Publications

Displaying 601 - 700 of 747
  • Sha, Z., Schijven, D., & Francks, C. (2021). Patterns of brain asymmetry associated with polygenic risks for autism and schizophrenia implicate language and executive functions but not brain masculinization. Molecular Psychiatry, 26(12), 7652-7660. doi:10.1038/s41380-021-01204-z.

    Abstract

    Autism spectrum disorder (ASD) and schizophrenia have been conceived as partly opposing disorders in terms of systemizing versus empathizing cognitive styles, with resemblances to male versus female average sex differences. Left-right asymmetry of the brain is an important aspect of its organization that shows average differences between the sexes, and can be altered in both ASD and schizophrenia. Here we mapped multivariate associations of polygenic risk scores for ASD and schizophrenia with asymmetries of regional cerebral cortical surface area, thickness and subcortical volume measures in 32,256 participants from the UK Biobank. Polygenic risks for the two disorders were positively correlated (r=0.08, p=7.13×10-50), and both were higher in females compared to males, consistent with biased participation against higher-risk males. Each polygenic risk score was associated with multivariate brain asymmetry after adjusting for sex, ASD r=0.03, p=2.17×10-9, schizophrenia r=0.04, p=2.61×10-11, but the multivariate patterns were mostly distinct for the two polygenic risks, and neither resembled average sex differences. Annotation based on meta-analyzed functional imaging data showed that both polygenic risks were associated with asymmetries of regions important for language and executive functions, consistent with behavioural associations that arose in phenome-wide association analysis. Overall, the results indicate that distinct patterns of subtly altered brain asymmetry may be functionally relevant manifestations of polygenic risks for ASD and schizophrenia, but do not support brain masculinization or feminization in their etiologies.
  • Sha, Z., Pepe, A., Schijven, D., Carrion Castillo, A., Roe, J. M., Westerhausen, R., Joliot, M., Fisher, S. E., Crivello, F., & Francks, C. (2021). Handedness and its genetic influences are associated with structural asymmetries of the cerebral cortex in 31,864 individuals. Proceedings of the National Academy of Sciences of the United States of America, 118(47): e2113095118. doi:10.1073/pnas.2113095118.

    Abstract

    Roughly 10% of the human population is left-handed, and this rate is increased in some brain-related disorders. The neuroanatomical correlates of hand preference have remained equivocal. We resampled structural brain image data from 28,802 right-handers and 3,062 left-handers (UK Biobank population dataset) to a symmetrical surface template, and mapped asymmetries for each of 8,681 vertices across the cerebral cortex in each individual. Left-handers compared to right-handers showed average differences of surface area asymmetry within the fusiform cortex, the anterior insula, the anterior middle cingulate cortex, and the precentral cortex. Meta-analyzed functional imaging data implicated these regions in executive functions and language. Polygenic disposition to left-handedness was associated with two of these regional asymmetries, and 18 loci previously linked with left-handedness by genome-wide screening showed associations with one or more of these asymmetries. Implicated genes included six encoding microtubule-related proteins: TUBB, TUBA1B, TUBB3, TUBB4A, MAP2, and NME7—mutations in the latter can cause left to right reversal of the visceral organs. There were also two cortical regions where average thickness asymmetry was altered in left-handedness: on the postcentral gyrus and the inferior occipital cortex, functionally annotated with hand sensorimotor and visual roles. These cortical thickness asymmetries were not heritable. Heritable surface area asymmetries of language-related regions may link the etiologies of hand preference and language, whereas nonheritable asymmetries of sensorimotor cortex may manifest as consequences of hand preference.
  • Sha, Z., Schijven, D., Carrion Castillo, A., Joliot, M., Mazoyer, B., Fisher, S. E., Crivello, F., & Francks, C. (2021). The genetic architecture of structural left–right asymmetry of the human brain. Nature Human Behaviour, 5, 1226-1236. doi:10.1038/s41562-021-01069-w.

    Abstract

    Left–right hemispheric asymmetry is an important aspect of healthy brain organization for many functions including language, and it can be altered in cognitive and psychiatric disorders. No mechanism has yet been identified for establishing the human brain’s left–right axis. We performed multivariate genome-wide association scanning of cortical regional surface area and thickness asymmetries, and subcortical volume asymmetries, using data from 32,256 participants from the UK Biobank. There were 21 significant loci associated with different aspects of brain asymmetry, with functional enrichment involving microtubule-related genes and embryonic brain expression. These findings are consistent with a known role of the cytoskeleton in left–right axis determination in other organs of invertebrates and frogs. Genetic variants associated with brain asymmetry overlapped with those associated with autism, educational attainment and schizophrenia. Comparably large datasets will likely be required in future studies, to replicate and further clarify the associations of microtubule-related genes with variation in brain asymmetry, behavioural and psychiatric traits.
  • Shapland, C. Y., Verhoef, E., Smith, G. D., Fisher, S. E., Verhulst, B., Dale, P. S., & St Pourcain, B. (2021). Multivariate genome-wide covariance analyses of literacy, language and working memory skills reveal distinct etiologies. npj Science of Learning, 6: 23. doi:10.1038/s41539-021-00101-y.

