort, P., & Ozyurek, A. (2015). Electrophysiological and kinematic correlates of communicative intent in the planning and production of pointing gestures and speech. Journal of Cognitive Neuroscience, 27(12), 2352-2368. doi:10.1162/jocn_a_00865.

    Abstract

    In everyday human communication, we often express our communicative intentions by manually pointing out referents in the material world around us to an addressee, often in tight synchronization with referential speech. This study investigated whether and how the kinematic form of index finger pointing gestures is shaped by the gesturer's communicative intentions and how this is modulated by the presence of concurrently produced speech. Furthermore, we explored the neural mechanisms underpinning the planning of communicative pointing gestures and speech. Two experiments were carried out in which participants pointed at referents for an addressee while the informativeness of their gestures and speech was varied. Kinematic and electrophysiological data were recorded online. It was found that participants prolonged the duration of the stroke and poststroke hold phase of their gesture to be more communicative, in particular when the gesture was carrying the main informational burden in their multimodal utterance. Frontal and P300 effects in the ERPs suggested the importance of intentional and modality-independent attentional mechanisms during the planning phase of informative pointing gestures. These findings contribute to a better understanding of the complex interplay between action, attention, intention, and language in the production of pointing gestures, a communicative act core to human interaction.
  • Peeters, D., Hagoort, P., & Ozyurek, A. (2015). Electrophysiological evidence for the role of shared space in online comprehension of spatial demonstratives. Cognition, 136, 64-84. doi:10.1016/j.cognition.2014.10.010.

    Abstract

    A fundamental property of language is that it can be used to refer to entities in the extra-linguistic physical context of a conversation in order to establish a joint focus of attention on a referent. Typological and psycholinguistic work across a wide range of languages has put forward at least two different theoretical views on demonstrative reference. Here we contrasted and tested these two accounts by investigating the electrophysiological brain activity underlying the construction of indexical meaning in comprehension. In two EEG experiments, participants watched pictures of a speaker who referred to one of two objects using speech and an index-finger pointing gesture. In contrast with separately collected native speakers’ linguistic intuitions, N400 effects showed a preference for a proximal demonstrative when speaker and addressee were in a face-to-face orientation and all possible referents were located in the shared space between them, irrespective of the physical proximity of the referent to the speaker. These findings reject egocentric proximity-based accounts of demonstrative reference, support a sociocentric approach to deixis, suggest that interlocutors construe a shared space during conversation, and imply that the psychological proximity of a referent may be more important than its physical proximity.
  • Samur, D., Lai, V. T., Hagoort, P., & Willems, R. M. (2015). Emotional context modulates embodied metaphor comprehension. Neuropsychologia, 78, 108-114. doi:10.1016/j.neuropsychologia.2015.10.003.

    Abstract

    Emotions are often expressed metaphorically, and both emotion and metaphor are ways through which abstract meaning can be grounded in language. Here we investigate specifically whether motion-related verbs when used metaphorically are differentially sensitive to a preceding emotional context, as compared to when they are used in a literal manner. Participants read stories that ended with ambiguous action/motion sentences (e.g., he got it), in which the action/motion could be interpreted metaphorically (he understood the idea) or literally (he caught the ball) depending on the preceding story. Orthogonal to the metaphorical manipulation, the stories were high or low in emotional content. The results showed that emotional context modulated the neural response in visual motion areas to the metaphorical interpretation of the sentences, but not to their literal interpretations. In addition, literal interpretations of the target sentences led to stronger activation in the visual motion areas as compared to metaphorical readings of the sentences. We interpret our results as suggesting that emotional context specifically modulates mental simulation during metaphor processing
  • Simanova, I., Van Gerven, M. A., Oostenveld, R., & Hagoort, P. (2015). Predicting the semantic category of internally generated words from neuromagnetic recordings. Journal of Cognitive Neuroscience, 27(1), 35-45. doi:10.1162/jocn_a_00690.

    Abstract

    In this study, we explore the possibility to predict the semantic category of words from brain signals in a free word generation task. Participants produced single words from different semantic categories in a modified semantic fluency task. A Bayesian logistic regression classifier was trained to predict the semantic category of words from single-trial MEG data. Significant classification accuracies were achieved using sensor-level MEG time series at the time interval of conceptual preparation. Semantic category prediction was also possible using source-reconstructed time series, based on minimum norm estimates of cortical activity. Brain regions that contributed most to classification on the source level were identified. These were the left inferior frontal gyrus, left middle frontal gyrus, and left posterior middle temporal gyrus. Additionally, the temporal dynamics of brain activity underlying the semantic preparation during word generation was explored. These results provide important insights about central aspects of language production
  • Todorovic, A., Schoffelen, J.-M., van Ede, F., Maris, E., & de Lange, F. P. (2015). Temporal expectation and attention jointly modulate auditory oscillatory activity in the beta band. PLoS One, 10(3): e0120288. doi:10.1371/journal.pone.0120288.

