Publications

Displaying 1101 - 1189 of 1189
  • Vanlangendonck, F., Takashima, A., Willems, R. M., & Hagoort, P. (2018). Distinguishable memory retrieval networks for collaboratively and non-collaboratively learned information. Neuropsychologia, 111, 123-132. doi:10.1016/j.neuropsychologia.2017.12.008.

    Abstract

    Learning often occurs in communicative and collaborative settings, yet almost all research into the neural basis of memory relies on participants encoding and retrieving information on their own. We investigated whether learning linguistic labels in a collaborative context at least partly relies on cognitively and neurally distinct representations, as compared to learning in an individual context. Healthy human participants learned labels for sets of abstract shapes in three different tasks. They came up with labels with another person in a collaborative communication task (collaborative condition), by themselves (individual condition), or were given pre-determined unrelated labels to learn by themselves (arbitrary condition). Immediately after learning, participants retrieved and produced the labels aloud during a communicative task in the MRI scanner. The fMRI results show that the retrieval of collaboratively generated labels as compared to individually learned labels engages brain regions involved in understanding others (mentalizing or theory of mind) and autobiographical memory, including the medial prefrontal cortex, the right temporoparietal junction and the precuneus. This study is the first to show that collaboration during encoding affects the neural networks involved in retrieval.
  • Vanlangendonck, F., Willems, R. M., & Hagoort, P. (2018). Taking common ground into account: Specifying the role of the mentalizing network in communicative language production. PLoS One, 13(10): e0202943. doi:10.1371/journal.pone.0202943.
  • 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
  • Varma, S., Takashima, A., Krewinkel, S., Van Kooten, M., Fu, L., Medendorp, W. P., Kessels, R. P. C., & Daselaar, S. M. (2017). Non-interfering effects of active post-encoding tasks on episodic memory consolidation in humans. Frontiers in Behavioral Neuroscience, 11: 54. doi:10.3389/fnbeh.2017.00054.

    Abstract

    So far, studies that investigated interference effects of post-learning processes on episodic memory consolidation in humans have used tasks involving only complex and meaningful information. Such tasks require reallocation of general or encoding-specific resources away from consolidation-relevant activities. The possibility that interference can be elicited using a task that heavily taxes our limited brain resources, but has low semantic and hippocampal related long-term memory processing demands, has never been tested. We address this question by investigating whether consolidation could persist in parallel with an active, encoding-irrelevant, minimally semantic task, regardless of its high resource demands for cognitive processing. We distinguish the impact of such a task on consolidation based on whether it engages resources that are: (1) general/executive, or (2) specific/overlapping with the encoding modality. Our experiments compared subsequent memory performance across two post-encoding consolidation periods: quiet wakeful rest and a cognitively demanding n-Back task. Across six different experiments (total N = 176), we carefully manipulated the design of the n-Back task to target general or specific resources engaged in the ongoing consolidation process. In contrast to previous studies that employed interference tasks involving conceptual stimuli and complex processing demands, we did not find any differences between n-Back and rest conditions on memory performance at delayed test, using both recall and recognition tests. Our results indicate that: (1) quiet, wakeful rest is not a necessary prerequisite for episodic memory consolidation; and (2) post-encoding cognitive engagement does not interfere with memory consolidation when task-performance has minimal semantic and hippocampally-based episodic memory processing demands. We discuss our findings with reference to resource and reactivation-led interference theories
  • Varma, S., Daselaar, S. M., Kessels, R. P. C., & Takashima, A. (2018). Promotion and suppression of autobiographical thinking differentially affect episodic memory consolidation. PLoS One, 13(8): e0201780. doi:10.1371/journal.pone.0201780.

    Abstract

    During a post-encoding delay period, the ongoing consolidation of recently acquired memories can suffer interference if the delay period involves encoding of new memories, or sensory stimulation tasks. Interestingly, two recent independent studies suggest that (i) autobiographical thinking also interferes markedly with ongoing consolidation of recently learned wordlist material, while (ii) a 2-Back task might not interfere with ongoing consolidation, possibly due to the suppression of autobiographical thinking. In this study, we directly compare these conditions against a quiet wakeful rest baseline to test whether the promotion (via familiar sound-cues) or suppression (via a 2-Back task) of autobiographical thinking during the post-encoding delay period can affect consolidation of studied wordlists in a negative or a positive way, respectively. Our results successfully replicate previous studies and show a significant interference effect (as compared to the rest condition) when learning is followed by familiar sound-cues that promote autobiographical thinking, whereas no interference effect is observed when learning is followed by the 2-Back task. Results from a post-experimental experience-sampling questionnaire further show significant differences in the degree of autobiographical thinking reported during the three post-encoding periods: highest in the presence of sound-cues and lowest during the 2-Back task. In conclusion, our results suggest that varying levels of autobiographical thought during the post-encoding period may modulate episodic memory consolidation.
  • 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.
  • Verbeek, N. E., van der Maas, N. A. T., Sonsma, A. C. M., Ippel, E., Bondt, P., Hagebeuk, E., Jansen, F. E., Geesink, H. H., Braun, K. P., de Louw, A., Augustijn, P. B., Neuteboom, R. F., Schieving, J. H., Stroink, H., Vermeulen, R. J., Nicolai, J., Brouwer, O. F., Van Kempen, M., De Kovel, C. G. F., Kemmeren, J. M. and 5 moreVerbeek, N. E., van der Maas, N. A. T., Sonsma, A. C. M., Ippel, E., Bondt, P., Hagebeuk, E., Jansen, F. E., Geesink, H. H., Braun, K. P., de Louw, A., Augustijn, P. B., Neuteboom, R. F., Schieving, J. H., Stroink, H., Vermeulen, R. J., Nicolai, J., Brouwer, O. F., Van Kempen, M., De Kovel, C. G. F., Kemmeren, J. M., Koeleman, B. P. C., Knoers, N. V., Lindhout, D., Gunning, W. B., & Brilstra, E. H. (2015). Effect of vaccinations on seizure risk and disease course in Dravet syndrome. Neurology, 85(7), 596-603. doi:10.1212/wnl.0000000000001855.

    Abstract

    Objective: To study the effect of vaccination-associated seizure onset on disease course and estimate the risk of subsequent seizures after infant pertussis combination and measles, mumps, and rubella (MMR) vaccinations in Dravet syndrome (DS). Methods: We retrospectively analyzed data from hospital medical files, child health clinics, and the vaccination register for children with DS and pathogenic SCN1A mutations. Seizures within 24 hours after infant whole-cell, acellular, or nonpertussis combination vaccination or within 5 to 12 days after MMR vaccination were defined as "vaccination-associated." Risks of vaccination-associated seizures for the different vaccines were analyzed in univariable and in multivariable logistic regression for pertussis combination vaccines and by a self-controlled case series analysis using parental seizure registries for MMR vaccines. Disease courses of children with and without vaccination-associated seizure onset were compared. Results: Children who had DS (n = 77) with and without vaccination-associated seizure onset (21% and 79%, respectively) differed in age at first seizure (median 3.7 vs 6.1 months, p < 0.001) but not in age at first nonvaccination-associated seizure, age at first report of developmental delay, or cognitive outcome. The risk of subsequent vaccination-associated seizures was significantly lower for acellular pertussis (9%; odds ratio 0.18, 95% confidence interval [CI] 0.05-0.71) and nonpertussis (8%; odds ratio 0.11, 95% CI 0.02-0.59) than whole-cell pertussis (37%; reference) vaccines. Self-controlled case series analysis showed an increased incidence rate ratio of seizures of 2.3 (95% CI 1.5-3.4) within the risk period of 5 to 12 days following MMR vaccination. Conclusions: Our results suggest that vaccination-associated earlier seizure onset does not alter disease course in DS, while the risk of subsequent vaccination-associated seizures is probably vaccine-specific.
  • Verdonschot, R. G., Tokimoto, S., & Miyaoka, Y. (2019). The fundamental phonological unit of Japanese word production: An EEG study using the picture-word interference paradigm. Journal of Neurolinguistics, 51, 184-193. doi:10.1016/j.jneuroling.2019.02.004.

    Abstract

    It has been shown that in Germanic languages (e.g. English, Dutch) phonemes are the primary (or proximate) planning units during the early stages of phonological encoding. Contrastingly, in Chinese and Japanese the phoneme does not seem to play an important role but rather the syllable (Chinese) and mora (Japanese) are essential. However, despite the lack of behavioral evidence, neurocorrelational studies in Chinese suggested that electrophysiological brain responses (i.e. preceding overt responses) may indicate some significance for the phoneme. We investigated this matter in Japanese and our data shows that unlike in Chinese (for which the literature shows mixed effects), in Japanese both the behavioral and neurocorrelational data indicate an important role only for the mora (and not the phoneme) during the early stages of phonological encoding.
  • Verdonschot, R. G., & Kinoshita, S. (2018). Mora or more? The phonological unit of Japanese word production in the Stroop color naming task. Memory & Cognition, 46(3), 410-425. doi:10.3758/s13421-017-0774-4.

    Abstract

    In English, Dutch, and other European languages, it is well established that the fundamental phonological unit in word production is the phoneme; in contrast, recent studies have shown that in Chinese it is the (atonal) syllable and in Japanese the mora. The present study investigated whether this cross-language variation in the size of the unit of word production is due to the type of script used in the language (i.e., alphabetic, morphosyllabic, or moraic). Capitalizing on the multiscriptal nature of Japanese, and using the Stroop color naming task, we show that the overlap in the initial mora between the color name and the written distractor facilitates color naming independent of script type. These results confirm the mora as the phonological unit of word production in Japanese, and establish the Stroop color naming task as a useful task for investigating the fundamental (or "proximate") phonological unit used in speech production.
  • Verdonschot, R. G., Lai, J., Chen, F., Tamaoka, K., & Schiller, N. O. (2015). Constructing initial phonology in Mandarin Chinese: Syllabic or subsyllabic? A masked priming investigation. Japanese Psychological Research, 57(1), 61-68. doi:10.1111/jpr.12064.

    Abstract

    Recent research has put forward the idea that Chinese speech production is governed by the syllable as the fundamental phonological unit. However, it may be that onset priming might be more difficult to obtain in Mandarin Chinese. Therefore, in this study, the degree of overlap between prime and target was increased from C to CV (i.e., extending beyond the phoneme) as well as whether primes and targets had an overlapping structure (CV vs. CVN). Subsyllabic priming effects were found (i.e., onset + vowel overlap but not purely onset overlap), contrasting with the claim that the syllable is the compulsory building block in the initial construction of Mandarin Chinese phonology.
  • Verdonschot, R. G., & Tamaoka, K. (2015). Editorial: The production of speech sounds across languages. Japanese Psychological Research, 57(1), 1-3. doi:10.1111/jpr.12073.
  • Verdonschot, R. G., Guillemaud, H., Rabenarivo, H., & Tamaoka, K. (2015). The Microsoft KINECT: A novel tool for psycholinguistic research. Open Journal of Modern Linguistics, 5, 291-301. doi:10.4236/ojml.2015.53026.

    Abstract

    The Microsoft KINECT is a 3D sensing device originally developed for the XBOX. The Microsoft KINECT opens up many exciting new opportunities for conducting experimental research on human behavior. We investigated some of these possibilities within the field of psycholinguistics (specifically: language production) by creating software, using C#, allowing for the KINECT to be used in a typical psycholinguistic experimental setting. The results of a naming experiment using this software confirmed that the KINECT was able to measure the effects of a robust psycholinguistic variable (word frequency) on naming latencies. However, although the current version of the software is able to measure psycholinguistic variables of interest, we also discuss several points where the software can still stand to be improved. The main aim of this paper is to make the software freely available for assessment and use by the psycholinguistic community and to illustrate the KINECT as a potentially valuable tool for investigating human behavior, especially in the field of psycholinguistics.
  • Verdonschot, R. G., & Tamaoka, K. (Eds.). (2015). The production of speech sounds across languages [Special Issue]. Japanese Psychological Research, 57(1).
  • Verga, L., & Kotz, S. A. (2017). Help me if I can't: Social interaction effects in adult contextual word learning. Cognition, 168, 76-90. doi:10.1016/j.cognition.2017.06.018.

    Abstract

    A major challenge in second language acquisition is to build up new vocabulary. How is it possible to identify the meaning of a new word among several possible referents? Adult learners typically use contextual information, which reduces the number of possible referents a new word can have. Alternatively, a social partner may facilitate word learning by directing the learner’s attention toward the correct new word meaning. While much is known about the role of this form of ‘joint attention’ in first language acquisition, little is known about its efficacy in second language acquisition. Consequently, we introduce and validate a novel visual word learning game to evaluate how joint attention affects the contextual learning of new words in a second language. Adult learners either acquired new words in a constant or variable sentence context by playing the game with a knowledgeable partner, or by playing the game alone on a computer. Results clearly show that participants who learned new words in social interaction (i) are faster in identifying a correct new word referent in variable sentence contexts, and (ii) temporally coordinate their behavior with a social partner. Testing the learned words in a post-learning recall or recognition task showed that participants, who learned interactively, better recognized words originally learned in a variable context. While this result may suggest that interactive learning facilitates the allocation of attention to a target referent, the differences in the performance during recognition and recall call for further studies investigating the effect of social interaction on learning performance. In summary, we provide first evidence on the role joint attention in second language learning. Furthermore, the new interactive learning game offers itself to further testing in complex neuroimaging research, where the lack of appropriate experimental set-ups has so far limited the investigation of the neural basis of adult word learning in social interaction.
  • Verga, L., & Kotz, S. A. (2019). Putting language back into ecological communication contexts. Language, Cognition and Neuroscience, 34(4), 536-544. doi:10.1080/23273798.2018.1506886.

