Publications

Displaying 901 - 967 of 967
  • 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, T., & Ravignani, A. (2021). Melodic universals emerge or are sustained through cultural evolution. Frontiers in Psychology, 12: 668300. doi:10.3389/fpsyg.2021.668300.

    Abstract

    To understand why music is structured the way it is, we need an explanation that accounts for both the universality and variability found in musical traditions. Here we test whether statistical universals that have been identified for melodic structures in music can emerge as a result of cultural adaptation to human biases through iterated learning. We use data from an experiment in which artificial whistled systems, where sounds were produced with a slide whistle, were learned by human participants and transmitted multiple times from person to person. These sets of whistled signals needed to be memorized and recalled and the reproductions of one participant were used as the input set for the next. We tested for the emergence of seven different melodic features, such as discrete pitches, motivic patterns, or phrase repetition, and found some evidence for the presence of most of these statistical universals. We interpret this as promising evidence that, similarly to rhythmic universals, iterated learning experiments can also unearth melodic statistical universals. More, ideally cross-cultural, experiments are nonetheless needed. Simulating the cultural transmission of artificial proto-musical systems can help unravel the origins of universal tendencies in musical structures.
  • Verhoef, E., Grove, J., Shapland, C. Y., Demontis, D., Burgess, S., Rai, D., Børglum, A. D., & St Pourcain, B. (2021). Discordant associations of educational attainment with ASD and ADHD implicate a polygenic form of pleiotropy. Nature Communications, 12: 6534. doi:10.1038/s41467-021-26755-1.

    Abstract

    Autism Spectrum Disorder (ASD) and Attention-Deficit/Hyperactivity Disorder (ADHD) are complex co-occurring neurodevelopmental conditions. Their genetic architectures reveal striking similarities but also differences, including strong, discordant polygenic associations with educational attainment (EA). To study genetic mechanisms that present as ASD-related positive and ADHD-related negative genetic correlations with EA, we carry out multivariable regression analyses using genome-wide summary statistics (N = 10,610–766,345). Our results show that EA-related genetic variation is shared across ASD and ADHD architectures, involving identical marker alleles. However, the polygenic association profile with EA, across shared marker alleles, is discordant for ASD versus ADHD risk, indicating independent effects. At the single-variant level, our results suggest either biological pleiotropy or co-localisation of different risk variants, implicating MIR19A/19B microRNA mechanisms. At the polygenic level, they point to a polygenic form of pleiotropy that contributes to the detectable genome-wide correlation between ASD and ADHD and is consistent with effect cancellation across EA-related regions.

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  • Verhoef, E., Shapland, C. Y., Fisher, S. E., Dale, P. S., & St Pourcain, B. (2021). The developmental origins of genetic factors influencing language and literacy: Associations with early-childhood vocabulary. Journal of Child Psychology and Psychiatry, 62(6), 728-738. doi:10.1111/jcpp.13327.

    Abstract

    Background

    The heritability of language and literacy skills increases from early‐childhood to adolescence. The underlying mechanisms are little understood and may involve (a) the amplification of genetic influences contributing to early language abilities, and/or (b) the emergence of novel genetic factors (innovation). Here, we investigate the developmental origins of genetic factors influencing mid‐childhood/early‐adolescent language and literacy. We evaluate evidence for the amplification of early‐childhood genetic factors for vocabulary, in addition to genetic innovation processes.
    Methods

    Expressive and receptive vocabulary scores at 38 months, thirteen language‐ and literacy‐related abilities and nonverbal cognition (7–13 years) were assessed in unrelated children from the Avon Longitudinal Study of Parents and Children (ALSPAC, Nindividuals ≤ 6,092). We investigated the multivariate genetic architecture underlying early‐childhood expressive and receptive vocabulary, and each of 14 mid‐childhood/early‐adolescent language, literacy or cognitive skills with trivariate structural equation (Cholesky) models as captured by genome‐wide genetic relationship matrices. The individual path coefficients of the resulting structural models were finally meta‐analysed to evaluate evidence for overarching patterns.
    Results

    We observed little support for the emergence of novel genetic sources for language, literacy or cognitive abilities during mid‐childhood or early adolescence. Instead, genetic factors of early‐childhood vocabulary, especially those unique to receptive skills, were amplified and represented the majority of genetic variance underlying many of these later complex skills (≤99%). The most predictive early genetic factor accounted for 29.4%(SE = 12.9%) to 45.1%(SE = 7.6%) of the phenotypic variation in verbal intelligence and literacy skills, but also for 25.7%(SE = 6.4%) in performance intelligence, while explaining only a fraction of the phenotypic variation in receptive vocabulary (3.9%(SE = 1.8%)).
    Conclusions

    Genetic factors contributing to many complex skills during mid‐childhood and early adolescence, including literacy, verbal cognition and nonverbal cognition, originate developmentally in early‐childhood and are captured by receptive vocabulary. This suggests developmental genetic stability and overarching aetiological mechanisms.

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  • 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
  • Verhoef, E., Shapland, C. Y., Fisher, S. E., Dale, P. S., & St Pourcain, B. (2021). The developmental genetic architecture of vocabulary skills during the first three years of life: Capturing emerging associations with later-life reading and cognition. PLoS Genetics, 17(2): e1009144. doi:10.1371/journal.pgen.1009144.

    Abstract

    Individual differences in early-life vocabulary measures are heritable and associated with subsequent reading and cognitive abilities, although the underlying mechanisms are little understood. Here, we (i) investigate the developmental genetic architecture of expressive and receptive vocabulary in early-life and (ii) assess timing of emerging genetic associations with mid-childhood verbal and non-verbal skills. We studied longitudinally assessed early-life vocabulary measures (15–38 months) and later-life verbal and non-verbal skills (7–8 years) in up to 6,524 unrelated children from the population-based Avon Longitudinal Study of Parents and Children (ALSPAC) cohort. We dissected the phenotypic variance of rank-transformed scores into genetic and residual components by fitting multivariate structural equation models to genome-wide genetic-relationship matrices. Our findings show that the genetic architecture of early-life vocabulary involves multiple distinct genetic factors. Two of these genetic factors are developmentally stable and also contribute to genetic variation in mid-childhood skills: One genetic factor emerging with expressive vocabulary at 24 months (path coefficient: 0.32(SE = 0.06)) was also related to later-life reading (path coefficient: 0.25(SE = 0.12)) and verbal intelligence (path coefficient: 0.42(SE = 0.13)), explaining up to 17.9% of the phenotypic variation. A second, independent genetic factor emerging with receptive vocabulary at 38 months (path coefficient: 0.15(SE = 0.07)), was more generally linked to verbal and non-verbal cognitive abilities in mid-childhood (reading path coefficient: 0.57(SE = 0.07); verbal intelligence path coefficient: 0.60(0.10); performance intelligence path coefficient: 0.50(SE = 0.08)), accounting for up to 36.1% of the phenotypic variation and the majority of genetic variance in these later-life traits (≥66.4%). Thus, the genetic foundations of mid-childhood reading and cognitive abilities are diverse. They involve at least two independent genetic factors that emerge at different developmental stages during early language development and may implicate differences in cognitive processes that are already detectable during toddlerhood.