    Abstract

    The neural response to a stimulus is influenced by endogenous factors such as expectation and attention. Current research suggests that expectation and attention exert their effects in opposite directions, where expectation decreases neural activity in sensory areas, while attention increases it. However, expectation and attention are usually studied either in isolation or confounded with each other. A recent study suggests that expectation and attention may act jointly on sensory processing, by increasing the neural response to expected events when they are attended, but decreasing it when they are unattended. Here we test this hypothesis in an auditory temporal cueing paradigm using magnetoencephalography in humans. In our study participants attended to, or away from, tones that could arrive at expected or unexpected moments. We found a decrease in auditory beta band synchrony to expected (versus unexpected) tones if they were unattended, but no difference if they were attended. Modulations in beta power were already evident prior to the expected onset times of the tones. These findings suggest that expectation and attention jointly modulate sensory processing.
  • Van den Bos, E., & Poletiek, F. H. (2015). Learning simple and complex artificial grammars in the presence of a semantic reference field: Effects on performance and awareness. Frontiers in Psychology, 6: 158. doi:10.3389/fpsyg.2015.00158.

    Abstract

    This study investigated whether the negative effect of complexity on artificial grammar learning could be compensated by adding semantics. Participants were exposed to exemplars from a simple or a complex finite state grammar presented with or without a semantic reference field. As expected, performance on a grammaticality judgment test was higher for the simple grammar than for the complex grammar. For the simple grammar, the results also showed that participants presented with a reference field and instructed to decode the meaning of each exemplar (decoding condition) did better than participants who memorized the exemplars without semantic referents (memorize condition). Contrary to expectations, however, there was no significant difference between the decoding condition and the memorize condition for the complex grammar. These findings indicated that the negative effect of complexity remained, despite the addition of semantics. To clarify how the presence of a reference field influenced the learning process, its effects on the acquisition of two types of knowledge (first- and second-order dependencies) and on participants’ awareness of their knowledge were examined. The results tentatively suggested that the reference field enhanced the learning of second-order dependencies. In addition, participants in the decoding condition realized when they had knowledge relevant to making a grammaticality judgment, whereas participants in the memorize condition demonstrated some knowledge of which they were unaware. These results are in line with the view that the reference field enhanced structure learning by making certain dependencies more salient. Moreover, our findings stress the influence of complexity on artificial grammar learning

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  • Veenstra, A., Meyer, A. S., & Acheson, D. J. (2015). Effects of parallel planning on agreement production. Acta Psychologica, 162, 29-39. doi:10.1016/j.actpsy.2015.09.011.

    Abstract

    An important issue in current psycholinguistics is how the time course of utterance planning affects the generation of grammatical structures. The current study investigated the influence of parallel activation of the components of complex noun phrases on the generation of subject-verb agreement. Specifically, the lexical interference account (Gillespie, M. and Pearlmutter, N. J., 2011b and Solomon, E. S. and Pearlmutter, N. J., 2004) predicts more agreement errors (i.e., attraction) for subject phrases in which the head and local noun mismatch in number (e.g., the apple next to the pears) when nouns are planned in parallel than when they are planned in sequence. We used a speeded picture description task that yielded sentences such as the apple next to the pears is red. The objects mentioned in the noun phrase were either semantically related or unrelated. To induce agreement errors, pictures sometimes mismatched in number. In order to manipulate the likelihood of parallel processing of the objects and to test the hypothesized relationship between parallel processing and the rate of agreement errors, the pictures were either placed close together or far apart. Analyses of the participants' eye movements and speech onset latencies indicated slower processing of the first object and stronger interference from the related (compared to the unrelated) second object in the close than in the far condition. Analyses of the agreement errors yielded an attraction effect, with more errors in mismatching than in matching conditions. However, the magnitude of the attraction effect did not differ across the close and far conditions. Thus, spatial proximity encouraged parallel processing of the pictures, which led to interference of the associated conceptual and/or lexical representation, but, contrary to the prediction, it did not lead to more attraction errors.
  • Wang, L., Bastiaansen, M. C. M., & Yang, Y. (2015). ERP responses to person names as a measure of trait inference in person perception. Social Neuroscience, 10, 89-99. doi:10.1080/17470919.2014.944995.