    Abstract

    Language is a multi-faceted form of communication. It is not until recently though that language research moved on from simple stimuli and protocols toward a more ecologically valid approach, namely “shifting” from words and simple sentences to stories with varying degrees of contextual complexity. While much needed, the use of ecologically valid stimuli such as stories should also be explored in interactive rather than individualistic experimental settings leading the way to an interactive neuroscience of language. Indeed, mounting evidence suggests that cognitive processes and their underlying neural activity significantly differ between social and individual experiences. We aim at reviewing evidence, which indicates that the characteristics of linguistic and extra-linguistic contexts may significantly influence communication–including spoken language comprehension. In doing so, we provide evidence on the use of new paradigms and methodological advancements that may enable the study of complex language features in a truly interactive, ecological way.
  • Verga, L., & Kotz, S. A. (2019). Spatial attention underpins social word learning in the right fronto-parietal network. NeuroImage, 195, 165-173. doi:10.1016/j.neuroimage.2019.03.071.

    Abstract

    In a multi- and inter-cultural world, we daily encounter new words. Adult learners often rely on a situational context to learn and understand a new word's meaning. Here, we explored whether interactive learning facilitates word learning by directing the learner's attention to a correct new word referent when a situational context is non-informative. We predicted larger involvement of inferior parietal, frontal, and visual cortices involved in visuo-spatial attention during interactive learning. We scanned participants while they played a visual word learning game with and without a social partner. As hypothesized, interactive learning enhanced activity in the right Supramarginal Gyrus when the situational context provided little information. Activity in the right Inferior Frontal Gyrus during interactive learning correlated with post-scanning behavioral test scores, while these scores correlated with activity in the Fusiform Gyrus in the non-interactive group. These results indicate that attention is involved in interactive learning when the situational context is minimal and suggest that individual learning processes may be largely different from interactive ones. As such, they challenge the ecological validity of what we know about individual learning and advocate the exploration of interactive learning in naturalistic settings.
  • Verga, L., Bigand, E., & Kotz, S. A. (2015). Play along: Effects of music and social interaction on word learning. Frontiers in Psychology, 6: 1316. doi:10.3389/fpsyg.2015.01316.

    Abstract

    Learning new words is an increasingly common necessity in everyday life. External factors, among which music and social interaction are particularly debated, are claimed to facilitate this task. Due to their influence on the learner’s temporal behavior, these stimuli are able to drive the learner’s attention to the correct referent of new words at the correct point in time. However, do music and social interaction impact learning behavior in the same way? The current study aims to answer this question. Native German speakers (N = 80) were requested to learn new words (pseudo-words) during a contextual learning game. This learning task was performed alone with a computer or with a partner, with or without music. Results showed that music and social interaction had a different impact on the learner’s behavior: Participants tended to temporally coordinate their behavior more with a partner than with music, and in both cases more than with a computer. However, when both music and social interaction were present, this temporal coordination was hindered. These results suggest that while music and social interaction do influence participants’ learning behavior, they have a different impact. Moreover, impaired behavior when both music and a partner are present suggests that different mechanisms are employed to coordinate with the two types of stimuli. Whether one or the other approach is more efficient for word learning, however, is a question still requiring further investigation, as no differences were observed between conditions in a retrieval phase, which took place immediately after the learning session. This study contributes to the literature on word learning in adults by investigating two possible facilitating factors, and has important implications for situations such as music therapy, in which music and social interaction are present at the same time.
  • Verhees, M. W. F. T., Chwilla, D. J., Tromp, J., & Vissers, C. T. W. M. (2015). Contributions of emotional state and attention to the processing of syntactic agreement errors: evidence from P600. Frontiers in Psychology, 6: 388. doi:10.3389%2Ffpsyg.2015.00388.

    Abstract

    The classic account of language is that language processing occurs in isolation from other cognitive systems, like perception, motor action, and emotion. The central theme of this paper is the relationship between a participant’s emotional state and language comprehension. Does emotional context affect how we process neutral words? Recent studies showed that processing of word meaning – traditionally conceived as an automatic process – is affected by emotional state. The influence of emotional state on syntactic processing is less clear. One study reported a mood-related P600 modulation, while another study did not observe an effect of mood on syntactic processing. The goals of this study were: First, to clarify whether and if so how mood affects syntactic processing. Second, to shed light on the underlying mechanisms by separating possible effects of mood from those of attention on syntactic processing. Event-related potentials (ERPs) were recorded while participants read syntactically correct or incorrect sentences. Mood (happy vs. sad) was manipulated by presenting film clips. Attention was manipulated by directing attention to syntactic features vs. physical features. The mood induction was effective. Interactions between mood, attention and syntactic correctness were obtained, showing that mood and attention modulated P600. The mood manipulation led to a reduction in P600 for sad as compared to happy mood when attention was directed at syntactic features. The attention manipulation led to a reduction in P600 when attention was directed at physical features compared to syntactic features for happy mood. From this we draw two conclusions: First, emotional state does affect syntactic processing. We propose mood-related differences in the reliance on heuristics as the underlying mechanism. Second, attention can contribute to emotion-related ERP effects in syntactic language processing. Therefore, future studies on the relation between language and emotion will have to control for effects of attention
  • Verheijen, J., Van der Zee, J., Gijselinck, I., Van den Bossche, T., Dillen, L., Heeman, B., Gómez-Tortosa, E., Lladó, A., Sanchez-Valle, R., Graff, C., Pastor, P., Pastor, M. A., Benussi, L., Ghidoni, R., Binetti, G., Clarimon, J., De Mendonça, A., Gelpi, E., Tsolaki, M., Diehl-Schmid, J. and 12 moreVerheijen, J., Van der Zee, J., Gijselinck, I., Van den Bossche, T., Dillen, L., Heeman, B., Gómez-Tortosa, E., Lladó, A., Sanchez-Valle, R., Graff, C., Pastor, P., Pastor, M. A., Benussi, L., Ghidoni, R., Binetti, G., Clarimon, J., De Mendonça, A., Gelpi, E., Tsolaki, M., Diehl-Schmid, J., Nacmias, B., Almeida, M. R., Borroni, B., Matej, R., Ruiz, A., Engelborghs, S., Vandenberghe, R., De Deyn, P. P., Cruts, M., Van Broeckhoven, C., Sleegers, K., BELNEU Consortium, & EU EOD Consortium (2018). Common and rare TBK1 variants in early-onset Alzheimer disease in a European cohort. Neurobiology of Aging, 62, 245.e1-245.e7. doi:10.1016/j.neurobiolaging.2017.10.012.

    Abstract

    TANK-binding kinase 1 (TBK1) loss-of-function (LoF) mutations are known to cause frontotemporal dementia (FTD) and amyotrophic lateral sclerosis (ALS), often combined with memory deficits early in the disease course. We performed targeted resequencing of TBK1 in 1253 early onset Alzheimer's disease (EOAD) patients from 8 European countries to investigate whether pathogenic TBK1 mutations are enriched among patients with clinical diagnosis of EOAD. Variant frequencies were compared against 2117 origin-matched controls. We identified only 1 LoF mutation (p.Thr79del) in a patient clinically diagnosed with Alzheimer's disease and a positive family history of ALS. We did not observe enrichment of rare variants in EOAD patients compared to controls, nor of rare variants affecting NFκB induction. Of 3 common coding variants, rs7486100 showed evidence of association (OR 1.46 [95% CI 1.13–1.9]; p-value 0.01). Homozygous carriers of the risk allele showed reduced expression of TBK1 (p-value 0.03). Our findings are not indicative of a significant role for TBK1 mutations in EOAD. The association between common variants in TBK1, disease risk and reduced TBK1 expression warrants follow-up in FTD/ALS cohorts. © 2017 The Author(s)

    Additional information

    Supplementary data
  • Verheijen, J., & Sleegers, K. (2018). Understanding Alzheimer Disease at the interface between genetics and transcriptomics. Trends in Genetics, 34(6), 434-447. doi:10.1016/j.tig.2018.02.007.

    Abstract

    Over 25 genes are known to affect the risk of developing Alzheimer disease (AD), the most common neurodegenerative dementia. However, mechanistic insights and improved disease management remains limited, due to difficulties in determining the functional consequences of genetic associations. Transcriptomics is increasingly being used to corroborate or enhance interpretation of genetic discoveries. These approaches, which include second and third generation sequencing, single-cell sequencing, and bioinformatics, reveal allele-specific events connecting AD risk genes to expression profiles, and provide converging evidence of pathophysiological pathways underlying AD. Simultaneously, they highlight brain region- and cell-type-specific expression patterns, and alternative splicing events that affect the straightforward relation between a genetic variant and AD, re-emphasizing the need for an integrated approach of genetics and transcriptomics in understanding AD. © 2018 The Authors
  • Verhoef, E., Demontis, D., Burgess, S., Shapland, C. Y., Dale, P. S., Okbay, A., Neale, B. M., Faraone, S. V., iPSYCH-Broad-PGC ADHD Consortium, Stergiakouli, E., Davey Smith, G., Fisher, S. E., Borglum, A., & St Pourcain, B. (2019). Disentangling polygenic associations between Attention-Deficit/Hyperactivity Disorder, educational attainment, literacy and language. Translational Psychiatry, 9: 35. doi:10.1038/s41398-018-0324-2.

    Abstract

    Interpreting polygenic overlap between ADHD and both literacy-related and language-related impairments is challenging as genetic associations might be influenced by indirectly shared genetic factors. Here, we investigate genetic overlap between polygenic ADHD risk and multiple literacy-related and/or language-related abilities (LRAs), as assessed in UK children (N ≤ 5919), accounting for genetically predictable educational attainment (EA). Genome-wide summary statistics on clinical ADHD and years of schooling were obtained from large consortia (N ≤ 326,041). Our findings show that ADHD-polygenic scores (ADHD-PGS) were inversely associated with LRAs in ALSPAC, most consistently with reading-related abilities, and explained ≤1.6% phenotypic variation. These polygenic links were then dissected into both ADHD effects shared with and independent of EA, using multivariable regressions (MVR). Conditional on EA, polygenic ADHD risk remained associated with multiple reading and/or spelling abilities, phonemic awareness and verbal intelligence, but not listening comprehension and non-word repetition. Using conservative ADHD-instruments (P-threshold < 5 × 10−8), this corresponded, for example, to a 0.35 SD decrease in pooled reading performance per log-odds in ADHD-liability (P = 9.2 × 10−5). Using subthreshold ADHD-instruments (P-threshold < 0.0015), these effects became smaller, with a 0.03 SD decrease per log-odds in ADHD risk (P = 1.4 × 10−6), although the predictive accuracy increased. However, polygenic ADHD-effects shared with EA were of equal strength and at least equal magnitude compared to those independent of EA, for all LRAs studied, and detectable using subthreshold instruments. Thus, ADHD-related polygenic links with LRAs are to a large extent due to shared genetic effects with EA, although there is evidence for an ADHD-specific association profile, independent of EA, that primarily involves literacy-related impairments.

    Additional information

    41398_2018_324_MOESM1_ESM.docx
  • Verkerk, A. (2015). Where do all the motion verbs come from? The speed of development of manner verbs and path verbs in Indo-European. Diachronica, 32(1), 69-104. doi:10.1075/dia.32.1.03ver.

    Abstract

    The last four decades have seen huge progress in the description and analysis of cross-linguistic diversity in the encoding of motion (Talmy 1985, 1991, Slobin 1996, 2004). Comparisons between satellite-framed and verb-framed languages suggest that satellite-framed languages typically have a larger manner of motion verb lexicon (swim, dash), while verb-framed languages typically have a larger path of motion verb lexicon (enter, cross) (Slobin 2004, Verkerk 2013, 2014b). This paper investigates how differences between the motion verb lexicons of satellite-framed and verb-framed languages emerge. Phylogenetic comparative methods adopted from biology and an etymological study are used to investigate manner verb lexicons and path verb lexicons in an Indo-European dataset. I show that manner verbs and path verbs typically have different types of etymological origins and that manner verbs emerge faster in satellite-framed subgroups, while path verbs emerge faster in verb-framed subgroups.
  • Vernes, S. C. (2017). What bats have to say about speech and language. Psychonomic Bulletin & Review, 24(1), 111-117. doi:10.3758/s13423-016-1060-3.