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  • Verhoef, E., Allegrini, A. G., Jansen, P. R., Lange, K., Wang, C. A., Morgan, A. T., Ahluwalia, T. S., Symeonides, C., EAGLE-Working Group, Eising, E., Franken, M.-C., Hypponen, E., Mansell, T., Olislagers, M., Omerovic, E., Rimfeld, K., Schlag, F., Selzam, S., Shapland, C. Y., Tiemeier, H., Whitehouse, A. J. O. Verhoef, E., Allegrini, A. G., Jansen, P. R., Lange, K., Wang, C. A., Morgan, A. T., Ahluwalia, T. S., Symeonides, C., EAGLE-Working Group, Eising, E., Franken, M.-C., Hypponen, E., Mansell, T., Olislagers, M., Omerovic, E., Rimfeld, K., Schlag, F., Selzam, S., Shapland, C. Y., Tiemeier, H., Whitehouse, A. J. O., Saffery, R., Bønnelykke, K., Reilly, S., Pennell, C. E., Wake, M., Cecil, C. A., Plomin, R., Fisher, S. E., & St Pourcain, B. (2024). Genome-wide analyses of vocabulary size in infancy and toddlerhood: Associations with Attention-Deficit/Hyperactivity Disorder and cognition-related traits. Biological Psychiatry, 95(1), 859-869. doi:10.1016/j.biopsych.2023.11.025.

    Abstract

    Background

    The number of words children produce (expressive vocabulary) and understand (receptive vocabulary) changes rapidly during early development, partially due to genetic factors. Here, we performed a meta–genome-wide association study of vocabulary acquisition and investigated polygenic overlap with literacy, cognition, developmental phenotypes, and neurodevelopmental conditions, including attention-deficit/hyperactivity disorder (ADHD).

    Methods

    We studied 37,913 parent-reported vocabulary size measures (English, Dutch, Danish) for 17,298 children of European descent. Meta-analyses were performed for early-phase expressive (infancy, 15–18 months), late-phase expressive (toddlerhood, 24–38 months), and late-phase receptive (toddlerhood, 24–38 months) vocabulary. Subsequently, we estimated single nucleotide polymorphism–based heritability (SNP-h2) and genetic correlations (rg) and modeled underlying factor structures with multivariate models.

    Results

    Early-life vocabulary size was modestly heritable (SNP-h2 = 0.08–0.24). Genetic overlap between infant expressive and toddler receptive vocabulary was negligible (rg = 0.07), although each measure was moderately related to toddler expressive vocabulary (rg = 0.69 and rg = 0.67, respectively), suggesting a multifactorial genetic architecture. Both infant and toddler expressive vocabulary were genetically linked to literacy (e.g., spelling: rg = 0.58 and rg = 0.79, respectively), underlining genetic similarity. However, a genetic association of early-life vocabulary with educational attainment and intelligence emerged only during toddlerhood (e.g., receptive vocabulary and intelligence: rg = 0.36). Increased ADHD risk was genetically associated with larger infant expressive vocabulary (rg = 0.23). Multivariate genetic models in the ALSPAC (Avon Longitudinal Study of Parents and Children) cohort confirmed this finding for ADHD symptoms (e.g., at age 13; rg = 0.54) but showed that the association effect reversed for toddler receptive vocabulary (rg = −0.74), highlighting developmental heterogeneity.

    Conclusions

    The genetic architecture of early-life vocabulary changes during development, shaping polygenic association patterns with later-life ADHD, literacy, and cognition-related traits.
  • Vernes, S. C., Kriengwatana, B. P., Beeck, V. C., Fischer, J., Tyack, P. L., Ten Cate, C., & Janik, V. M. (2021). The multi-dimensional nature of vocal learning. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376: 20200236. doi:10.1098/rstb.2020.0236.

    Abstract

    How learning affects vocalizations is a key question in the study of animal
    communication and human language. Parallel efforts in birds and humans
    have taught us much about how vocal learning works on a behavioural
    and neurobiological level. Subsequent efforts have revealed a variety of
    cases among mammals in which experience also has a major influence on
    vocal repertoires. Janik and Slater (Anim. Behav. 60, 1–11. (doi:10.1006/
    anbe.2000.1410)) introduced the distinction between vocal usage and pro-
    duction learning, providing a general framework to categorize how
    different types of learning influence vocalizations. This idea was built on
    by Petkov and Jarvis (Front. Evol. Neurosci. 4, 12. (doi:10.3389/fnevo.2012.
    00012)) to emphasize a more continuous distribution between limited and
    more complex vocal production learners. Yet, with more studies providing
    empirical data, the limits of the initial frameworks become apparent.
    We build on these frameworks to refine the categorization of vocal learning
    in light of advances made since their publication and widespread agreement
    that vocal learning is not a binary trait. We propose a novel classification
    system, based on the definitions by Janik and Slater, that deconstructs
    vocal learning into key dimensions to aid in understanding the mechanisms
    involved in this complex behaviour. We consider how vocalizations can
    change without learning, and a usage learning framework that considers
    context specificity and timing. We identify dimensions of vocal production
    learning, including the copying of auditory models (convergence/
    divergence on model sounds, accuracy of copying), the degree of change
    (type and breadth of learning) and timing (when learning takes place, the
    length of time it takes and how long it is retained). We consider grey
    areas of classification and current mechanistic understanding of these beha-
    viours. Our framework identifies research needs and will help to inform
    neurobiological and evolutionary studies endeavouring to uncover the
    multi-dimensional nature of vocal learning.
    This article is part of the theme issue ‘Vocal learning in animals and
    humans’.
  • Vernes, S. C., Janik, V. M., Fitch, W. T., & Slater, P. J. B. (2021). Vocal learning in animals and humans. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376: 20200234. doi:10.1098/rstb.2020.0234.
  • 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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    Supplementary file
  • 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.