    Abstract

    Using event-related potentials (ERPs), this study examines how trait information inferred from behaviors is associated with person names. In linguistic discourses, person names were associated with descriptions of either positive or negative behaviors. In a subsequent explicit evaluation task, the previously described person names were presented in isolation, and the participants were asked to judge the emotional valence of these names. We found that the names associated with positive descriptions elicited a larger positivity in the ERP than the names associated with negative descriptions. The results indicate that the emotional valence of person names attached to person perception can be dynamically influenced by short descriptions of the target person, probably due to trait inference based on the provided behavioral descriptions

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  • Xiang, H., Van Leeuwen, T. M., Dediu, D., Roberts, L., Norris, D. G., & Hagoort, P. (2015). L2-proficiency-dependent laterality shift in structural connectivity of brain language pathways. Brain Connectivity, 5(6), 349-361. doi:10.1089/brain.2013.0199.

    Abstract

    Diffusion tensor imaging (DTI) and a longitudinal language learning approach were applied to investigate the relationship between the achieved second language (L2) proficiency during L2 learning and the reorganization of structural connectivity between core language areas. Language proficiency tests and DTI scans were obtained from German students before and after they completed an intensive 6-week course of the Dutch language. In the initial learning stage, with increasing L2 proficiency, the hemispheric dominance of the BA6-temporal pathway (mainly along the arcuate fasciculus) shifted from the left to the right hemisphere. With further increased proficiency, however, lateralization dominance was again found in the left BA6-temporal pathway. This result is consistent with reports in the literature that imply a stronger involvement of the right hemisphere in L2-processing especially for less proficient L2-speakers. This is the first time that a L2-proficiency-dependent laterality shift in structural connectivity of language pathways during L2 acquisition has been observed to shift from left to right, and back to left hemisphere dominance with increasing L2-proficiency. We additionally find that changes in fractional anisotropy values after the course are related to the time elapsed between the two scans. The results suggest that structural connectivity in (at least part of) the perisylvian language network may be subject to fast dynamic changes following language learning
  • Acheson, D. J. (2013). Signatures of response conflict monitoring in language production. Procedia - Social and Behavioral Sciences, 94, 214-215. doi:10.1016/j.sbspro.2013.09.106.
  • Acheson, D. J., & Hagoort, P. (2013). Stimulating the brain's language network: Syntactic ambiguity resolution after TMS to the IFG and MTG. Journal of Cognitive Neuroscience, 25(10), 1664-1677. doi:10.1162/jocn_a_00430.

    Abstract

    The posterior middle temporal gyrus (MTG) and inferior frontal gyrus (IFG) are two critical nodes of the brain's language network. Previous neuroimaging evidence has supported a dissociation in language comprehension in which parts of the MTG are involved in the retrieval of lexical syntactic information and the IFG is involved in unification operations that maintain, select, and integrate multiple sources of information over time. In the present investigation, we tested for causal evidence of this dissociation by modulating activity in IFG and MTG using an offline TMS procedure: continuous theta-burst stimulation. Lexical–syntactic retrieval was manipulated by using sentences with and without a temporarily word-class (noun/verb) ambiguity (e.g., run). In one group of participants, TMS was applied to the IFG and MTG, and in a control group, no TMS was applied. Eye movements were recorded and quantified at two critical sentence regions: a temporarily ambiguous region and a disambiguating region. Results show that stimulation of the IFG led to a modulation of the ambiguity effect (ambiguous–unambiguous) at the disambiguating sentence region in three measures: first fixation durations, total reading times, and regressive eye movements into the region. Both IFG and MTG stimulation modulated the ambiguity effect for total reading times in the temporarily ambiguous sentence region relative to a control group. The current results demonstrate that an offline repetitive TMS protocol can have influences at a different point in time during online processing and provide causal evidence for IFG involvement in unification operations during sentence comprehension.
  • Andics, A., McQueen, J. M., & Petersson, K. M. (2013). Mean-based neural coding of voices. NeuroImage, 79, 351-360. doi:10.1016/j.neuroimage.2013.05.002.