    Abstract

    Understanding the biological foundations of language is vital to gaining insight into how the capacity for language may have evolved in humans. Animal models can be exploited to learn about the biological underpinnings of shared human traits, and although no other animals display speech or language, a range of behaviors found throughout the animal kingdom are relevant to speech and spoken language. To date, such investigations have been dominated by studies of our closest primate relatives searching for shared traits, or more distantly related species that are sophisticated vocal communicators, like songbirds. Herein I make the case for turning our attention to the Chiropterans, to shed new light on the biological encoding and evolution of human language-relevant traits. Bats employ complex vocalizations to facilitate navigation as well as social interactions, and are exquisitely tuned to acoustic information. Furthermore, bats display behaviors such as vocal learning and vocal turn-taking that are directly pertinent for human spoken language. Emerging technologies are now allowing the study of bat vocal communication, from the behavioral to the neurobiological and molecular level. Although it is clear that no single animal model can reflect the complexity of human language, by comparing such findings across diverse species we can identify the shared biological mechanisms likely to have influenced the evolution of human language. Keywords
  • Versace, E., Rogge, J. R., Shelton-May, N., & Ravignani, A. (2019). Positional encoding in cotton-top tamarins (Saguinus oedipus). Animal Cognition, 22, 825-838. doi:10.1007/s10071-019-01277-y.

    Abstract

    Strategies used in artificial grammar learning can shed light into the abilities of different species to extract regularities from the environment. In the A(X)nB rule, A and B items are linked, but assigned to different positional categories and separated by distractor items. Open questions are how widespread is the ability to extract positional regularities from A(X)nB patterns, which strategies are used to encode positional regularities and whether individuals exhibit preferences for absolute or relative position encoding. We used visual arrays to investigate whether cotton-top tamarins (Saguinusoedipus) can learn this rule and which strategies they use. After training on a subset of exemplars, two of the tested monkeys successfully generalized to novel combinations. These tamarins discriminated between categories of tokens with different properties (A, B, X) and detected a positional relationship between non-adjacent items even in the presence of novel distractors. The pattern of errors revealed that successful subjects used visual similarity with training stimuli to solve the task and that successful tamarins extracted the relative position of As and Bs rather than their absolute position, similarly to what has been observed in other species. Relative position encoding appears to be favoured in different tasks and taxa. Generalization, though, was incomplete, since we observed a failure with items that during training had always been presented in reinforced arrays, showing the limitations in grasping the underlying positional rule. These results suggest the use of local strategies in the extraction of positional rules in cotton-top tamarins.

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  • Verspeek, J., Staes, N., Van Leeuwen, E. J. C., Eens, M., & Stevens, J. M. G. (2019). Bonobo personality predicts friendship. Scientific Reports, 9: 19245. doi:10.1038/s41598-019-55884-3.

    Abstract

    In bonobos, strong bonds have been documented between unrelated females and between mothers
    and their adult sons, which can have important fitness benefits. Often age, sex or kinship similarity
    have been used to explain social bond strength variation. Recent studies in other species also stress
    the importance of personality, but this relationship remains to be investigated in bonobos. We used
    behavioral observations on 39 adult and adolescent bonobos housed in 5 European zoos to study the
    role of personality similarity in dyadic relationship quality. Dimension reduction analyses on individual
    and dyadic behavioral scores revealed multidimensional personality (Sociability, Openness, Boldness,
    Activity) and relationship quality components (value, compatibility). We show that, aside from
    relatedness and sex combination of the dyad, relationship quality is also associated with personality
    similarity of both partners. While similarity in Sociability resulted in higher relationship values, lower
    relationship compatibility was found between bonobos with similar Activity scores. The results of this
    study expand our understanding of the mechanisms underlying social bond formation in anthropoid
    apes. In addition, we suggest that future studies in closely related species like chimpanzees should
    implement identical methods for assessing bond strength to shed further light on the evolution of this
    phenomenon.

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  • Viebahn, M., McQueen, J. M., Ernestus, M., Frauenfelder, U. H., & Bürki, A. (2018). How much does orthography influence the processing of reduced word forms? Evidence from novel-word learning about French schwa deletion. The Quarterly Journal of Experimental Psychology, 71(11), 2378-2394. doi:10.1177/1747021817741859.

    Abstract

    This study examines the influence of orthography on the processing of reduced word forms. For this purpose, we compared the impact of phonological variation with the impact of spelling-sound consistency on the processing of words that may be produced with or without the vowel schwa. Participants learnt novel French words in which the vowel schwa was present or absent in the first syllable. In Experiment 1, the words were consistently produced without schwa or produced in a variable manner (i.e., sometimes produced with and sometimes produced without schwa). In Experiment 2, words were always produced in a consistent manner, but an orthographic exposure phase was included in which words that were produced without schwa were either spelled with or without the letter . Results from naming and eye-tracking tasks suggest that both phonological variation and spelling-sound consistency influence the processing of spoken novel words. However, the influence of phonological variation outweighs the effect of spelling-sound consistency. Our findings therefore suggest that the influence of orthography on the processing of reduced word forms is relatively small.
  • Viebahn, M., Ernestus, M., & McQueen, J. M. (2017). Speaking style influences the brain’s electrophysiological response to grammatical errors in speech comprehension. Journal of Cognitive Neuroscience, 29(7), 1132-1146. doi:10.1162/jocn_a_01095.

    Abstract

    This electrophysiological study asked whether the brain processes grammatical gender
    violations in casual speech differently than in careful speech. Native speakers of Dutch were
    presented with utterances that contained adjective-noun pairs in which the adjective was either
    correctly inflected with a word-final schwa (e.g. een spannende roman “a suspenseful novel”) or
    incorrectly uninflected without that schwa (een spannend roman). Consistent with previous
    findings, the uninflected adjectives elicited an electrical brain response sensitive to syntactic
    violations when the talker was speaking in a careful manner. When the talker was speaking in a
    casual manner, this response was absent. A control condition showed electrophysiological responses
    for carefully as well as casually produced utterances with semantic anomalies, showing that
    listeners were able to understand the content of both types of utterance. The results suggest that
    listeners take information about the speaking style of a talker into account when processing the
    acoustic-phonetic information provided by the speech signal. Absent schwas in casual speech are
    effectively not grammatical gender violations. These changes in syntactic processing are evidence
    of contextually-driven neural flexibility.

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  • Viebahn, M., Ernestus, M., & McQueen, J. M. (2015). Syntactic predictability in the recognition of carefully and casually produced speech. Journal of Experimental Psychology: Learning, Memory, and Cognition, 41(6), 1684-1702. doi:10.1037/a0039326.
  • Villanueva, P., Nudel, R., Hoischen, A., Fernández, M. A., Simpson, N. H., Gilissen, C., Reader, R. H., Jara, L., Echeverry, M., Francks, C., Baird, G., Conti-Ramsden, G., O’Hare, A., Bolton, P., Hennessy, E. R., the SLI Consortium, Palomino, H., Carvajal-Carmona Veltman J.A., L., Veltman, J. A., Cazier, J.-B. and 3 moreVillanueva, P., Nudel, R., Hoischen, A., Fernández, M. A., Simpson, N. H., Gilissen, C., Reader, R. H., Jara, L., Echeverry, M., Francks, C., Baird, G., Conti-Ramsden, G., O’Hare, A., Bolton, P., Hennessy, E. R., the SLI Consortium, Palomino, H., Carvajal-Carmona Veltman J.A., L., Veltman, J. A., Cazier, J.-B., De Barbieri, Z., Fisher, S. E., & Newbury, D. (2015). Exome sequencing in an admixed isolated population indicates NFXL1 variants confer a risk for Specific Language Impairment. PLoS Genetics, 11(3): e1004925. doi:10.1371/journal.pgen.1004925.
  • Vogels, J., & Van Bergen, G. (2017). Where to place inaccessible subjects in Dutch: The role of definiteness and animacy. Corpus linguistics and linguistic theory, 13(2), 369-398. doi:10.1515/cllt-2013-0021.

    Abstract

    Cross-linguistically, both subjects and topical information tend to be placed at the beginning of a sentence. Subjects are generally highly topical, causing both tendencies to converge on the same word order. However, subjects that lack prototypical topic properties may give rise to an incongruence between the preference to start a sentence with the subject and the preference to start a sentence with the most accessible information. We present a corpus study in which we investigate in what syntactic position (preverbal or postverbal) such low-accessible subjects are typically found in Dutch natural language. We examine the effects of both discourse accessibility (definiteness) and inherent accessibility (animacy). Our results show that definiteness and animacy interact in determining subject position in Dutch. Non-referential (bare) subjects are less likely to occur in preverbal position than definite subjects, and this tendency is reinforced when the subject is inanimate. This suggests that these two properties that make the subject less accessible together can ‘gang up’ against the subject first preference. The results support a probabilistic multifactorial account of syntactic variation.
  • Volker-Touw, C. M., de Koning, H. D., Giltay, J., De Kovel, C. G. F., van Kempen, T. S., Oberndorff, K., Boes, M., van Steensel, M. A., van Well, G. T., Blokx, W. A., Schalkwijk, J., Simon, A., Frenkel, J., & van Gijn, M. E. (2017). Erythematous nodes, urticarial rash and arthralgias in a large pedigree with NLRC4-related autoinflammatory disease, expansion of the phenotype. British Journal of Dermatology, 176(1), 244-248. doi:10.1111/bjd.14757.

    Abstract

    Autoinflammatory disorders (AID) are a heterogeneous group of diseases, characterized by an unprovoked innate immune response, resulting in recurrent or ongoing systemic inflammation and fever1-3. Inflammasomes are protein complexes with an essential role in pyroptosis and the caspase-1-mediated activation of the proinflammatory cytokines IL-1β, IL-17 and IL-18.
  • De Vos, J., Schriefers, H., Nivard, M. C., & Lemhöfer, K. (2018). A meta‐analysis and meta‐regression of incidental second language word learning from spoken input. Language Learning, 68(4), 906-941. doi:10.1111/lang.12296.

    Abstract

    We meta‐analyzed the effectiveness of incidental second language word learning from spoken input. Our sample contained 105 effect sizes from 32 primary studies employing meaning‐focused word‐learning activities with 1,964 participants with typical cognitive functioning. The random‐effects meta‐analysis yielded a mean effect size of g = 1.05, reflecting generally large vocabulary gains from spoken input in meaning‐focused activities. A meta‐regression with three substantive and two methodological predictors also revealed that adult participants outperformed children in terms of word learning and that interactive learning tasks were more effective than noninteractive ones. Furthermore, learning scores were higher when measured with recognition than with recall tests. Methodologically, the use of a no‐input control group seemed to protect against an overestimation of learning effects, evidenced by smaller effect sizes. Finally, whether a pretest–posttest design was used did not influence effect sizes. All data and the analysis script are publicly available.
  • De Vos, J., Schriefers, H., Bosch, L. t., & Lemhöfer, K. (2019). Interactive L2 vocabulary acquisition in a lab-based immersion setting. Language, Cognition and Neuroscience, 34(7), 916-935. doi:10.1080/23273798.2019.1599127.

    Abstract

    ABSTRACTWe investigated to what extent L2 word learning in spoken interaction takes place when learners are unaware of taking part in a language learning study. Using a novel paradigm for approximating naturalistic (but not necessarily non-intentional) L2 learning in the lab, German learners of Dutch were led to believe that the study concerned judging the price of objects. Dutch target words (object names) were selected individually such that these words were unknown to the respective participant. Then, in a dialogue-like task with the experimenter, the participants were first exposed to and then tested on the target words. In comparison to a no-input control group, we observed a clear learning effect especially from the first two exposures, and better learning for cognates than for non-cognates, but no modulating effect of the exposure-production lag. Moreover, some of the acquired knowledge persisted over a six-month period.
  • De Vos, C., & Nyst, V.A.S (2018). Introduction: The time-depth and typology of rural sign languages. Sign Language Studies, 18(4), 477-487.
  • De Vos, J., Schriefers, H., & Lemhöfer, K. (2018). Noticing vocabulary holes aids incidental second language word learning: An experimental study. Bilingualism: Language and Cognition, 22(3), 500-515. doi:10.1017/S1366728918000019.

    Abstract

    Noticing the hole (NTH) occurs when speakers want to say something, but realise they do not know the right word(s). Such awareness of lacking knowledge supposedly facilitates the acquisition of the unknown word(s) from later input (Swain, 1993). We tested this claim by experimentally inducing NTH in a second language (L2) for some participants (experimental), but not others (control). Then, in a price comparison game, all participants were exposed to spoken L2 input containing the to-be-learned words. They were unaware of taking part in an L2 study. Post-tests showed that participants who had noticed holes in their vocabulary had indeed learned more words compared to participants who had not. This held both for the experimental group as well as those participants in the control group who later reported to have noticed holes. Thus, when we become aware of vocabulary holes, the first step to improve our vocabulary is already taken.
  • De Vos, C., & Pfau, R. (2015). Sign Language Typology: The contribution of rural sign languages. Annual Review of Linguistics, 1, 265-288. doi:10.1146/annurev-linguist-030514-124958.

    Abstract

    Since the 1990s, the field of sign language typology has shown that sign languages exhibit typological variation at all relevant levels of linguistic description. These initial typological comparisons were heavily skewed toward the urban sign languages of developed countries, mostly in the Western world. This review reports on the recent contributions made by rural signing varieties, that is, sign languages that have evolved in village communities, often in developing countries, due to a high incidence of deafness. With respect to a number of structural properties, rural sign languages fit into previously established typological classifications. However, they also exhibit unique and typologically marked features that challenge received views on possible sign languages. At the same time, the shared features of geographically dispersed rural signing varieties provide a unique window into the social dynamics that may shape the structures of modern human languages.
  • De Vos, C. (2015). The Kata Kolok pointing system: Morphemization and syntactic integration. Topics in Cognitive Science, 7(1), 150-168. doi:10.1111/tops.12124.