    Additional information

    Supplementary material
  • 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.
  • Von Holzen, K., & Bergmann, C. (2021). The development of infants’ responses to mispronunciations: A meta-analysis. Developmental Psychology, 57(1), 1-18. doi:10.1037/dev0001141.

    Abstract

    As they develop into mature speakers of their native language, infants must not only learn words but also the sounds that make up those words. To do so, they must strike a balance between accepting speaker dependent variation (e.g. mood, voice, accent), but appropriately rejecting variation when it (potentially) changes a word's meaning (e.g. cat vs. hat). This meta-analysis focuses on studies investigating infants' ability to detect mispronunciations in familiar words, or mispronunciation sensitivity. Our goal was to evaluate the development of infants' phonological representations for familiar words as well as explore the role of experimental manipulations related to theoretical questions and analysis choices. The results show that although infants are sensitive to mispronunciations, they still accept these altered forms as labels for target objects. Interestingly, this ability is not modulated by age or vocabulary size, suggesting that a mature understanding of native language phonology may be present in infants from an early age, possibly before the vocabulary explosion. These results also support several theoretical assumptions made in the literature, such as sensitivity to mispronunciation size and position of the mispronunciation. We also shed light on the impact of data analysis choices that may lead to different conclusions regarding the development of infants' mispronunciation sensitivity. Our paper concludes with recommendations for improved practice in testing infants' word and sentence processing on-line.
  • 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 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.

    Additional information

    Data available (link to Figshare)
  • Wagner, M. A., Broersma, M., McQueen, J. M., Dhaene, S., & Lemhöfer, K. (2021). Phonetic convergence to non-native speech: Acoustic and perceptual evidence. Journal of Phonetics, 88: 101076. doi:10.1016/j.wocn.2021.101076.

    Abstract

    While the tendency of speakers to align their speech to that of others acoustic-phonetically has been widely studied among native speakers, very few studies have examined whether natives phonetically converge to non-native speakers. Here we measured native Dutch speakers’ convergence to a non-native speaker with an unfamiliar accent in a novel non-interactive task. Furthermore, we assessed the role of participants’ perceptions of the non-native accent in their tendency to converge. In addition to a perceptual measure (AXB ratings), we examined convergence on different acoustic dimensions (e.g., vowel spectra, fricative CoG, speech rate, overall f0) to determine what dimensions, if any, speakers converge to. We further combined these two types of measures to discover what dimensions weighed in raters’ judgments of convergence. The results reveal overall convergence to our non-native speaker, as indexed by both perceptual and acoustic measures. However, the ratings suggest the stronger participants rated the non-native accent to be, the less likely they were to converge. Our findings add to the growing body of evidence that natives can phonetically converge to non-native speech, even without any apparent socio-communicative motivation to do so. We argue that our results are hard to integrate with a purely social view of convergence.
  • 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.
  • Wang, M.-Y., Korbmacher, M., Eikeland, R., Craven, A. R., & Specht, K. (2024). The intra‐individual reliability of1H‐MRSmeasurement in the anterior cingulate cortex across 1 year. Human Brain Mapping, 45(1): e26531. doi:10.1002/hbm.26531.

    Abstract

    Magnetic resonance spectroscopy (MRS) is the primary method that can measure the levels of metabolites in the brain in vivo. To achieve its potential in clinical usage, the reliability of the measurement requires further articulation. Although there are many studies that investigate the reliability of gamma-aminobutyric acid (GABA), comparatively few studies have investigated the reliability of other brain metabolites, such as glutamate (Glu), N-acetyl-aspartate (NAA), creatine (Cr), phosphocreatine (PCr), or myo-inositol (mI), which all play a significant role in brain development and functions. In addition, previous studies which predominately used only two measurements (two data points) failed to provide the details of the time effect (e.g., time-of-day) on MRS measurement within subjects. Therefore, in this study, MRS data located in the anterior cingulate cortex (ACC) were repeatedly recorded across 1 year leading to at least 25 sessions for each subject with the aim of exploring the variability of other metabolites by using the index coefficient of variability (CV); the smaller the CV, the more reliable the measurements. We found that the metabolites of NAA, tNAA, and tCr showed the smallest CVs (between 1.43% and 4.90%), and the metabolites of Glu, Glx, mI, and tCho showed modest CVs (between 4.26% and 7.89%). Furthermore, we found that the concentration reference of the ratio to water results in smaller CVs compared to the ratio to tCr. In addition, we did not find any time-of-day effect on the MRS measurements. Collectively, the results of this study indicate that the MRS measurement is reasonably reliable in quantifying the levels of metabolites.

    Additional information

    tables and figures data
  • Wang, X., Jahagirdar, S., Bakker, W., Lute, C., Kemp, B., Knegsel, A. v., & Saccenti, E. (2024). Discrimination of Lipogenic or Glucogenic Diet Effects in Early-Lactation Dairy Cows Using Plasma Metabolite Abundances and Ratios in Combination with Machine Learning. Metabolites, 14(4): 230. doi:10.3390/metabo14040230.

    Abstract

    During early lactation, dairy cows have a negative energy balance since their energy demands exceed their energy intake: in this study, we aimed to investigate the association between diet and plasma metabolomics profiles and how these relate to energy unbalance of course in the early-lactation stage. Holstein-Friesian cows were randomly assigned to a glucogenic (n = 15) or lipogenic (n = 15) diet in early lactation. Blood was collected in week 2 and week 4 after calving. Plasma metabolite profiles were detected using liquid chromatography–mass spectrometry (LC-MS), and a total of 39 metabolites were identified. Two plasma metabolomic profiles were available every week for each cow. Metabolite abundance and metabolite ratios were used for the analysis using the XGboost algorithm to discriminate between diet treatment and lactation week. Using metabolite ratios resulted in better discrimination performance compared with the metabolite abundances in assigning cows to a lipogenic diet or a glucogenic diet. The quality of the discrimination of performance of lipogenic diet and glucogenic diet effects improved from 0.606 to 0.753 and from 0.696 to 0.842 in week 2 and week 4 (as measured by area under the curve, AUC), when the metabolite abundance ratios were used instead of abundances. The top discriminating ratios for diet were the ratio of arginine to tyrosine and the ratio of aspartic acid to valine in week 2 and week 4, respectively. For cows fed the lipogenic diet, choline and the ratio of creatinine to tryptophan were top features to discriminate cows in week 2 vs. week 4. For cows fed the glucogenic diet, methionine and the ratio of 4-hydroxyproline to choline were top features to discriminate dietary effects in week 2 or week 4. This study shows the added value of using metabolite abundance ratios to discriminate between lipogenic and glucogenic diet and lactation weeks in early-lactation cows when using metabolomics data. The application of this research will help to accurately regulate the nutrition of lactating dairy cows and promote sustainable agricultural development.
  • 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
  • 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.
  • 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
  • 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.