    Abstract

    The social significance of recognizing the person who talks to us is obvious, but the neural mechanisms that mediate talker identification are unclear. Regions along the bilateral superior temporal sulcus (STS) and the inferior frontal cortex (IFC) of the human brain are selective for voices, and they are sensitive to rapid voice changes. Although it has been proposed that voice recognition is supported by prototype-centered voice representations, the involvement of these category-selective cortical regions in the neural coding of such "mean voices" has not previously been demonstrated. Using fMRI in combination with a voice identity learning paradigm, we show that voice-selective regions are involved in the mean-based coding of voice identities. Voice typicality is encoded on a supra-individual level in the right STS along a stimulus-dependent, identity-independent (i.e., voice-acoustic) dimension, and on an intra-individual level in the right IFC along a stimulus-independent, identity-dependent (i.e., voice identity) dimension. Voice recognition therefore entails at least two anatomically separable stages, each characterized by neural mechanisms that reference the central tendencies of voice categories.
  • Asaridou, S. S., & McQueen, J. M. (2013). Speech and music shape the listening brain: Evidence for shared domain-general mechanisms. Frontiers in Psychology, 4: 321. doi:10.3389/fpsyg.2013.00321.

    Abstract

    Are there bi-directional influences between speech perception and music perception? An answer to this question is essential for understanding the extent to which the speech and music that we hear are processed by domain-general auditory processes and/or by distinct neural auditory mechanisms. This review summarizes a large body of behavioral and neuroscientific findings which suggest that the musical experience of trained musicians does modulate speech processing, and a sparser set of data, largely on pitch processing, which suggest in addition that linguistic experience, in particular learning a tone language, modulates music processing. Although research has focused mostly on music on speech effects, we argue that both directions of influence need to be studied, and conclude that the picture which thus emerges is one of mutual interaction across domains. In particular, it is not simply that experience with spoken language has some effects on music perception, and vice versa, but that because of shared domain-general subcortical and cortical networks, experiences in both domains influence behavior in both domains.
  • De Boer, M., Toni, I., & Willems, R. M. (2013). What drives successful verbal communication? Frontiers in Human Neuroscience, 7: 622. doi:10.3389/fnhum.2013.00622.

    Abstract

    There is a vast amount of potential mappings between behaviors and intentions in communication: a behavior can indicate a multitude of different intentions, and the same intention can be communicated with a variety of behaviors. Humans routinely solve these many-to-many referential problems when producing utterances for an Addressee. This ability might rely on social cognitive skills, for instance, the ability to manipulate unobservable summary variables to disambiguate ambiguous behavior of other agents (“mentalizing”) and the drive to invest resources into changing and understanding the mental state of other agents (“communicative motivation”). Alternatively, the ambiguities of verbal communicative interactions might be solved by general-purpose cognitive abilities that process cues that are incidentally associated with the communicative interaction. In this study, we assess these possibilities by testing which cognitive traits account for communicative success during a verbal referential task. Cognitive traits were assessed with psychometric scores quantifying motivation, mentalizing abilities, and general-purpose cognitive abilities, taxing abstract visuo-spatial abilities. Communicative abilities of participants were assessed by using an on-line interactive task that required a speaker to verbally convey a concept to an Addressee. The communicative success of the utterances was quantified by measuring how frequently a number of Evaluators would infer the correct concept. Speakers with high motivational and general-purpose cognitive abilities generated utterances that were more easily interpreted. These findings extend to the domain of verbal communication the notion that motivational and cognitive factors influence the human ability to rapidly converge on shared communicative innovations.
  • Campisi, E., & Ozyurek, A. (2013). Iconicity as a communicative strategy: Recipient design in multimodal demonstrations for adults and children. Journal of Pragmatics, 47, 14-27. doi:10.1016/j.pragma.2012.12.007.

    Abstract

    Humans are the only species that uses communication to teach new knowledge to novices, usually to children (Tomasello, 1999 and Csibra and Gergely, 2006). This context of communication can employ “demonstrations” and it takes place with or without the help of objects (Clark, 1996). Previous research has focused on understanding the nature of demonstrations for very young children and with objects involved. However, little is known about the strategies used in demonstrating an action to an older child in comparison to another adult and without the use of objects, i.e., with gestures only. We tested if during demonstration of an action speakers use different degrees of iconicity in gestures for a child compared to an adult. 18 Italian subjects described to a camera how to make coffee imagining the listener as a 12-year-old child, a novice or an expert adult. While speech was found more informative both for the novice adult and for the child compared to the expert adult, the rate of iconic gestures increased and they were more informative and bigger only for the child compared to both of the adult conditions. Iconicity in gestures can be a powerful communicative strategy in teaching new knowledge to children in demonstrations and this is in line with claims that it can be used as a scaffolding device in grounding knowledge in experience (Perniss et al., 2010).
  • Cappuccio, M. L., Chu, M., & Kita, S. (2013). Pointing as an instrumental gesture: Gaze representation through indication. Humana.Mente: Journal of Philosophical Studies, 24, 125-149.