    Abstract

    Signed utterances are densely packed with pointing signs, reaching a frequency of one in six signs in spontaneous conversations (de Vos, 2012; Johnston, 2013a; Morford & MacFarlane, 2003). These pointing signs attain a wide range of functions and are formally highly diversified. Based on corpus analysis of spontaneous pointing signs in Kata Kolok, a rural signing variety of Bali, this paper argues that the full meaning potentials of pointing signs come about through the integration of a varied set of linguistic and extralinguistic cues. Taking this hybrid nature of point- ing phenomena into account, it is argued that pointing signs may become an intrinsic aspect of sign language grammars through two mechanisms: morphemization and syntactic integration. Although not entailed in this research, this approach could implicate that some highly systema- tized pointing systems of speaking communities may to a degree be grammatical as well.

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    tops12124-sup-0001-AppenidxSa-Sb.docx
  • De Vos, C., Torreira, F., & Levinson, S. C. (2015). Turn-timing in signed conversations: Coordinating stroke-to-stroke turn boundaries. Frontiers in Psychology, 6: 268. doi:10.3389/fpsyg.2015.00268.

    Abstract

    In spoken interactions, interlocutors carefully plan and time their utterances, minimising gaps and overlaps between consecutive turns. Cross-linguistic comparison has indicated that spoken languages vary only minimally in terms of turn-timing, and language acquisition research has shown pre-linguistic vocal turn-taking in the first half year of life. These observations suggest that the turn-taking system may provide a fundamental basis for our linguistic capacities. The question remains however to what extent our capacity for rapid turn-taking is determined by modality constraints. The avoidance of overlapping turns could be motivated by the difficulty of hearing and speaking at the same time. If so, turn-taking in sign might show greater toleration for overlap. Alternatively, signed conversations may show a similar distribution of turn-timing as spoken languages, thus avoiding both gaps and overlaps. To address this question we look at turn-timing in question-answer sequences in spontaneous conversations of Sign Language of the Netherlands. The findings indicate that although there is considerable overlap in two or more signers' articulators in conversation, when proper allowance is made for onset preparation, post-utterance retraction and the intentional holding of signs for response, turn-taking latencies in sign look remarkably like those reported for spoken language. This is consistent with the possibility that, at least with regard to responses to questions, speakers and signers follow similar time courses in planning and producing their utterances in on-going conversation. This suggests that turn-taking systems may well be a shared cognitive infrastructure underlying all modern human languages, both spoken and signed.
  • Vosse, T., & Kempen, G. (2000). Syntactic structure assembly in human parsing: A computational model based on competitive inhibition and a lexicalist grammar. Cognition, 75, 105-143.

    Abstract

    We present the design, implementation and simulation results of a psycholinguistic model of human syntactic processing that meets major empirical criteria. The parser operates in conjunction with a lexicalist grammar and is driven by syntactic information associated with heads of phrases. The dynamics of the model are based on competition by lateral inhibition ('competitive inhibition'). Input words activate lexical frames (i.e. elementary trees anchored to input words) in the mental lexicon, and a network of candidate 'unification links' is set up between frame nodes. These links represent tentative attachments that are graded rather than all-or-none. Candidate links that, due to grammatical or 'treehood' constraints, are incompatible, compete for inclusion in the final syntactic tree by sending each other inhibitory signals that reduce the competitor's attachment strength. The outcome of these local and simultaneous competitions is controlled by dynamic parameters, in particular by the Entry Activation and the Activation Decay rate of syntactic nodes, and by the Strength and Strength Build-up rate of Unification links. In case of a successful parse, a single syntactic tree is returned that covers the whole input string and consists of lexical frames connected by winning Unification links. Simulations are reported of a significant range of psycholinguistic parsing phenomena in both normal and aphasic speakers of English: (i) various effects of linguistic complexity (single versus double, center versus right-hand self-embeddings of relative clauses; the difference between relative clauses with subject and object extraction; the contrast between a complement clause embedded within a relative clause versus a relative clause embedded within a complement clause); (ii) effects of local and global ambiguity, and of word-class and syntactic ambiguity (including recency and length effects); (iii) certain difficulty-of-reanalysis effects (contrasts between local ambiguities that are easy to resolve versus ones that lead to serious garden-path effects); (iv) effects of agrammatism on parsing performance, in particular the performance of various groups of aphasic patients on several sentence types.
  • De Vries, C., Reijnierse, W. G., & Willems, R. M. (2018). Eye movements reveal readers’ sensitivity to deliberate metaphors during narrative reading. Scientific Study of Literature, 8(1), 135-164. doi:10.1075/ssol.18008.vri.

    Abstract

    Metaphors occur frequently in literary texts. Deliberate Metaphor Theory (DMT; e.g., Steen, 2017) proposes that metaphors that serve a communicative function as metaphor are radically different from metaphors that do not have this function. We investigated differences in processing between deliberate and non-deliberate metaphors, compared to non-metaphorical words in literary reading. Using the Deliberate Metaphor Identification Procedure (Reijnierse et al., 2018), we identified metaphors in two literary stories. Then, eye-tracking was used to investigate participants’ (N = 72) reading behavior. Deliberate metaphors were read slower than non-deliberate metaphors, and both metaphor types were read slower than non-metaphorical words. Differences were controlled for several psycholinguistic variables. Differences in reading behavior were related to individual differences in reading experience and absorption and appreciation of the story. These results are in line with predictions from DMT and underline the importance of distinguishing between metaphor types in the experimental study of literary reading.
  • Vromans, R. D., & Jongman, S. R. (2018). The interplay between selective and nonselective inhibition during single word production. PLoS One, 13(5): e0197313. doi:10.1371/journal.pone.0197313.

    Abstract

    The present study investigated the interplay between selective inhibition (the ability to suppress specific competing responses) and nonselective inhibition (the ability to suppress any inappropriate response) during single word production. To this end, we combined two well-established research paradigms: the picture-word interference task and the stop-signal task. Selective inhibition was assessed by instructing participants to name target pictures (e.g., dog) in the presence of semantically related (e.g., cat) or unrelated (e.g., window) distractor words. Nonselective inhibition was tested by occasionally presenting a visual stop-signal, indicating that participants should withhold their verbal response. The stop-signal was presented early (250 ms) aimed at interrupting the lexical selection stage, and late (325 ms) to influence the word-encoding stage of the speech production process. We found longer naming latencies for pictures with semantically related distractors than with unrelated distractors (semantic interference effect). The results further showed that, at both delays, stopping latencies (i.e., stop-signal RTs) were prolonged for naming pictures with semantically related distractors compared to pictures with unrelated distractors. Taken together, our findings suggest that selective and nonselective inhibition, at least partly, share a common inhibitory mechanism during different stages of the speech production process.

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    Data available (link to Figshare)
  • Wain, L. V., Shrine, N., Miller, S., Jackson, V. E., Ntalla, I., Artigas, M. S., Billington, C. K., Kheirallah, A. K., Allen, R., Cook, J. P., Probert, K., Obeidat, M., Bossé, Y., Hao, K., Postma, D. S., Paré, P. D., Ramasamy, A., UK Brain Expression Consortium (UKBEC), Mägi, R., Mihailov, E., Reinmaa, E. and 20 moreWain, L. V., Shrine, N., Miller, S., Jackson, V. E., Ntalla, I., Artigas, M. S., Billington, C. K., Kheirallah, A. K., Allen, R., Cook, J. P., Probert, K., Obeidat, M., Bossé, Y., Hao, K., Postma, D. S., Paré, P. D., Ramasamy, A., UK Brain Expression Consortium (UKBEC), Mägi, R., Mihailov, E., Reinmaa, E., Melén, E., O’Connell, J., Frangou, E., Delaneau, O., OxGSK, C., Freeman, C., Petkova, D., McCarthy, M., Sayers, I., Deloukas, P., Hubbard, R., Pavord, I., Hansell, A. L., Thomson, N. C., Zeggini, E., Morris, A. P., Marchini, J., Strachan, D. P., Tobin, M. D., & Hall, I. P. (2015). Novel insights into the genetics of smoking behaviour, lung function, and chronic obstructive pulmonary disease (UK BiLEVE): a genetic association study in UK Biobank. The Lancet Respiratory Medicine, 3(10), 769-781. doi:10.1016/S2213-2600(15)00283-0.

    Abstract

    Understanding the genetic basis of airflow obstruction and smoking behaviour is key to determining the pathophysiology of chronic obstructive pulmonary disease (COPD). We used UK Biobank data to study the genetic causes of smoking behaviour and lung health.

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  • 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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  • Wang, L., Hagoort, P., & Jensen, O. (2018). Language prediction is reflected by coupling between frontal gamma and posterior alpha oscillations. Journal of Cognitive Neuroscience, 30(3), 432-447. doi:10.1162/jocn_a_01190.

    Abstract

    Readers and listeners actively predict upcoming words during language processing. These predictions might serve to support the unification of incoming words into sentence context and thus rely on interactions between areas in the language network. In the current magnetoencephalography study, participants read sentences that varied in contextual constraints so that the predictability of the sentence-final words was either high or low. Before the sentence-final words, we observed stronger alpha power suppression for the highly compared with low constraining sentences in the left inferior frontal cortex, left posterior temporal region, and visual word form area. Importantly, the temporal and visual word form area alpha power correlated negatively with left frontal gamma power for the highly constraining sentences. We suggest that the correlation between alpha power decrease in temporal language areas and left prefrontal gamma power reflects the initiation of an anticipatory unification process in the language network.
  • Wang, L., Hagoort, P., & Jensen, O. (2018). Gamma oscillatory activity related to language prediction. Journal of Cognitive Neuroscience, 30(8), 1075-1085. doi:10.1162/jocn_a_01275.

    Abstract

    Using magnetoencephalography, the current study examined gamma activity associated with language prediction. Participants read high- and low-constraining sentences in which the final word of the sentence was either expected or unexpected. Although no consistent gamma power difference induced by the sentence-final words was found between the expected and unexpected conditions, the correlation of gamma power during the prediction and activation intervals of the sentence-final words was larger when the presented words matched with the prediction compared with when the prediction was violated or when no prediction was available. This suggests that gamma magnitude relates to the match between predicted and perceived words. Moreover, the expected words induced activity with a slower gamma frequency compared with that induced by unexpected words. Overall, the current study establishes that prediction is related to gamma power correlations and a slowing of the gamma frequency.
  • Wang, M., Shao, Z., Chen, Y., & Schiller, N. O. (2018). Neural correlates of spoken word production in semantic and phonological blocked cyclic naming. Language, Cognition and Neuroscience, 33(5), 575-586. doi:10.1080/23273798.2017.1395467.

    Abstract

    The blocked cyclic naming paradigm has been increasingly employed to investigate the mechanisms underlying spoken word production. Semantic homogeneity typically elicits longer naming latencies than heterogeneity; however, it is debated whether competitive lexical selection or incremental learning underlies this effect. The current study manipulated both semantic and phonological homogeneity and used behavioural and electrophysiological measurements to provide evidence that can distinguish between the two accounts. Results show that naming latencies are longer in semantically homogeneous blocks, but shorter in phonologically homogeneous blocks, relative to heterogeneity. The semantic factor significantly modulates electrophysiological waveforms from 200 ms and the phonological factor from 350 ms after picture presentation. A positive component was demonstrated in both manipulations, possibly reflecting a task-related top-down bias in performing blocked cyclic naming. These results provide novel insights into the neural correlates of blocked cyclic naming and further contribute to the understanding of spoken word production.
  • Wanke, K., Devanna, P., & Vernes, S. C. (2018). Understanding neurodevelopmental disorders: The promise of regulatory variation in the 3’UTRome. Biological Psychiatry, 83(7), 548-557. doi:10.1016/j.biopsych.2017.11.006.

    Abstract

    Neurodevelopmental disorders have a strong genetic component, but despite widespread efforts, the specific genetic factors underlying these disorders remain undefined for a large proportion of affected individuals. Given the accessibility of exome-sequencing, this problem has thus far been addressed from a protein-centric standpoint; however, protein-coding regions only make up ∼1-2% of the human genome. With the advent of whole-genome sequencing we are in the midst of a paradigm shift as it is now possible to interrogate the entire sequence of the human genome (coding and non-coding) to fill in the missing heritability of complex disorders. These new technologies bring new challenges, as the number of non-coding variants identified per individual can be overwhelming, making it prudent to focus on non-coding regions of known function, for which the effects of variation can be predicted and directly tested to assess pathogenicity. The 3’UTRome is a region of the non-coding genome that perfectly fulfils these criteria and is of high interest when searching for pathogenic variation related to complex neurodevelopmental disorders. Herein, we review the regulatory roles of the 3’UTRome as binding sites for microRNAs, RNA binding proteins or during alternative polyadenylation. We detail existing evidence that these regions contribute to neurodevelopmental disorders and outline strategies for identification and validation of novel putatively pathogenic variation in these regions. This evidence suggests that studying the 3’UTRome will lead to the identification of new risk factors, new candidate disease genes and a better understanding of the molecular mechanisms contributing to NDDs.