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    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.
  • Wesseldijk, L. W., Henechowicz, T. L., Baker, D. J., Bignardi, G., Karlsson, R., Gordon, R. L., Mosing, M. A., Ullén, F., & Fisher, S. E. (2024). Notes from Beethoven’s genome. Current Biology, 34(6), R233-R234. doi:10.1016/j.cub.2024.01.025.

    Abstract

    Rapid advances over the last decade in DNA sequencing and statistical genetics enable us to investigate the genomic makeup of individuals throughout history. In a recent notable study, Begg et al.1 used Ludwig van Beethoven’s hair strands for genome sequencing and explored genetic predispositions for some of his documented medical issues. Given that it was arguably Beethoven’s skills as a musician and composer that made him an iconic figure in Western culture, we here extend the approach and apply it to musicality. We use this as an example to illustrate the broader challenges of individual-level genetic predictions.

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  • Wilkinson, G. S., Adams, D. M., Haghani, A., Lu, A. T., Zoller, J., Breeze, C. E., Arnold, B. D., Ball, H. C., Carter, G. G., Cooper, L. N., Dechmann, D. K. N., Devanna, P., Fasel, N. J., Galazyuk, A. V., Günther, L., Hurme, E., Jones, G., Knörnschild, M., Lattenkamp, E. Z., Li, C. Z. and 17 moreWilkinson, G. S., Adams, D. M., Haghani, A., Lu, A. T., Zoller, J., Breeze, C. E., Arnold, B. D., Ball, H. C., Carter, G. G., Cooper, L. N., Dechmann, D. K. N., Devanna, P., Fasel, N. J., Galazyuk, A. V., Günther, L., Hurme, E., Jones, G., Knörnschild, M., Lattenkamp, E. Z., Li, C. Z., Mayer, F., Reinhardt, J. A., Medellin, R. A., Nagy, M., Pope, B., Power, M. L., Ransome, R. D., Teeling, E. C., Vernes, S. C., Zamora-Mejías, D., Zhang, J., Faure, P. A., Greville, L. J., Herrera M., L. G., Flores-Martínez, J. J., & Horvath, S. (2021). DNA methylation predicts age and provides insight into exceptional longevity of bats. Nature Communications, 12: 1615. doi:10.1038/s41467-021-21900-2.

    Abstract

    Exceptionally long-lived species, including many bats, rarely show overt signs of aging, making it difficult to determine why species differ in lifespan. Here, we use DNA methylation (DNAm) profiles from 712 known-age bats, representing 26 species, to identify epigenetic changes associated with age and longevity. We demonstrate that DNAm accurately predicts chronological age. Across species, longevity is negatively associated with the rate of DNAm change at age-associated sites. Furthermore, analysis of several bat genomes reveals that hypermethylated age- and longevity-associated sites are disproportionately located in promoter regions of key transcription factors (TF) and enriched for histone and chromatin features associated with transcriptional regulation. Predicted TF binding site motifs and enrichment analyses indicate that age-related methylation change is influenced by developmental processes, while longevity-related DNAm change is associated with innate immunity or tumorigenesis genes, suggesting that bat longevity results from augmented immune response and cancer suppression.

    Additional information

    supplementary information
  • Willems, R. M., & Peelen, M. V. (2021). How context changes the neural basis of perception and language. iScience, 24(5): 102392. doi:10.1016/j.isci.2021.102392.

    Abstract

    Cognitive processes—from basic sensory analysis to language understanding—are typically contextualized. While the importance of considering context for understanding cognition has long been recognized in psychology and philosophy, it has not yet had much impact on cognitive neuroscience research, where cognition is often studied in decontextualized paradigms. Here, we present examples of recent studies showing that context changes the neural basis of diverse cognitive processes, including perception, attention, memory, and language. Within the domains of perception and language, we review neuroimaging results showing that context interacts with stimulus processing, changes activity in classical perception and language regions, and recruits additional brain regions that contribute crucially to naturalistic perception and language. We discuss how contextualized cognitive neuroscience will allow for discovering new principles of the mind and brain.
  • 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.
  • Winter, B., Lupyan, G., Perry, L. K., Dingemanse, M., & Perlman, M. (2024). Iconicity ratings for 14,000+ English words. Behavior Research Methods, 56, 1640-1655. doi:10.3758/s13428-023-02112-6.

    Abstract

    Iconic words and signs are characterized by a perceived resemblance between aspects of their form and aspects of their meaning. For example, in English, iconic words include peep and crash, which mimic the sounds they denote, and wiggle and zigzag, which mimic motion. As a semiotic property of words and signs, iconicity has been demonstrated to play a role in word learning, language processing, and language evolution. This paper presents the results of a large-scale norming study for more than 14,000 English words conducted with over 1400 American English speakers. We demonstrate the utility of these ratings by replicating a number of existing findings showing that iconicity ratings are related to age of acquisition, sensory modality, semantic neighborhood density, structural markedness, and playfulness. We discuss possible use cases and limitations of the rating dataset, which is made publicly available.
  • 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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  • Woensdregt, M., Cummins, C., & Smith, K. (2021). A computational model of the cultural co-evolution of language and mindreading. Synthese, 199, 1347-1385. doi:10.1007/s11229-020-02798-7.