    Abstract

    We call those gestures “instrumental” that can enhance certain thinking processes of an agent by offering him representational models of his actions in a virtual space of imaginary performative possibilities. We argue that pointing is an instrumental gesture in that it represents geometrical information on one’s own gaze direction (i.e., a spatial model for attentional/ocular fixation/orientation), and provides a ritualized template for initiating gaze coordination and joint attention. We counter two possible objections, asserting respectively that the representational content of pointing is not constitutive, but derived from language, and that pointing directly solicits gaze coordination, without representing it. We consider two studies suggesting that attention and spatial perception are actively modified by one’s own pointing activity: the first study shows that pointing gestures help children link sets of objects to their corresponding number words; the second, that adults are faster and more accurate in counting when they point.
  • Cristia, A., Dupoux, E., Hakuno, Y., Lloyd-Fox, S., Schuetze, M., Kivits, J., Bergvelt, T., Van Gelder, M., Filippin, L., Charron, S., & Minagawa-Kawai, Y. (2013). An online database of infant functional Near InfraRed Spectroscopy studies: A community-augmented systematic review. PLoS One, 8(3): e58906. doi:10.1371/journal.pone.0058906.

    Abstract

    Until recently, imaging the infant brain was very challenging. Functional Near InfraRed Spectroscopy (fNIRS) is a promising, relatively novel technique, whose use is rapidly expanding. As an emergent field, it is particularly important to share methodological knowledge to ensure replicable and robust results. In this paper, we present a community-augmented database which will facilitate precisely this exchange. We tabulated articles and theses reporting empirical fNIRS research carried out on infants below three years of age along several methodological variables. The resulting spreadsheet has been uploaded in a format allowing individuals to continue adding new results, and download the most recent version of the table. Thus, this database is ideal to carry out systematic reviews. We illustrate its academic utility by focusing on the factors affecting three key variables: infant attrition, the reliability of oxygenated and deoxygenated responses, and signal-to-noise ratios. We then discuss strengths and weaknesses of the DBIfNIRS, and conclude by suggesting a set of simple guidelines aimed to facilitate methodological convergence through the standardization of reports.
  • Cristia, A. (2013). Input to language: The phonetics of infant-directed speech. Language and Linguistics Compass, 7, 157-170. doi:10.1111/lnc3.12015.

    Abstract

    Over the first year of life, infant perception changes radically as the child learns the phonology of the ambient language from the speech she is exposed to. Since infant-directed speech attracts the child's attention more than other registers, it is necessary to describe that input in order to understand language development, and to address questions of learnability. In this review, evidence from corpora analyses, experimental studies, and observational paradigms is brought together to outline the first comprehensive empirical picture of infant-directed speech and its effects on language acquisition. The ensuing landscape suggests that infant-directed speech provides an emotionally and linguistically rich input to language acquisition

    Additional information

    Cristia_Suppl_Material.xls
  • Cristia, A., Mielke, J., Daland, R., & Peperkamp, S. (2013). Similarity in the generalization of implicitly learned sound patterns. Journal of Laboratory Phonology, 4(2), 259-285.

    Abstract

    A core property of language is the ability to generalize beyond observed examples. In two experiments, we explore how listeners generalize implicitly learned sound patterns to new nonwords and to new sounds, with the goal of shedding light on how similarity affects treatment of potential generalization targets. During the exposure phase, listeners heard nonwords whose onset consonant was restricted to a subset of a natural class (e.g., /d g v z Z/). During the test phase, listeners were presented with new nonwords and asked to judge how frequently they had been presented before; some of the test items began with a consonant from the exposure set (e.g., /d/), and some began with novel consonants with varying relations to the exposure set (e.g., /b/, which is highly similar to all onsets in the training set; /t/, which is highly similar to one of the training onsets; and /p/, which is less similar than the other two). The exposure onset was rated most frequent, indicating that participants encoded onset attestation in the exposure set, and generalized it to new nonwords. Participants also rated novel consonants as somewhat frequent, indicating generalization to onsets that did not occur in the exposure phase. While generalization could be accounted for in terms of featural distance, it was insensitive to natural class structure. Generalization to new sounds was predicted better by models requiring prior linguistic knowledge (either traditional distinctive features or articulatory phonetic information) than by a model based on a linguistically naïve measure of acoustic similarity.

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