    Additional information

    1-s2.0-S0006322317321911-mmc1.pdf
  • Warner, N., & Cutler, A. (2017). Stress effects in vowel perception as a function of language-specific vocabulary patterns. Phonetica, 74, 81-106. doi:10.1159/000447428.

    Abstract

    Background/Aims: Evidence from spoken word recognition suggests that for English listeners, distinguishing full versus reduced vowels is important, but discerning stress differences involving the same full vowel (as in mu- from music or museum) is not. In Dutch, in contrast, the latter distinction is important. This difference arises from the relative frequency of unstressed full vowels in the two vocabularies. The goal of this paper is to determine how this difference in the lexicon influences the perception of stressed versus unstressed vowels. Methods: All possible sequences of two segments (diphones) in Dutch and in English were presented to native listeners in gated fragments. We recorded identification performance over time throughout the speech signal. The data were here analysed specifically for patterns in perception of stressed versus unstressed vowels. Results: The data reveal significantly larger stress effects (whereby unstressed vowels are harder to identify than stressed vowels) in English than in Dutch. Both language-specific and shared patterns appear regarding which vowels show stress effects. Conclusion: We explain the larger stress effect in English as reflecting the processing demands caused by the difference in use of unstressed vowels in the lexicon. The larger stress effect in English is due to relative inexperience with processing unstressed full vowels
  • 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.
  • Warrier, V., Chakrabarti, B., Murphy, L., Chan, A., Craig, I., Mallya, U., Lakatošová, S., Rehnstrom, K., Peltonen, L., Wheelwright, S., Allison, C., Fisher, S. E., & Baron-Cohen, S. (2015). A pooled genome-wide association study of Asperger Syndrome. PLoS One, 10(7): e0131202. doi: 10.1371/journal.pone.0131202.

    Abstract

    Asperger Syndrome (AS) is a neurodevelopmental condition characterized by impairments in social interaction and communication, alongside the presence of unusually repetitive, restricted interests and stereotyped behaviour. Individuals with AS have no delay in cognitive and language development. It is a subset of Autism Spectrum Conditions (ASC), which are highly heritable and has a population prevalence of approximately 1%. Few studies have investigated the genetic basis of AS. To address this gap in the literature, we performed a genome-wide pooled DNA association study to identify candidate loci in 612 individuals (294 cases and 318 controls) of Caucasian ancestry, using the Affymetrix GeneChip Human Mapping version 6.0 array. We identified 11 SNPs that had a p-value below 1x10-5. These SNPs were independently genotyped in the same sample. Three of the SNPs (rs1268055, rs7785891 and rs2782448) were nominally significant, though none remained significant after Bonferroni correction. Two of our top three SNPs (rs7785891 and rs2782448) lie in loci previously implicated in ASC. However, investigation of the three SNPs in the ASC genome-wide association dataset from the Psychiatric Genomics Consortium indicated that these three SNPs were not significantly associated with ASC. The effect sizes of the variants were modest, indicating that our study was not sufficiently powered to identify causal variants with precision.
  • Warrington, N. M., Howe, L. D., Paternoster, L., Kaakinen, M., Herrala, S., Huikari, V., Wu, Y. Y., Kemp, J. P., Timpson, N. J., St Pourcain, B., Smith, G. D., Tilling, K., Jarvelin, M.-R., Pennell, C. E., Evans, D. M., Lawlor, D. A., Briollais, L., & Palmer, L. J. (2015). A genome-wide association study of body mass index across early life and childhood. International Journal of Epidemiology, 44(2), 700-712. doi:10.1093/ije/dyv077.

    Abstract

    Background: Several studies have investigated the effect of known adult body mass index (BMI) associated single nucleotide polymorphisms (SNPs) on BMI in childhood. There has been no genome-wide association study (GWAS) of BMI trajectories over childhood.
    Methods: We conducted a GWAS meta-analysis of BMI trajectories from 1 to 17 years of age in 9377 children (77 967 measurements) from the Avon Longitudinal Study of Parents and Children (ALSPAC) and the Western Australian Pregnancy Cohort (Raine) Study. Genome-wide significant loci were examined in a further 3918 individuals (48 530 measurements) from Northern Finland. Linear mixed effects models with smoothing splines were used in each cohort for longitudinal modelling of BMI.
    Results: A novel SNP, downstream from the FAM120AOS gene on chromosome 9, was detected in the meta-analysis of ALSPAC and Raine. This association was driven by a difference in BMI at 8 years (T allele of rs944990 increased BMI; PSNP = 1.52 × 10−8), with a modest association with change in BMI over time (PWald(Change) = 0.006). Three known adult BMI-associated loci (FTO, MC4R and ADCY3) and one childhood obesity locus (OLFM4) reached genome-wide significance (PWald < 1.13 × 10−8) with BMI at 8 years and/or change over time.
    Conclusions: This GWAS of BMI trajectories over childhood identified a novel locus that warrants further investigation. We also observed genome-wide significance with previously established obesity loci, making the novel observation that these loci affected both the level and the rate of change in BMI. We have demonstrated that the use of repeated measures data can increase power to allow detection of genetic loci with smaller sample sizes.
  • Warrington, N. M., Zhu, G., Dy, V., Heath, A. C., Madden, P. A. F., Hemani, G., Kemp, J. P., McMahon, G., St Pourcain, B., Timpson, N. J., Taylor, C. M., Golding, J., Lawlor, D. A., Steer, C., Montgomery, G. W., Martin, N. G., Smith, G. D., Evans, D. M., & Whitfield, J. B. (2015). Genome-wide association study of blood lead shows multiple associations near ALAD. Human Molecular Genetics, 24(13), 3871-3879. doi:10.1093/hmg/ddv112.

    Abstract

    Exposure to high levels of environmental lead, or biomarker evidence of high body lead content, is associated with anaemia, developmental and neurological deficits in children, and increased mortality in adults. Adverse effects of lead still occur despite substantial reduction in environmental exposure. There is genetic variation between individuals in blood lead concentration but the polymorphisms contributing to this have not been defined. We measured blood or erythrocyte lead content, and carried out genome-wide association analysis, on population-based cohorts of adult volunteers from Australia and UK (N = 5433). Samples from Australia were collected in two studies, in 1993–1996 and 2002–2005 and from UK in 1991–1992. One locus, at ALAD on chromosome 9, showed consistent association with blood lead across countries and evidence for multiple independent allelic effects. The most significant single nucleotide polymorphism (SNP), rs1805313 (P = 3.91 × 10−14 for lead concentration in a meta-analysis of all data), is known to have effects on ALAD expression in blood cells but other SNPs affecting ALAD expression did not affect blood lead. Variants at 12 other loci, including ABO, showed suggestive associations (5 × 10−6 >} P {> 5 × 10−8). Identification of genetic polymorphisms affecting blood lead reinforces the view that genetic factors, as well as environmental ones, are important in determining blood lead levels. The ways in which ALAD variation affects lead uptake or distribution are still to be determined.
  • Watson, L. M., Wong, M. M. K., Vowles, J., Cowley, S. A., & Becker, E. B. E. (2018). A simplified method for generating purkinje cells from human-induced pluripotent stem cells. The Cerebellum, 17(4), 419-427. doi:10.1007/s12311-017-0913-2.

    Abstract

    The establishment of a reliable model for the study of Purkinje cells in vitro is of particular importance, given their central role in cerebellar function and pathology. Recent advances in induced pluripotent stem cell (iPSC) technology offer the opportunity to generate multiple neuronal subtypes for study in vitro. However, to date, only a handful of studies have generated Purkinje cells from human pluripotent stem cells, with most of these protocols proving challenging to reproduce. Here, we describe a simplified method for the reproducible generation of Purkinje cells from human iPSCs. After 21 days of treatment with factors selected to mimic the self-inductive properties of the isthmic organiser—insulin, fibroblast growth factor 2 (FGF2), and the transforming growth factor β (TGFβ)-receptor blocker SB431542—hiPSCs could be induced to form En1-positive cerebellar progenitors at efficiencies of up to 90%. By day 35 of differentiation, subpopulations of cells representative of the two cerebellar germinal zones, the rhombic lip (Atoh1-positive) and ventricular zone (Ptf1a-positive), could be identified, with the latter giving rise to cells positive for Purkinje cell progenitor-specific markers, including Lhx5, Kirrel2, Olig2 and Skor2. Further maturation was observed following dissociation and co-culture of these cerebellar progenitors with mouse cerebellar cells, with 10% of human cells staining positive for the Purkinje cell marker calbindin by day 70 of differentiation. This protocol, which incorporates modifications designed to enhance cell survival and maturation and improve the ease of handling, should serve to make existing models more accessible, in order to enable future advances in the field.

    Additional information

    12311_2017_913_MOESM1_ESM.docx
  • Watson, L. M., Wong, M. M. K., & Becker, E. B. E. (2015). Induced pluripotent stem cell technology for modelling and therapy of cerebellar ataxia. Open Biology, 5: 150056. doi:10.1098/rsob.150056.

    Abstract

    Induced pluripotent stem cell (iPSC) technology has emerged as an important tool in understanding, and potentially reversing, disease pathology. This is particularly true in the case of neurodegenerative diseases, in which the affected cell types are not readily accessible for study. Since the first descriptions of iPSC-based disease modelling, considerable advances have been made in understanding the aetiology and progression of a diverse array of neurodegenerative conditions, including Parkinson's disease and Alzheimer's disease. To date, however, relatively few studies have succeeded in using iPSCs to model the neurodegeneration observed in cerebellar ataxia. Given the distinct neurodevelopmental phenotypes associated with certain types of ataxia, iPSC-based models are likely to provide significant insights, not only into disease progression, but also to the development of early-intervention therapies. In this review, we describe the existing iPSC-based disease models of this heterogeneous group of conditions and explore the challenges associated with generating cerebellar neurons from iPSCs, which have thus far hindered the expansion of this research.
  • 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
  • Weekes, B. S., Abutalebi, J., Mak, H.-K.-F., Borsa, V., Soares, S. M. P., Chiu, P. W., & Zhang, L. (2018). Effect of monolingualism and bilingualism in the anterior cingulate cortex: a proton magnetic resonance spectroscopy study in two centers. Letras de Hoje, 53(1), 5-12. doi:10.15448/1984-7726.2018.1.30954.

    Abstract

    Reports of an advantage of bilingualism on brain structure in young adult participants
    are inconsistent. Abutalebi et al. (2012) reported more efficient monitoring of conflict during the
    Flanker task in young bilinguals compared to young monolingual speakers. The present study
    compared young adult (mean age = 24) Cantonese-English bilinguals in Hong Kong and young
    adult monolingual speakers. We expected (a) differences in metabolites in neural tissue to result
    from bilingual experience, as measured by 1H-MRS at 3T, (b) correlations between metabolic
    levels and Flanker conflict and interference effects (c) different associations in bilingual and
    monolingual speakers. We found evidence of metabolic differences in the ACC due to bilingualism,
    specifically in metabolites Cho, Cr, Glx and NAA. However, we found no significant correlations
    between metabolic levels and conflict and interference effects and no significant evidence of
    differential relationships between bilingual and monolingual speakers. Furthermore, we found no
    evidence of significant differences in the mean size of conflict and interference effects between
    groups i.e. no bilingual advantage. Lower levels of Cho, Cr, Glx and NAA in bilingual adults
    compared to monolingual adults suggest that the brains of bilinguals develop greater adaptive
    control during conflict monitoring because of their extensive bilingual experience.
  • Wegman, J., Tyborowska, A., Hoogman, M., Vasquez, A. A., & Janzen, G. (2017). The brain-derived neurotrophic factor Val66Met polymorphism affects encoding of object locations during active navigation. European Journal of Neuroscience, 45(12), 1501-1511. doi:10.1111/ejn.13416.

    Abstract

    The brain-derived neurotrophic factor (BDNF) was shown to be involved in spatial memory and spatial strategy preference. A naturally occurring single nucleotide polymorphism of the BDNF gene (Val66Met) affects activity-dependent secretion of BDNF. The current event-related fMRI study on preselected groups of ‘Met’ carriers and homozygotes of the ‘Val’ allele investigated the role of this polymorphism on encoding and retrieval in a virtual navigation task in 37 healthy volunteers. In each trial, participants navigated toward a target object. During encoding, three positional cues (columns) with directional cues (shadows) were available. During retrieval, the invisible target had to be replaced while either two objects without shadows (objects trial) or one object with a shadow (shadow trial) were available. The experiment consisted of blocks, informing participants of which trial type would be most likely to occur during retrieval. We observed no differences between genetic groups in task performance or time to complete the navigation tasks. The imaging results show that Met carriers compared to Val homozygotes activate the left hippocampus more during successful object location memory encoding. The observed effects were independent of non-significant performance differences or volumetric differences in the hippocampus. These results indicate that variations of the BDNF gene affect memory encoding during spatial navigation, suggesting that lower levels of BDNF in the hippocampus results in less efficient spatial memory processing
  • Whelpton, M., Guðmundsdóttir Beck, þ., & Jordan, F. (2015). The semantics and morphology of household container names in Icelandic and Dutch. Language Sciences, 49, 67-81. doi:10.1016/j.langsci.2014.07.014.