    Abstract

    Several evolutionary accounts of human social cognition posit that language has co-evolved with the sophisticated mindreading abilities of modern humans. It has also been argued that these mindreading abilities are the product of cultural, rather than biological, evolution. Taken together, these claims suggest that the evolution of language has played an important role in the cultural evolution of human social cognition. Here we present a new computational model which formalises the assumptions that underlie this hypothesis, in order to explore how language and mindreading interact through cultural evolution. This model treats communicative behaviour as an interplay between the context in which communication occurs, an agent’s individual perspective on the world, and the agent’s lexicon. However, each agent’s perspective and lexicon are private mental representations, not directly observable to other agents. Learners are therefore confronted with the task of jointly inferring the lexicon and perspective of their cultural parent, based on their utterances in context. Simulation results show that given these assumptions, an informative lexicon evolves not just under a pressure to be successful at communicating, but also under a pressure for accurate perspective-inference. When such a lexicon evolves, agents become better at inferring others’ perspectives; not because their innate ability to learn about perspectives changes, but because sharing a language (of the right type) with others helps them to do so.
  • Wolf, M. C., Meyer, A. S., Rowland, C. F., & Hintz, F. (2021). The effects of input modality, word difficulty and reading experience on word recognition accuracy. Collabra: Psychology, 7(1): 24919. doi:10.1525/collabra.24919.

    Abstract

    Language users encounter words in at least two different modalities. Arguably, the most frequent encounters are in spoken or written form. Previous research has shown that – compared to the spoken modality – written language features more difficult words. Thus, frequent reading might have effects on word recognition. In the present study, we investigated 1) whether input modality (spoken, written, or bimodal) has an effect on word recognition accuracy, 2) whether this modality effect interacts with word difficulty, 3) whether the interaction of word difficulty and reading experience impacts word recognition accuracy, and 4) whether this interaction is influenced by input modality. To do so, we re-analysed a dataset that was collected in the context of a vocabulary test development to assess in which modality test words should be presented. Participants had carried out a word recognition task, where non-words and words of varying difficulty were presented in auditory, visual and audio-visual modalities. In addition to this main experiment, participants had completed a receptive vocabulary and an author recognition test to measure their reading experience. Our re-analyses did not reveal evidence for an effect of input modality on word recognition accuracy, nor for interactions with word difficulty or language experience. Word difficulty interacted with reading experience in that frequent readers were more accurate in recognizing difficult words than individuals who read less frequently. Practical implications are discussed.
  • 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.
  • Wolna, A., Szewczyk, J., Diaz, M., Domagalik, A., Szwed, M., & Wodniecka, Z. (2024). Domain-general and language-specific contributions to speech production in a second language: An fMRI study using functional localizers. Scientific Reports, 14: 57. doi:10.1038/s41598-023-49375-9.

    Abstract

    For bilinguals, speaking in a second language (L2) compared to the native language (L1) is usually more difficult. In this study we asked whether the difficulty in L2 production reflects increased demands imposed on domain-general or core language mechanisms. We compared the brain response to speech production in L1 and L2 within two functionally-defined networks in the brain: the Multiple Demand (MD) network and the language network. We found that speech production in L2 was linked to a widespread increase of brain activity in the domain-general MD network. The language network did not show a similarly robust differences in processing speech in the two languages, however, we found increased response to L2 production in the language-specific portion of the left inferior frontal gyrus (IFG). To further explore our results, we have looked at domain-general and language-specific response within the brain structures postulated to form a Bilingual Language Control (BLC) network. Within this network, we found a robust increase in response to L2 in the domain-general, but also in some language-specific voxels including in the left IFG. Our findings show that L2 production strongly engages domain-general mechanisms, but only affects language sensitive portions of the left IFG. These results put constraints on the current model of bilingual language control by precisely disentangling the domain-general and language-specific contributions to the difficulty in speech production in L2.

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  • Wolna, A., Szewczyk, J., Diaz, M., Domagalik, A., Szwed, M., & Wodniecka, Z. (2024). Tracking components of bilingual language control in speech production: An fMRI study using functional localizers. Neurobiology of Language, 5(2), 315-340. doi:10.1162/nol_a_00128.

    Abstract

    When bilingual speakers switch back to speaking in their native language (L1) after having used their second language (L2), they often experience difficulty in retrieving words in their L1. This phenomenon is referred to as the L2 after-effect. We used the L2 after-effect as a lens to explore the neural bases of bilingual language control mechanisms. Our goal was twofold: first, to explore whether bilingual language control draws on domain-general or language-specific mechanisms; second, to investigate the precise mechanism(s) that drive the L2 after-effect. We used a precision fMRI approach based on functional localizers to measure the extent to which the brain activity that reflects the L2 after-effect overlaps with the language network (Fedorenko et al., 2010) and the domain-general multiple demand network (Duncan, 2010), as well as three task-specific networks that tap into interference resolution, lexical retrieval, and articulation. Forty-two Polish–English bilinguals participated in the study. Our results show that the L2 after-effect reflects increased engagement of domain-general but not language-specific resources. Furthermore, contrary to previously proposed interpretations, we did not find evidence that the effect reflects increased difficulty related to lexical access, articulation, and the resolution of lexical interference. We propose that difficulty of speech production in the picture naming paradigm—manifested as the L2 after-effect—reflects interference at a nonlinguistic level of task schemas or a general increase of cognitive control engagement during speech production in L1 after L2.

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    supplementary materials
  • 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.

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  • Wongratwanich, P., Shimabukuro, K., Konishi, M., Nagasaki, T., Ohtsuka, M., Suei, Y., Nakamoto, T., Verdonschot, R. G., Kanesaki, T., Sutthiprapaporn, P., & Kakimoto, N. (2021). Do various imaging modalities provide potential early detection and diagnosis of medication-related osteonecrosis of the jaw? A review. Dentomaxillofacial Radiology, 50: 20200417. doi:10.1259/dmfr.20200417.

    Abstract


    Objective: Patients with medication-related osteonecrosis of the jaw (MRONJ) often visit their dentists at advanced stages and subsequently require treatments that greatly affect quality of life. Currently, no clear diagnostic criteria exist to assess MRONJ, and the definitive diagnosis solely relies on clinical bone exposure. This ambiguity leads to a diagnostic delay, complications, and unnecessary burden. This article aims to identify imaging modalities' usage and findings of MRONJ to provide possible approaches for early detection.

    Methods: Literature searches were conducted using PubMed, Web of Science, Scopus, and Cochrane Library to review all diagnostic imaging modalities for MRONJ.

    Results: Panoramic radiography offers a fundamental understanding of the lesions. Imaging findings were comparable between non-exposed and exposed MRONJ, showing osteolysis, osteosclerosis, and thickened lamina dura. Mandibular cortex index Class II could be a potential early MRONJ indicator. While three-dimensional modalities, CT and CBCT, were able to show more features unique to MRONJ such as a solid type periosteal reaction, buccal predominance of cortical perforation, and bone-within-bone appearance. MRI signal intensities of vital bones are hypointense on T1WI and hyperintense on T2WI and STIR when necrotic bone shows hypointensity on all T1WI, T2WI, and STIR. Functional imaging is the most sensitive method but is usually performed in metastasis detection rather than being a diagnostic tool for early MRONJ.

    Conclusion: Currently, MRONJ-specific imaging features cannot be firmly established. However, the current data are valuable as it may lead to a more efficient diagnostic procedure along with a more suitable selection of imaging modalities.
  • 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., & Lupker, S. J. (2021). Orthographic properties of distractors do influence phonological Stroop effects: Evidence from Japanese Romaji distractors. Memory & Cognition, 49(3), 600-612. doi:10.3758/s13421-020-01103-8.

    Abstract

    In attempting to understand mental processes, it is important to use a task that appropriately reflects the underlying processes being investigated. Recently, Verdonschot and Kinoshita (Memory & Cognition, 46,410-425, 2018) proposed that a variant of the Stroop task-the "phonological Stroop task"-might be a suitable tool for investigating speech production. The major advantage of this task is that the task is apparently not affected by the orthographic properties of the stimuli, unlike other, commonly used, tasks (e.g., associative-cuing and word-reading tasks). The viability of this proposal was examined in the present experiments by manipulating the script types of Japanese distractors. For Romaji distractors (e.g., "kushi"), color-naming responses were faster when the initial phoneme was shared between the color name and the distractor than when the initial phonemes were different, thereby showing a phoneme-based phonological Stroop effect (Experiment1). In contrast, no such effect was observed when the same distractors were presented in Katakana (e.g., "< ") pound, replicating Verdonschot and Kinoshita's original results (Experiment2). A phoneme-based effect was again found when the Katakana distractors used in Verdonschot and Kinoshita's original study were transcribed and presented in Romaji (Experiment3). Because the observation of a phonemic effectdirectly depended on the orthographic properties of the distractor stimuli, we conclude that the phonological Stroop task is also susceptible to orthographic influences.
  • Zaadnoordijk, L., Buckler, H., Cusack, R., Tsuji, S., & Bergmann, C. (2021). A global perspective on testing infants online: Introducing ManyBabies-AtHome. Frontiers in Psychology, 12: 703234. doi:10.3389/fpsyg.2021.703234.

    Abstract

    Online testing holds great promise for infant scientists. It could increase participant diversity, improve reproducibility and collaborative possibilities, and reduce costs for researchers and participants. However, despite the rise of platforms and participant databases, little work has been done to overcome the challenges of making this approach available to researchers across the world. In this paper, we elaborate on the benefits of online infant testing from a global perspective and identify challenges for the international community that have been outside of the scope of previous literature. Furthermore, we introduce ManyBabies-AtHome, an international, multi-lab collaboration that is actively working to facilitate practical and technical aspects of online testing as well as address ethical concerns regarding data storage and protection, and cross-cultural variation. The ultimate goal of this collaboration is to improve the method of testing infants online and make it globally available.
  • Zettersten, M., Cox, C., Bergmann, C., Tsui, A. S. M., Soderstrom, M., Mayor, J., Lundwall, R. A., Lewis, M., Kosie, J. E., Kartushina, N., Fusaroli, R., Frank, M. C., Byers-Heinlein, K., Black, A. K., & Mathur, M. B. (2024). Evidence for infant-directed speech preference is consistent across large-scale, multi-site replication and meta-analysis. Open Mind, 8, 439-461. doi:10.1162/opmi_a_00134.

    Abstract

    There is substantial evidence that infants prefer infant-directed speech (IDS) to adult-directed speech (ADS). The strongest evidence for this claim has come from two large-scale investigations: i) a community-augmented meta-analysis of published behavioral studies and ii) a large-scale multi-lab replication study. In this paper, we aim to improve our understanding of the IDS preference and its boundary conditions by combining and comparing these two data sources across key population and design characteristics of the underlying studies. Our analyses reveal that both the meta-analysis and multi-lab replication show moderate effect sizes (d ≈ 0.35 for each estimate) and that both of these effects persist when relevant study-level moderators are added to the models (i.e., experimental methods, infant ages, and native languages). However, while the overall effect size estimates were similar, the two sources diverged in the effects of key moderators: both infant age and experimental method predicted IDS preference in the multi-lab replication study, but showed no effect in the meta-analysis. These results demonstrate that the IDS preference generalizes across a variety of experimental conditions and sampling characteristics, while simultaneously identifying key differences in the empirical picture offered by each source individually and pinpointing areas where substantial uncertainty remains about the influence of theoretically central moderators on IDS preference. Overall, our results show how meta-analyses and multi-lab replications can be used in tandem to understand the robustness and generalizability of developmental phenomena.

    Additional information

    supplementary data link to preprint
  • Yu, C., Zhang, Y., Slone, L. K., & Smith, L. B. (2021). The infant’s view redefines the problem of referential uncertainty in early word learning. Proceedings of the National Academy of Sciences of the United States of America, 118(52): e2107019118. doi:10.1073/pnas.2107019118.

    Abstract

    The learning of first object names is deemed a hard problem due to the uncertainty inherent in mapping a heard name to the intended referent in a cluttered and variable world. However, human infants readily solve this problem. Despite considerable theoretical discussion, relatively little is known about the uncertainty infants face in the real world. We used head-mounted eye tracking during parent–infant toy play and quantified the uncertainty by measuring the distribution of infant attention to the potential referents when a parent named both familiar and unfamiliar toy objects. The results show that infant gaze upon hearing an object name is often directed to a single referent which is equally likely to be a wrong competitor or the intended target. This bimodal gaze distribution clarifies and redefines the uncertainty problem and constrains possible solutions.
  • Zhang, Y., Yurovsky, D., & Yu, C. (2021). Cross-situational learning from ambiguous egocentric input is a continuous process: Evidence using the human simulation paradigm. Cognitive Science, 45(7): e13010. doi:10.1111/cogs.13010.