    Abstract

    In this paper, we report an experiment on the naming of household containers in Dutch and Icelandic carried out as part of the Evolution of Semantic Systems project (EoSS; Majid et al., 2011). This naming experiment allows us to support and elaborate on a hypothesis by Malt et al. (2003) that productive morphology in the naming domain can have an influence on boundary placement within the extensional space. Specifically, we demonstrate that the Dutch diminutive -(t)je favours a cut between small items versus others, whereas Icelandic, which does not use the diminutive in this domain, favours a cut between large items and others. This is not a typological effect, as Dutch and Icelandic are both Germanic languages and both have diminutive morphology available in principle. We find no evidence that the diminutive produces a proliferation of terms and/or fine-grained nesting within the extensional domain. Rather, the Dutch diminutive favours a more even distribution of terms across the space whereas Icelandic favours broad inclusive terms with a number of narrower specialist terms. Further, the extensional space defined by the diminutive is not associated with its own clear prototypical exemplar. Using evidence from compounding and modification, we also consider which semantic features are prominent in differentiating categories within the domain. By far the most prominent in both languages is the inferred contents of the container. Other than contents, however, the languages differ in the range and prominence of features such as intended usage or material of composition. Our results demonstrate that in order to understand the processes that produce semantic divisions of basic object classes, we should consider fine-grained analyses of closely related languages alongside analyses of typologically different languages.
  • Wiese, R., Orzechowska, P., Alday, P. M., & Ulbrich, C. (2017). Structural Principles or Frequency of Use? An ERP Experiment on the Learnability of Consonant Clusters. Frontiers in Psychology, 7: 2005. doi:10.3389/fpsyg.2016.02005.

    Abstract

    Phonological knowledge of a language involves knowledge about which segments can be combined under what conditions. Languages vary in the quantity and quality of licensed combinations, in particular sequences of consonants, with Polish being a language with a large inventory of such combinations. The present paper reports on a two-session experiment in which Polish-speaking adult participants learned nonce words with final consonant clusters. The aim was to study the role of two factors which potentially play a role in the learning of phonotactic structures: the phonological principle of sonority (ordering sound segments within the syllable according to their inherent loudness) and the (non-) existence as a usage-based phenomenon. EEG responses in two different time windows (adversely to behavioral responses) show linguistic processing by native speakers of Polish to be sensitive to both distinctions, in spite of the fact that Polish is rich in sonority-violating clusters. In particular, a general learning effect in terms of an N400 effect was found which was demonstrated to be different for sonority-obeying clusters than for sonority-violating clusters. Furthermore, significant interactions of formedness and session, and of existence and session, demonstrate that both factors, the sonority principle and the frequency pattern, play a role in the learning process.
  • Winsvold, B. S., Palta, P., Eising, E., Page, C. M., The International Headache Genetics Consortium, Van den Maagdenberg, A. M. J. M., Palotie, A., & Zwart, J.-A. (2018). Epigenetic DNA methylation changes associated with headache chronification: A retrospective case-control study. Cephalalgia, 38(2), 312-322. doi:10.1177/0333102417690111.

    Abstract

    Background

    The biological mechanisms of headache chronification are poorly understood. We aimed to identify changes in DNA methylation associated with the transformation from episodic to chronic headache.
    Methods

    Participants were recruited from the population-based Norwegian HUNT Study. Thirty-six female headache patients who transformed from episodic to chronic headache between baseline and follow-up 11 years later were matched against 35 controls with episodic headache. DNA methylation was quantified at 485,000 CpG sites, and changes in methylation level at these sites were compared between cases and controls by linear regression analysis. Data were analyzed in two stages (Stages 1 and 2) and in a combined meta-analysis.
    Results

    None of the top 20 CpG sites identified in Stage 1 replicated in Stage 2 after multiple testing correction. In the combined meta-analysis the strongest associated CpG sites were related to SH2D5 and NPTX2, two brain-expressed genes involved in the regulation of synaptic plasticity. Functional enrichment analysis pointed to processes including calcium ion binding and estrogen receptor pathways.
    Conclusion

    In this first genome-wide study of DNA methylation in headache chronification several potentially implicated loci and processes were identified. The study exemplifies the use of prospectively collected population cohorts to search for epigenetic mechanisms of disease
  • Winter, B., Perlman, M., & Majid, A. (2018). Vision dominates in perceptual language: English sensory vocabulary is optimized for usage. Cognition, 179, 213-220. doi:10.1016/j.cognition.2018.05.008.

    Abstract

    Researchers have suggested that the vocabularies of languages are oriented towards the communicative needs of language users. Here, we provide evidence demonstrating that the higher frequency of visual words in a large variety of English corpora is reflected in greater lexical differentiation—a greater number of unique words—for the visual domain in the English lexicon. In comparison, sensory modalities that are less frequently talked about, particularly taste and smell, show less lexical differentiation. In addition, we show that even though sensory language can be expected to change across historical time and between contexts of use (e.g., spoken language versus fiction), the pattern of visual dominance is a stable property of the English language. Thus, we show that across the board, precisely those semantic domains that are more frequently talked about are also more lexically differentiated, for perceptual experiences. This correlation between type and token frequencies suggests that the sensory lexicon of English is geared towards communicative efficiency.
  • Wirthlin, M., Chang, E. F., Knörnschild, M., Krubitzer, L. A., Mello, C. V., Miller, C. T., Pfenning, A. R., Vernes, S. C., Tchernichovski, O., & Yartsev, M. M. (2019). A modular approach to vocal learning: Disentangling the diversity of a complex behavioral trait. Neuron, 104(1), 87-99. doi:10.1016/j.neuron.2019.09.036.

    Abstract

    Vocal learning is a behavioral trait in which the social and acoustic environment shapes the vocal repertoire of individuals. Over the past century, the study of vocal learning has progressed at the intersection of ecology, physiology, neuroscience, molecular biology, genomics, and evolution. Yet, despite the complexity of this trait, vocal learning is frequently described as a binary trait, with species being classified as either vocal learners or vocal non-learners. As a result, studies have largely focused on a handful of species for which strong evidence for vocal learning exists. Recent studies, however, suggest a continuum in vocal learning capacity across taxa. Here, we further suggest that vocal learning is a multi-component behavioral phenotype comprised of distinct yet interconnected modules. Discretizing the vocal learning phenotype into its constituent modules would facilitate integration of findings across a wider diversity of species, taking advantage of the ways in which each excels in a particular module, or in a specific combination of features. Such comparative studies can improve understanding of the mechanisms and evolutionary origins of vocal learning. We propose an initial set of vocal learning modules supported by behavioral and neurobiological data and highlight the need for diversifying the field in order to disentangle the complexity of the vocal learning phenotype.

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  • De Wit, S. J., van der Werf, Y. D., Mataix-Cols, D., Trujillo, J. P., van Oppen, P., Veltman, D. J., & van den Heuvel, O. A. (2015). Emotion regulation before and after transcranial magnetic stimulation in obsessive compulsive disorder. Psychological Medicine, 45(14), 3059-3073. doi:10.1017/S0033291715001026.

    Abstract

    Impaired emotion regulation may underlie exaggerated emotional reactivity in patients with obsessive compulsive disorder (OCD), yet instructed emotion regulation has never been studied in the disorder. METHOD: This study aimed to assess the neural correlates of emotion processing and regulation in 43 medication-free OCD patients and 38 matched healthy controls, and additionally test if these can be modulated by stimulatory (patients) and inhibitory (controls) repetitive transcranial magnetic stimulation (rTMS) over the left dorsolateral prefrontal cortex (dlPFC). Participants performed an emotion regulation task during functional magnetic resonance imaging before and after a single session of randomly assigned real or sham rTMS. Effect of group and rTMS were assessed on self-reported distress ratings and brain activity in frontal-limbic regions of interest. RESULTS: Patients had higher distress ratings than controls during emotion provocation, but similar rates of distress reduction after voluntary emotion regulation. OCD patients compared with controls showed altered amygdala responsiveness during symptom provocation and diminished left dlPFC activity and frontal-amygdala connectivity during emotion regulation. Real v. sham dlPFC stimulation differentially modulated frontal-amygdala connectivity during emotion regulation in OCD patients. CONCLUSIONS: We propose that the increased emotional reactivity in OCD may be due to a deficit in emotion regulation caused by a failure of cognitive control exerted by the dorsal frontal cortex. Modulatory rTMS over the left dlPFC may influence automatic emotion regulation capabilities by influencing frontal-limbic connectivity.
  • Witteman, M. J., Bardhan, N. P., Weber, A., & McQueen, J. M. (2015). Automaticity and stability of adaptation to foreign-accented speech. Language and Speech, 52(2), 168-189. doi:10.1177/0023830914528102.

    Abstract

    In three cross-modal priming experiments we asked whether adaptation to a foreign-accented speaker is automatic, and whether adaptation can be seen after a long delay between initial exposure and test. Dutch listeners were exposed to a Hebrew-accented Dutch speaker with two types of Dutch words: those that contained [ɪ] (globally accented words), and those in which the Dutch [i] was shortened to [ɪ] (specific accent marker words). Experiment 1, which served as a baseline, showed that native Dutch participants showed facilitatory priming for globally accented, but not specific accent, words. In experiment 2, participants performed a 3.5-minute phoneme monitoring task, and were tested on their comprehension of the accented speaker 24 hours later using the same cross-modal priming task as in experiment 1. During the phoneme monitoring task, listeners were asked to detect a consonant that was not strongly accented. In experiment 3, the delay between exposure and test was extended to 1 week. Listeners in experiments 2 and 3 showed facilitatory priming for both globally accented and specific accent marker words. Together, these results show that adaptation to a foreign-accented speaker can be rapid and automatic, and can be observed after a prolonged delay in testing.
  • Wnuk, E., De Valk, J. M., Huisman, J. L. A., & Majid, A. (2017). Hot and cold smells: Odor-temperature associations across cultures. Frontiers in Psychology, 8: 1373. doi:10.3389/fpsyg.2017.01373.

    Abstract

    It is often assumed odors are associated with hot and cold temperature, since odor processing may trigger thermal sensations, such as coolness in the case of mint. It is unknown, however, whether people make consistent temperature associations for a variety of everyday odors, and, if so, what determines them. Previous work investigating the bases of cross-modal associations suggests a number of possibilities, including universal forces (e.g., perception), as well as culture-specific forces (e.g., language and cultural beliefs). In this study, we examined odor-temperature associations in three cultures—Maniq (N = 11), Thai (N = 24), and Dutch (N = 24)—who differ with respect to their cultural preoccupation with odors, their odor lexicons, and their beliefs about the relationship of odors (and odor objects) to temperature. Participants matched 15 odors to temperature by touching cups filled with hot or cold water, and described the odors in their native language. The results showed no consistent associations among the Maniq, and only a handful of consistent associations between odor and temperature among the Thai and Dutch. The consistent associations differed across the two groups, arguing against their universality. Further analysis revealed cross-modal associations could not be explained by language, but could be the result of cultural beliefs
  • Li, Q., Wojciechowski, R., Simpson, C. L., Hysi, P. G., Verhoeven, V. J. M., Ikram, M. K., Höhn, R., Vitart, V., Hewitt, A. W., Oexle, K., Mäkelä, K.-M., MacGregor, S., Pirastu, M., Fan, Q., Cheng, C.-Y., St Pourcain, B., McMahon, G., Kemp, J. P., Northstone, K., Rahi, J. S. and 69 moreLi, Q., Wojciechowski, R., Simpson, C. L., Hysi, P. G., Verhoeven, V. J. M., Ikram, M. K., Höhn, R., Vitart, V., Hewitt, A. W., Oexle, K., Mäkelä, K.-M., MacGregor, S., Pirastu, M., Fan, Q., Cheng, C.-Y., St Pourcain, B., McMahon, G., Kemp, J. P., Northstone, K., Rahi, J. S., Cumberland, P. M., Martin, N. G., Sanfilippo, P. G., Lu, Y., Wang, Y. X., Hayward, C., Polašek, O., Campbell, H., Bencic, G., Wright, A. F., Wedenoja, J., Zeller, T., Schillert, A., Mirshahi, A., Lackner, K., Yip, S. P., Yap, M. K. H., Ried, J. S., Gieger, C., Murgia, F., Wilson, J. F., Fleck, B., Yazar, S., Vingerling, J. R., Hofman, A., Uitterlinden, A., Rivadeneira, F., Amin, N., Karssen, L., Oostra, B. A., Zhou, X., Teo, Y.-Y., Tai, E. S., Vithana, E., Barathi, V., Zheng, Y., Siantar, R. G., Neelam, K., Shin, Y., Lam, J., Yonova-Doing, E., Venturini, C., Hosseini, S. M., Wong, H.-S., Lehtimäki, T., Kähönen, M., Raitakari, O., Timpson, N. J., Evans, D. M., Khor, C.-C., Aung, T., Young, T. L., Mitchell, P., Klein, B., van Duijn, C. M., Meitinger, T., Jonas, J. B., Baird, P. N., Mackey, D. A., Wong, T. Y., Saw, S.-M., Pärssinen, O., Stambolian, D., Hammond, C. J., Klaver, C. C. W., Williams, C., Paterson, A. D., Bailey-Wilson, J. E., & Guggenheim, J. A. (2015). Genome-wide association study for refractive astigmatism reveals genetic co-determination with spherical equivalent refractive error: the CREAM consortium. Human Genetics, 134, 131-146. doi:10.1007/s00439-014-1500-y.