    Abstract

    Recent laboratory experiments have shown that both infant and adult learners can acquire word-referent mappings using cross-situational statistics. The vast majority of the work on this topic has used unfamiliar objects presented on neutral backgrounds as the visual contexts for word learning. However, these laboratory contexts are much different than the real-world contexts in which learning occurs. Thus, the feasibility of generalizing cross-situational learning beyond the laboratory is in question. Adapting the Human Simulation Paradigm, we conducted a series of experiments examining cross-situational learning from children's egocentric videos captured during naturalistic play. Focusing on individually ambiguous naming moments that naturally occur during toy play, we asked how statistical learning unfolds in real time through accumulating cross-situational statistics in naturalistic contexts. We found that even when learning situations were individually ambiguous, learners' performance gradually improved over time. This improvement was driven in part by learners' use of partial knowledge acquired from previous learning situations, even when they had not yet discovered correct word-object mappings. These results suggest that word learning is a continuous process by means of real-time information integration.
  • 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.
  • Zhong, S., Wei, L., Zhao, C., Yang, L., Di, Z., Francks, C., & Gong, G. (2021). Interhemispheric relationship of genetic influence on human brain connectivity. Cerebral Cortex, 31(1), 77-88. doi:10.1093/cercor/bhaa207.

    Abstract

    To understand the origins of interhemispheric differences and commonalities/coupling in human brain wiring, it is crucial to determine how homologous interregional connectivities of the left and right hemispheres are genetically determined and related. To address this, in the present study, we analyzed human twin and pedigree samples with high-quality diffusion magnetic resonance imaging tractography and estimated the heritability and genetic correlation of homologous left and right white matter (WM) connections. The results showed that the heritability of WM connectivity was similar and coupled between the 2 hemispheres and that the degree of overlap in genetic factors underlying homologous WM connectivity (i.e., interhemispheric genetic correlation) varied substantially across the human brain: from complete overlap to complete nonoverlap. Particularly, the heritability was significantly stronger and the chance of interhemispheric complete overlap in genetic factors was higher in subcortical WM connections than in cortical WM connections. In addition, the heritability and interhemispheric genetic correlations were stronger for long-range connections than for short-range connections. These findings highlight the determinants of the genetics underlying WM connectivity and its interhemispheric relationships, and provide insight into genetic basis of WM connectivity asymmetries in both healthy and disease states.

    Additional information

    Supplementary data
  • Zhou, W., Broersma, M., & Cutler, A. (2021). Asymmetric memory for birth language perception versus production in young international adoptees. Cognition, 213: 104788. doi:10.1016/j.cognition.2021.104788.

    Abstract

    Adults who as children were adopted into a different linguistic community retain knowledge of their birth language. The possession (without awareness) of such knowledge is known to facilitate the (re)learning of birth-language speech patterns; this perceptual learning predicts such adults' production success as well, indicating that the retained linguistic knowledge is abstract in nature. Adoptees' acquisition of their adopted language is fast and complete; birth-language mastery disappears rapidly, although this latter process has been little studied. Here, 46 international adoptees from China aged four to 10 years, with Dutch as their new language, plus 47 matched non-adopted Dutch-native controls and 40 matched non-adopted Chinese controls, undertook across a two-week period 10 blocks of training in perceptually identifying Chinese speech contrasts (one segmental, one tonal) which were unlike any Dutch contrasts. Chinese controls easily accomplished all these tasks. The same participants also provided speech production data in an imitation task. In perception, adoptees and Dutch controls scored equivalently poorly at the outset of training; with training, the adoptees significantly improved while the Dutch controls did not. In production, adoptees' imitations both before and after training could be better identified, and received higher goodness ratings, than those of Dutch controls. The perception results confirm that birth-language knowledge is stored and can facilitate re-learning in post-adoption childhood; the production results suggest that although processing of phonological category detail appears to depend on access to the stored knowledge, general articulatory dimensions can at this age also still be remembered, and may facilitate spoken imitation.

    Additional information

    stimulus materials
  • Zhou, H., Van der Ham, S., De Boer, B., Bogaerts, L., & Raviv, L. (2024). Modality and stimulus effects on distributional statistical learning: Sound vs. sight, time vs. space. Journal of Memory and Language, 138: 104531. doi:10.1016/j.jml.2024.104531.

    Abstract

    Statistical learning (SL) is postulated to play an important role in the process of language acquisition as well as in other cognitive functions. It was found to enable learning of various types of statistical patterns across different sensory modalities. However, few studies have distinguished distributional SL (DSL) from sequential and spatial SL, or examined DSL across modalities using comparable tasks. Considering the relevance of such findings to the nature of SL, the current study investigated the modality- and stimulus-specificity of DSL. Using a within-subject design we compared DSL performance in auditory and visual modalities. For each sensory modality, two stimulus types were used: linguistic versus non-linguistic auditory stimuli and temporal versus spatial visual stimuli. In each condition, participants were exposed to stimuli that varied in their length as they were drawn from two categories (short versus long). DSL was assessed using a categorization task and a production task. Results showed that learners’ performance was only correlated for tasks in the same sensory modality. Moreover, participants were better at categorizing the temporal signals in the auditory conditions than in the visual condition, where in turn an advantage of the spatial condition was observed. In the production task participants exaggerated signal length more for linguistic signals than non-linguistic signals. Together, these findings suggest that DSL is modality- and stimulus-sensitive.

    Additional information

    link to preprint
  • 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.
  • Zimianiti, E. (2021). Adjective-noun constructions in Griko: Focusing on measuring adjectives and their placement in the nominal domain. LingUU Journal, 5(2), 62-75.

    Abstract

    This paper examines adjectival placement in Griko, an Italian-Greek lan-
    guage variety. Guardiano and Stavrou (2019, 2014) have argued that
    there is a gap of evidence in the diachrony of adjectives in prenominal
    position and in particular, of measuring adjectives. Evidence is presented
    in this paper contradicting the aforementioned claims. After considering
    the placement of adjectives in Greek and Italian, and their similarities
    and differences, the adjectival pattern of Griko is analysed. The analysis
    is based mostly on written data from the early 20th century proving the
    prenominal position of adjectives and adding to the diachronic schema of
    adjectival placement in Griko.
  • Zinken, J., Kaiser, J., Weidner, M., Mondada, L., Rossi, G., & Sorjonen, M.-L. (2021). Rule talk: Instructing proper play with impersonal deontic statements. Frontiers in Communication, 6: 660394. doi:10.3389/fcomm.2021.660394.

    Abstract

    The present paper explores how rules are enforced and talked about in everyday life. Drawing on a corpus of board game recordings across European languages, we identify a sequential and praxeological context for rule talk. After a game rule is breached, a participant enforces proper play and then formulates a rule with an impersonal deontic statement (e.g. ‘It’s not allowed to do this’). Impersonal deontic statements express what may or may not be done without tying the obligation to a particular individual. Our analysis shows that such statements are used as part of multi-unit and multi-modal turns where rule talk is accomplished through both grammatical and embodied means. Impersonal deontic statements serve multiple interactional goals: they account for having changed another’s behavior in the moment and at the same time impart knowledge for the future. We refer to this complex action as an “instruction”. The results of this study advance our understanding of rules and rule-following in everyday life, and of how resources of language and the body are combined to enforce and formulate rules.
  • Zioga, I., Zhou, Y. J., Weissbart, H., Martin, A. E., & Haegens, S. (2024). Alpha and beta oscillations differentially support word production in a rule-switching task. eNeuro, 11(4): ENEURO.0312-23.2024. doi:10.1523/ENEURO.0312-23.2024.

    Abstract

    Research into the role of brain oscillations in basic perceptual and cognitive functions has suggested that the alpha rhythm reflects functional inhibition while the beta rhythm reflects neural ensemble (re)activation. However, little is known regarding the generalization of these proposed fundamental operations to linguistic processes, such as speech comprehension and production. Here, we recorded magnetoencephalography in participants performing a novel rule-switching paradigm. Specifically, Dutch native speakers had to produce an alternative exemplar from the same category or a feature of a given target word embedded in spoken sentences (e.g., for the word “tuna”, an exemplar from the same category—“seafood”—would be “shrimp”, and a feature would be “pink”). A cue indicated the task rule—exemplar or feature—either before (pre-cue) or after (retro-cue) listening to the sentence. Alpha power during the working memory delay was lower for retro-cue compared with that for pre-cue in the left hemispheric language-related regions. Critically, alpha power negatively correlated with reaction times, suggestive of alpha facilitating task performance by regulating inhibition in regions linked to lexical retrieval. Furthermore, we observed a different spatiotemporal pattern of beta activity for exemplars versus features in the right temporoparietal regions, in line with the proposed role of beta in recruiting neural networks for the encoding of distinct categories. Overall, our study provides evidence for the generalizability of the role of alpha and beta oscillations from perceptual to more “complex, linguistic processes” and offers a novel task to investigate links between rule-switching, working memory, and word production.
  • 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., Riad, T., Ylinen, S., & Csépe, V. (2021). Phonological variations are compensated at the lexical level: Evidence from auditory neural activity. Frontiers in Human Neuroscience, 15: 622904. doi:10.3389/fnhum.2021.622904.

    Abstract

    Dealing with phonological variations is important for speech processing. This article addresses whether phonological variations introduced by assimilatory processes are compensated for at the pre-lexical or lexical level, and whether the nature of variation and the phonological context influence this process. To this end, Swedish nasal regressive place assimilation was investigated using the mismatch negativity (MMN) component. In nasal regressive assimilation, the coronal nasal assimilates to the place of articulation of a following segment, most clearly with a velar or labial place of articulation, as in utan mej “without me” > [ʉːtam mɛjː]. In a passive auditory oddball paradigm, 15 Swedish speakers were presented with Swedish phrases with attested and unattested phonological variations and contexts for nasal assimilation. Attested variations – a coronal-to-labial change as in utan “without” > [ʉːtam] – were contrasted with unattested variations – a labial-to-coronal change as in utom “except” > ∗[ʉːtɔn] – in appropriate and inappropriate contexts created by mej “me” [mɛjː] and dej “you” [dɛjː]. Given that the MMN amplitude depends on the degree of variation between two stimuli, the MMN responses were expected to indicate to what extent the distance between variants was tolerated by the perceptual system. Since the MMN response reflects not only low-level acoustic processing but also higher-level linguistic processes, the results were predicted to indicate whether listeners process assimilation at the pre-lexical and lexical levels. The results indicated no significant interactions across variations, suggesting that variations in phonological forms do not incur any cost in lexical retrieval; hence such variation is compensated for at the lexical level. However, since the MMN response reached significance only for a labial-to-coronal change in a labial context and for a coronal-to-labial change in a coronal context, the compensation might have been influenced by the nature of variation and the phonological context. It is therefore concluded that while assimilation is compensated for at the lexical level, there is also some influence from pre-lexical processing. The present results reveal not only signal-based perception of phonological units, but also higher-level lexical processing, and are thus able to reconcile the bottom-up and top-down models of speech processing.
  • 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.
  • Zora, H., & Csépe, V. (2021). Perception of Prosodic Modulations of Linguistic and Paralinguistic Origin: Evidence From Early Auditory Event-Related Potentials. Frontiers in Neuroscience, 15: 797487. doi:10.3389/fnins.2021.797487.

    Abstract

    How listeners handle prosodic cues of linguistic and paralinguistic origin is a central question for spoken communication. In the present EEG study, we addressed this question by examining neural responses to variations in pitch accent (linguistic) and affective (paralinguistic) prosody in Swedish words, using a passive auditory oddball paradigm. The results indicated that changes in pitch accent and affective prosody elicited mismatch negativity (MMN) responses at around 200 ms, confirming the brain’s pre-attentive response to any prosodic modulation. The MMN amplitude was, however, statistically larger to the deviation in affective prosody in comparison to the deviation in pitch accent and affective prosody combined, which is in line with previous research indicating not only a larger MMN response to affective prosody in comparison to neutral prosody but also a smaller MMN response to multidimensional deviants than unidimensional ones. The results, further, showed a significant P3a response to the affective prosody change in comparison to the pitch accent change at around 300 ms, in accordance with previous findings showing an enhanced positive response to emotional stimuli. The present findings provide evidence for distinct neural processing of different prosodic cues, and statistically confirm the intrinsic perceptual and motivational salience of paralinguistic information in spoken communication.
  • 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

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