    Abstract

    To identify genetic variants associated with refractive astigmatism in the general population, meta-analyses of genome-wide association studies were performed for: White Europeans aged at least 25 years (20 cohorts, N = 31,968); Asian subjects aged at least 25 years (7 cohorts, N = 9,295); White Europeans aged <25 years (4 cohorts, N = 5,640); and all independent individuals from the above three samples combined with a sample of Chinese subjects aged <25 years (N = 45,931). Participants were classified as cases with refractive astigmatism if the average cylinder power in their two eyes was at least 1.00 diopter and as controls otherwise. Genome-wide association analysis was carried out for each cohort separately using logistic regression. Meta-analysis was conducted using a fixed effects model. In the older European group the most strongly associated marker was downstream of the neurexin-1 (NRXN1) gene (rs1401327, P = 3.92E−8). No other region reached genome-wide significance, and association signals were lower for the younger European group and Asian group. In the meta-analysis of all cohorts, no marker reached genome-wide significance: The most strongly associated regions were, NRXN1 (rs1401327, P = 2.93E−07), TOX (rs7823467, P = 3.47E−07) and LINC00340 (rs12212674, P = 1.49E−06). For 34 markers identified in prior GWAS for spherical equivalent refractive error, the beta coefficients for genotype versus spherical equivalent, and genotype versus refractive astigmatism, were highly correlated (r = −0.59, P = 2.10E−04). This work revealed no consistent or strong genetic signals for refractive astigmatism; however, the TOX gene region previously identified in GWAS for spherical equivalent refractive error was the second most strongly associated region. Analysis of additional markers provided evidence supporting widespread genetic co-susceptibility for spherical and astigmatic refractive errors.
  • Wolf, M. C. (2015). Het verschil tussen hardop en stillezen wat betreft leessnelheid en tekstbegrip en de invloed hierop van fonologisch bewustzijn, benoemsnelheid en visuele aandachtsspanne. Student Undergraduate Research E-journal, 1(1), 261-264. Retrieved from http://journals.library.tudelft.nl/index.php/sure/article/view/1025.

    Abstract

    In het onderwijs wordt aangenomen dat hardop en stillezen dezelfde processen zijn. In dit onderzoek wordt gekeken naar het verschil tussen hardop en stillezen wat betreft leessnelheid en tekstbegrip bij 90 kinderen uit groep 4. Ook wordt de invloed van de cognitieve vaardigheden fonologisch bewustzijn, benoemsnelheid en visuele aandachtsspanne op de verschillende leesmodi onderzocht. De participanten lazen stil sneller, maar begrepen de tekst beter hardop. De cognitieve vaardigheden correleerden met hardop en stillezen wat betreft leessnelheid, maar hingen in beide leesmodi niet samen met tekstbegrip. Hoewel hardop en stillezen samenhangen, onderstrepen deze bevindingen dat het verschillende leesmodi zijn.
  • Wolf, M. C., Muijselaar, M. M. L., Boonstra, A. M., & De Bree, E. H. (2019). The relationship between reading and listening comprehension: Shared and modality-specific components. Reading and Writing, 32(7), 1747-1767. doi:10.1007/s11145-018-9924-8.

    Abstract

    This study aimed to increase our understanding on the relationship between reading and listening comprehension. Both in comprehension theory and in educational practice, reading and listening comprehension are often seen as interchangeable, overlooking modality-specific aspects of them separately. Three questions were addressed. First, it was examined to what extent reading and listening comprehension comprise modality-specific, distinct skills or an overlapping, domain-general skill in terms of the amount of explained variance in one comprehension type by the opposite comprehension type. Second, general and modality-unique subskills of reading and listening comprehension were sought by assessing the contributions of the foundational skills word reading fluency, vocabulary, memory, attention, and inhibition to both comprehension types. Lastly, the practice of using either listening comprehension or vocabulary as a proxy of general comprehension was investigated. Reading and listening comprehension tasks with the same format were assessed in 85 second and third grade children. Analyses revealed that reading comprehension explained 34% of the variance in listening comprehension, and listening comprehension 40% of reading comprehension. Vocabulary and word reading fluency were found to be shared contributors to both reading and listening comprehension. None of the other cognitive skills contributed significantly to reading or listening comprehension. These results indicate that only part of the comprehension process is indeed domain-general and not influenced by the modality in which the information is provided. Especially vocabulary seems to play a large role in this domain-general part. The findings warrant a more prominent focus of modality-specific aspects of both reading and listening comprehension in research and education.
  • Wong, M. M. K., Hoekstra, S. D., Vowles, J., Watson, L. M., Fuller, G., Németh, A. H., Cowley, S. A., Ansorge, O., Talbot, K., & Becker, E. B. E. (2018). Neurodegeneration in SCA14 is associated with increased PKCγ kinase activity, mislocalization and aggregation. Acta Neuropathologica Communications, 6: 99. doi:10.1186/s40478-018-0600-7.

    Abstract

    Spinocerebellar ataxia type 14 (SCA14) is a subtype of the autosomal dominant cerebellar ataxias that is characterized by slowly progressive cerebellar dysfunction and neurodegeneration. SCA14 is caused by mutations in the PRKCG gene, encoding protein kinase C gamma (PKCγ). Despite the identification of 40 distinct disease-causing mutations in PRKCG, the pathological mechanisms underlying SCA14 remain poorly understood. Here we report the molecular neuropathology of SCA14 in post-mortem cerebellum and in human patient-derived induced pluripotent stem cells (iPSCs) carrying two distinct SCA14 mutations in the C1 domain of PKCγ, H36R and H101Q. We show that endogenous expression of these mutations results in the cytoplasmic mislocalization and aggregation of PKCγ in both patient iPSCs and cerebellum. PKCγ aggregates were not efficiently targeted for degradation. Moreover, mutant PKCγ was found to be hyper-activated, resulting in increased substrate phosphorylation. Together, our findings demonstrate that a combination of both, loss-of-function and gain-of-function mechanisms are likely to underlie the pathogenesis of SCA14, caused by mutations in the C1 domain of PKCγ. Importantly, SCA14 patient iPSCs were found to accurately recapitulate pathological features observed in post-mortem SCA14 cerebellum, underscoring their potential as relevant disease models and their promise as future drug discovery tools.

    Additional information

    additional file
  • Wong, M. M. K., Watson, L. M., & Becker, E. B. E. (2017). Recent advances in modelling of cerebellar ataxia using induced pluripotent stem cells. Journal of Neurology & Neuromedicine, 2(7), 11-15. doi:10.29245/2572.942X/2017/7.1134.

    Abstract

    The cerebellar ataxias are a group of incurable brain disorders that are caused primarily by the progressive dysfunction and degeneration of cerebellar Purkinje cells. The lack of reliable disease models for the heterogeneous ataxias has hindered the understanding of the underlying pathogenic mechanisms as well as the development of effective therapies for these devastating diseases. Recent advances in the field of induced pluripotent stem cell (iPSC) technology offer new possibilities to better understand and potentially reverse disease pathology. Given the neurodevelopmental phenotypes observed in several types of ataxias, iPSC-based models have the potential to provide significant insights into disease progression, as well as opportunities for the development of early intervention therapies. To date, however, very few studies have successfully used iPSC-derived cells to cerebellar ataxias. In this review, we focus on recent breakthroughs in generating human iPSC-derived Purkinje cells. We also highlight the future challenges that will need to be addressed in order to fully exploit these models for the modelling of the molecular mechanisms underlying cerebellar ataxias and the development of effective therapeutics.
  • 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
  • Yager, J., & Burenhult, N. (2017). Jedek: a newly discovered Aslian variety of Malaysia. Linguistic Typology, 21(3), 493-545. doi:10.1515/lingty-2017-0012.

    Abstract

    Jedek is a previously unrecognized variety of the Northern Aslian subgroup of the Aslian branch of the Austroasiatic language family. It is spoken by c. 280 individuals in the resettlement area of Sungai Rual, near Jeli in Kelantan state, Peninsular Malaysia. The community originally consisted of several bands of foragers along the middle reaches of the Pergau river. Jedek’s distinct status first became known during a linguistic survey carried out in the DOBES project Tongues of the Semang (2005-2011). This paper describes the process leading up to its discovery and provides an overview of its typological characteristics.
  • Yang, J., Zhu, H., & Tian, X. (2018). Group-level multivariate analysis in EasyEEG toolbox: Examining the temporal dynamics using topographic responses. Frontiers in Neuroscience, 12: 468. doi:10.3389/fnins.2018.00468.

    Abstract

    Electroencephalography (EEG) provides high temporal resolution cognitive information from non-invasive recordings. However, one of the common practices-using a subset of sensors in ERP analysis is hard to provide a holistic and precise dynamic results. Selecting or grouping subsets of sensors may also be subject to selection bias, multiple comparison, and further complicated by individual differences in the group-level analysis. More importantly, changes in neural generators and variations in response magnitude from the same neural sources are difficult to separate, which limit the capacity of testing different aspects of cognitive hypotheses. We introduce EasyEEG, a toolbox that includes several multivariate analysis methods to directly test cognitive hypotheses based on topographic responses that include data from all sensors. These multivariate methods can investigate effects in the dimensions of response magnitude and topographic patterns separately using data in the sensor space, therefore enable assessing neural response dynamics. The concise workflow and the modular design provide user-friendly and programmer-friendly features. Users of all levels can benefit from the open-sourced, free EasyEEG to obtain a straightforward solution for efficient processing of EEG data and a complete pipeline from raw data to final results for publication.
  • Yoshihara, M., Nakayama, M., Verdonschot, R. G., & Hino, Y. (2017). The phonological unit of Japanese Kanji compounds: A masked priming investigation. Journal of Experimental Psychology: Human Perception and Performance, 43(7), 1303-1328. doi:10.1037/xhp0000374.

    Abstract

    Using the masked priming paradigm, we examined which phonological unit is used when naming Kanji compounds. Although the phonological unit in the Japanese language has been suggested to be the mora, Experiment 1 found no priming for mora-related Kanji prime-target pairs. In Experiment 2, significant priming was only found when Kanji pairs shared the whole sound of their initial Kanji characters. Nevertheless, when the same Kanji pairs used in Experiment 2 were transcribed into Kana, significant mora priming was observed in Experiment 3. In Experiment 4, matching the syllable structure and pitch-accent of the initial Kanji characters did not lead to mora priming, ruling out potential alternative explanations for the earlier absence of the effect. A significant mora priming effect was observed, however, when the shared initial mora constituted the whole sound of their initial Kanji characters in Experiments 5. Lastly, these results were replicated in Experiment 6. Overall, these results indicate that the phonological unit involved when naming Kanji compounds is not the mora but the whole sound of each Kanji character. We discuss how different phonological units may be involved when processing Kanji and Kana words as well as the implications for theories dealing with language production processes.
  • Zhao, H., Zhou, W., Yao, Z., Wan, Y., Cao, J., Zhang, L., Zhao, J., Li, H., Zhou, R., Li, B., Wei, G., Zhang, Z., French, C. A., Dekker, J. D., Yang, Y., Fisher, S. E., Tucker, H. O., & Guo, X. (2015). Foxp1/2/4 regulate endochondral ossification as a suppresser complex. Developmental Biology, 398, 242-254. doi:10.1016/j.ydbio.2014.12.007.

    Abstract

    Osteoblast induction and differentiation in developing long bones is dynamically controlled by the opposing action of transcriptional activators and repressors. In contrast to the long list of activators that have been discovered over past decades, the network of repressors is not well-defined. Here we identify the expression of Foxp1/2/4 proteins, comprised of Forkhead-box (Fox) transcription factors of the Foxp subfamily, in both perichondrial skeletal progenitors and proliferating chondrocytes during endochondral ossification. Mice carrying loss-of-function and gain-of-function Foxp mutations had gross defects in appendicular skeleton formation. At the cellular level, over-expression of Foxp1/2/4 in chondroctyes abrogated osteoblast formation and chondrocyte hypertrophy. Conversely, single or compound deficiency of Foxp1/2/4 in skeletal progenitors or chondrocytes resulted in premature osteoblast differentiation in the perichondrium, coupled with impaired proliferation, survival, and hypertrophy of chondrocytes in the growth plate. Foxp1/2/4 and Runx2 proteins interacted in vitro and in vivo, and Foxp1/2/4 repressed Runx2 transactivation function in heterologous cells. This study establishes Foxp1/2/4 proteins as coordinators of osteogenesis and chondrocyte hypertrophy in developing long bones and suggests that a novel transcriptional repressor network involving Foxp1/2/4 may regulate Runx2 during endochondral ossification.
  • Zhen, Z., Yang, Z., Huang, L., Kong, X., Wang, X., Dang, X., Huang, Y., Song, Y., & Liu, J. (2015). Quantifying interindividual variability and asymmetry of face-selective regions: A probabilistic functional atlas. NeuroImage, 113, 13-25. doi:10.1016/j.neuroimage.2015.03.010.

    Abstract

    Face-selective regions (FSRs) are among the most widely studied functional regions in the human brain. However, individual variability of the FSRs has not been well quantified. Here we use functional magnetic resonance imaging (fMRI) to localize the FSRs and quantify their spatial and functional variabilities in 202 healthy adults. The occipital face area (OFA), posterior and anterior fusiform face areas (pFFA and aFFA), posterior continuation of the superior temporal sulcus (pcSTS), and posterior and anterior STS (pSTS and aSTS) were delineated for each individual with a semi-automated procedure. A probabilistic atlas was constructed to characterize their interindividual variability, revealing that the FSRs were highly variable in location and extent across subjects. The variability of FSRs was further quantified on both functional (i.e., face selectivity) and spatial (i.e., volume, location of peak activation, and anatomical location) features. Considerable interindividual variability and rightward asymmetry were found in all FSRs on these features. Taken together, our work presents the first effort to characterize comprehensively the variability of FSRs in a large sample of healthy subjects, and invites future work on the origin of the variability and its relation to individual differences in behavioral performance. Moreover, the probabilistic functional atlas will provide an adequate spatial reference for mapping the face network.
  • Zhen, Z., Kong, X., Huang, L., Yang, Z., Wang, X., Hao, X., Huang, T., Song, Y., & Liu, J. (2017). Quantifying the variability of scene-selective regions: Interindividual, interhemispheric, and sex differences. Human Brain Mapping, 38(4), 2260-2275. doi:10.1002/hbm.23519.

    Abstract

    Scene-selective regions (SSRs), including the parahippocampal place area (PPA), retrosplenial cortex (RSC), and transverse occipital sulcus (TOS), are among the most widely characterized functional regions in the human brain. However, previous studies have mostly focused on the commonality within each SSR, providing little information on different aspects of their variability. In a large group of healthy adults (N = 202), we used functional magnetic resonance imaging to investigate different aspects of topographical and functional variability within SSRs, including interindividual, interhemispheric, and sex differences. First, the PPA, RSC, and TOS were delineated manually for each individual. We then demonstrated that SSRs showed substantial interindividual variability in both spatial topography and functional selectivity. We further identified consistent interhemispheric differences in the spatial topography of all three SSRs, but distinct interhemispheric differences in scene selectivity. Moreover, we found that all three SSRs showed stronger scene selectivity in men than in women. In summary, our work thoroughly characterized the interindividual, interhemispheric, and sex variability of the SSRs and invites future work on the origin and functional significance of these variabilities. Additionally, we constructed the first probabilistic atlases for the SSRs, which provide the detailed anatomical reference for further investigations of the scene network.
  • Zheng, X., Roelofs, A., Farquhar, J., & Lemhöfer, K. (2018). Monitoring of language selection errors in switching: Not all about conflict. PLoS One, 13(11): e0200397. doi:10.1371/journal.pone.0200397.

    Abstract

    Although bilingual speakers are very good at selectively using one language rather than another, sometimes language selection errors occur. To investigate how bilinguals monitor their speech errors and control their languages in use, we recorded event-related potentials (ERPs) in unbalanced Dutch-English bilingual speakers in a cued language-switching task. We tested the conflict-based monitoring model of Nozari and colleagues by investigating the error-related negativity (ERN) and comparing the effects of the two switching directions (i.e., to the first language, L1 vs. to the second language, L2). Results show that the speakers made more language selection errors when switching from their L2 to the L1 than vice versa. In the EEG, we observed a robust ERN effect following language selection errors compared to correct responses, reflecting monitoring of speech errors. Most interestingly, the ERN effect was enlarged when the speakers were switching to their L2 (less conflict) compared to switching to the L1 (more conflict). Our findings do not support the conflict-based monitoring model. We discuss an alternative account in terms of error prediction and reinforcement learning.
  • Zheng, X., Roelofs, A., & Lemhöfer, K. (2018). Language selection errors in switching: language priming or cognitive control? Language, Cognition and Neuroscience, 33(2), 139-147. doi:10.1080/23273798.2017.1363401.

    Abstract

    Although bilingual speakers are very good at selectively using one language rather than another, sometimes language selection errors occur. We examined the relative contribution of top-down cognitive control and bottom-up language priming to these errors. Unbalanced Dutch-English bilinguals named pictures and were cued to switch between languages under time pressure. We also manipulated the number of same-language trials before a switch (long vs. short runs). Results show that speakers made more language selection errors when switching from their second language (L2) to the first language (L1) than vice versa. Furthermore, they made more errors when switching to the L1 after a short compared to a long run of L2 trials. In the reverse switching direction (L1 to L2), run length had no effect. These findings are most compatible with an account of language selection errors that assigns a strong role to top-down processes of cognitive control.

    Additional information

    plcp_a_1363401_sm2537.docx
  • Zheng, X., & Lemhöfer, K. (2019). The “semantic P600” in second language processing: When syntax conflicts with semantics. Neuropsychologia, 127, 131-147. doi:10.1016/j.neuropsychologia.2019.02.010.

    Abstract

    In sentences like “the mouse that chased the cat was hungry”, the syntactically correct interpretation (the mouse chases the cat) is contradicted by semantic and pragmatic knowledge. Previous research has shown that L1 speakers sometimes base sentence interpretation on this type of knowledge (so-called “shallow” or “good-enough” processing). We made use of both behavioural and ERP measurements to investigate whether L2 learners differ from native speakers in the extent to which they engage in “shallow” syntactic processing. German learners of Dutch as well as Dutch native speakers read sentences containing relative clauses (as in the example above) for which the plausible thematic roles were or were not reversed, and made plausibility judgments. The results show that behaviourally, L2 learners had more difficulties than native speakers to discriminate plausible from implausible sentences. In the ERPs, we replicated the previously reported finding of a “semantic P600” for semantic reversal anomalies in native speakers, probably reflecting the effort to resolve the syntax-semantics conflict. In L2 learners, though, this P600 was largely attenuated and surfaced only in those trials that were judged correctly for plausibility. These results generally point at a more prevalent, but not exclusive occurrence of shallow syntactic processing in L2 learners.
  • 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.
  • Zoefel, B., Ten Oever, S., & Sack, A. T. (2018). The involvement of endogenous neural oscillations in the processing of rhythmic input: More than a regular repetition of evoked neural responses. Frontiers in Neuroscience, 12: 95. doi:10.3389/fnins.2018.00095.

    Abstract

    It is undisputed that presenting a rhythmic stimulus leads to a measurable brain response that follows the rhythmic structure of this stimulus. What is still debated, however, is the question whether this brain response exclusively reflects a regular repetition of evoked responses, or whether it also includes entrained oscillatory activity. Here we systematically present evidence in favor of an involvement of entrained neural oscillations in the processing of rhythmic input while critically pointing out which questions still need to be addressed before this evidence could be considered conclusive. In this context, we also explicitly discuss the potential functional role of such entrained oscillations, suggesting that these stimulus-aligned oscillations reflect, and serve as, predictive processes, an idea often only implicitly assumed in the literature.
  • Zora, H., Schwarz, I.-C., & Heldner, M. (2015). Neural correlates of lexical stress: Mismatch negativity reflects fundamental frequency and intensity. NeuroReport, 26(13), 791-796. doi:10.1097/WNR.0000000000000426.

    Abstract

    Neural correlates of lexical stress were studied using the mismatch negativity (MMN) component in event-related potentials. The MMN responses were expected to reveal the encoding of stress information into long-term memory and the contributions of prosodic features such as fundamental frequency (F0) and intensity toward lexical access. In a passive oddball paradigm, neural responses to changes in F0, intensity, and in both features together were recorded for words and pseudowords. The findings showed significant differences not only between words and pseudowords but also between prosodic features. Early processing of prosodic information in words was indexed by an intensity-related MMN and an F0-related P200. These effects were stable at right-anterior and mid-anterior regions. At a later latency, MMN responses were recorded for both words and pseudowords at the mid-anterior and posterior regions. The P200 effect observed for F0 at the early latency for words developed into an MMN response. Intensity elicited smaller MMN for pseudowords than for words. Moreover, a larger brain area was recruited for the processing of words than for the processing of pseudowords. These findings suggest earlier and higher sensitivity to prosodic changes in words than in pseudowords, reflecting a language-related process. The present study, therefore, not only establishes neural correlates of lexical stress but also confirms the presence of long-term memory traces for prosodic information in the brain.
  • Zora, H., Riad, T., & Ylinen, S. (2019). Prosodically controlled derivations in the mental lexicon. Journal of Neurolinguistics, 52: 100856. doi:10.1016/j.jneuroling.2019.100856.

    Abstract

    Swedish morphemes are classified as prosodically specified or prosodically unspecified, depending on lexical or phonological stress, respectively. Here, we investigate the allomorphy of the suffix -(i)sk, which indicates the distinction between lexical and phonological stress; if attached to a lexically stressed morpheme, it takes a non-syllabic form (-sk), whereas if attached to a phonologically stressed morpheme, an epenthetic vowel is inserted (-isk). Using mismatch negativity (MMN), we explored the neural processing of this allomorphy across lexically stressed and phonologically stressed morphemes. In an oddball paradigm, participants were occasionally presented with congruent and incongruent derivations, created by the suffix -(i)sk, within the repetitive presentation of their monomorphemic stems. The results indicated that the congruent derivation of the lexically stressed stem elicited a larger MMN than the incongruent sequences of the same stem and the derivational suffix, whereas after the phonologically stressed stem a non-significant tendency towards an opposite pattern was observed. We argue that the significant MMN response to the congruent derivation in the lexical stress condition is in line with lexical MMN, indicating a holistic processing of the sequence of lexically stressed stem and derivational suffix. The enhanced MMN response to the incongruent derivation in the phonological stress condition, on the other hand, is suggested to reflect combinatorial processing of the sequence of phonologically stressed stem and derivational suffix. These findings bring a new aspect to the dual-system approach to neural processing of morphologically complex words, namely the specification of word stress.
  • Zormpa, E., Meyer, A. S., & Brehm, L. (2019). Slow naming of pictures facilitates memory for their names. Psychonomic Bulletin & Review, 26(5), 1675-1682. doi:10.3758/s13423-019-01620-x.

    Abstract

    Speakers remember their own utterances better than those of their interlocutors, suggesting that language production is beneficial to memory. This may be partly explained by a generation effect: The act of generating a word is known to lead to a memory advantage (Slamecka & Graf, 1978). In earlier work, we showed a generation effect for recognition of images (Zormpa, Brehm, Hoedemaker, & Meyer, 2019). Here, we tested whether the recognition of their names would also benefit from name generation. Testing whether picture naming improves memory for words was our primary aim, as it serves to clarify whether the representations affected by generation are visual or conceptual/lexical. A secondary aim was to assess the influence of processing time on memory. Fifty-one participants named pictures in three conditions: after hearing the picture name (identity condition), backward speech, or an unrelated word. A day later, recognition memory was tested in a yes/no task. Memory in the backward speech and unrelated conditions, which required generation, was superior to memory in the identity condition, which did not require generation. The time taken by participants for naming was a good predictor of memory, such that words that took longer to be retrieved were remembered better. Importantly, that was the case only when generation was required: In the no-generation (identity) condition, processing time was not related to recognition memory performance. This work has shown that generation affects conceptual/lexical representations, making an important contribution to the understanding of the relationship between memory and language.
  • Zormpa, E., Brehm, L., Hoedemaker, R. S., & Meyer, A. S. (2019). The production effect and the generation effect improve memory in picture naming. Memory, 27(3), 340-352. doi:10.1080/09658211.2018.1510966.

    Abstract

    The production effect (better memory for words read aloud than words read silently) and the picture superiority effect (better memory for pictures than words) both improve item memory in a picture naming task (Fawcett, J. M., Quinlan, C. K., & Taylor, T. L. (2012). Interplay of the production and picture superiority effects: A signal detection analysis. Memory (Hove, England), 20(7), 655–666. doi:10.1080/09658211.2012.693510). Because picture naming requires coming up with an appropriate label, the generation effect (better memory for generated than read words) may contribute to the latter effect. In two forced-choice memory experiments, we tested the role of generation in a picture naming task on later recognition memory. In Experiment 1, participants named pictures silently or aloud with the correct name or an unreadable label superimposed. We observed a generation effect, a production effect, and an interaction between the two. In Experiment 2, unreliable labels were included to ensure full picture processing in all conditions. In this experiment, we observed a production and a generation effect but no interaction, implying the effects are dissociable. This research demonstrates the separable roles of generation and production in picture naming and their impact on memory. As such, it informs the link between memory and language production and has implications for memory asymmetries between language production and comprehension.

    Additional information

    pmem_a_1510966_sm9257.pdf
  • De Zubicaray, G., & Fisher, S. E. (Eds.). (2017). Genes, brain and language [Special Issue]. Brain and Language, 172.
  • De Zubicaray, G., & Fisher, S. E. (2017). Genes, Brain, and Language: A brief introduction to the Special Issue. Brain and Language, 172, 1-2. doi:10.1016/j.bandl.2017.08.003.

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