    Abstract

    Several abilities outside literacy proper are associated with reading and spelling, both phenotypically and genetically, though our knowledge of multivariate genomic covariance structures is incomplete. Here, we introduce structural models describing genetic and residual influences between traits to study multivariate links across measures of literacy, phonological awareness, oral language, and phonological working memory (PWM) in unrelated UK youth (8-13 years, N=6,453). We find that all phenotypes share a large proportion of underlying genetic variation, although especially oral language and PWM reveal substantial differences in their genetic variance composition with substantial trait-specific genetic influences. Multivariate genetic and residual trait covariance showed concordant patterns, except for marked differences between oral language and literacy/phonological awareness, where strong genetic links contrasted near-zero residual overlap. These findings suggest differences in etiological mechanisms, acting beyond a pleiotropic set of genetic variants, and implicate variation in trait modifiability even among phenotypes that have high genetic correlations.

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    supplementary information
  • Shkaravska, O., & Van Eekelen, M. (2021). Polynomial solutions of algebraic difference equations and homogeneous symmetric polynomials. Journal of Symbolic Computation, 103, 22-45. doi:10.1016/j.jsc.2019.10.022.

    Abstract

    This article addresses the problem of computing an upper bound of
    the degree d of a polynomial solution P(x) of an algebraic differ-
    ence equation of the form Gx)(P(x −τ1), . . . , P(x −τs) + G0(x) =

    0 when such P(x) with the coefficients in a field K of character-
    istic zero exists and where G is a non-linear s-variable polynomial
    with coefficients in K[x] and G0 is a polynomial with coefficients
    in K.
    It will be shown that if G is a quadratic polynomial with constant
    coefficients then one can construct a countable family of polynomi-
    als fl(u0) such that if there exists a (minimal) index l0 with fl0(u0)
    being a non-zero polynomial, then the degree d is one of its roots
    or d ≤ l0, or d < deg(G0). Moreover, the existence of such l0 will
    be proven for K being the field of real numbers. These results are
    based on the properties of the modules generated by special fami-
    lies of homogeneous symmetric polynomials.
    A sufficient condition for the existence of a similar bound of the
    degree of a polynomial solution for an algebraic difference equation
    with G of arbitrary total degree and with variable coefficients will
    be proven as well.
  • Shopen, T., Reid, N., Shopen, G., & Wilkins, D. G. (1997). Ensuring the survival of Aboriginal and Torres Strait islander languages into the 21st century. Australian Review of Applied Linguistics, 10(1), 143-157.

    Abstract

    Aboriginal languages threatened by speakers poor economic and social conditions; some may survive through support for community development, language maintenance, bilingual education and training of Aboriginal teachers and linguists, and nonAboriginal teachers of Aboriginal and Islander students.
  • Simon-Thomas, E. R., Keltner, D. J., Sauter, D., Sinicropi-Yao, L., & Abramson, A. (2009). The voice conveys specific emotions: Evidence from vocal burst displays. Emotion, 9, 838-846. doi:10.1037/a0017810.

    Abstract

    Studies of emotion signaling inform claims about the taxonomic structure, evolutionary origins, and physiological correlates of emotions. Emotion vocalization research has tended to focus on a limited set of emotions: anger, disgust, fear, sadness, surprise, happiness, and for the voice, also tenderness. Here, we examine how well brief vocal bursts can communicate 22 different emotions: 9 negative (Study 1) and 13 positive (Study 2), and whether prototypical vocal bursts convey emotions more reliably than heterogeneous vocal bursts (Study 3). Results show that vocal bursts communicate emotions like anger, fear, and sadness, as well as seldom-studied states like awe, compassion, interest, and embarrassment. Ancillary analyses reveal family-wise patterns of vocal burst expression. Errors in classification were more common within emotion families (e.g., ‘self-conscious,’ ‘pro-social’) than between emotion families. The three studies reported highlight the voice as a rich modality for emotion display that can inform fundamental constructs about emotion.
  • Slim, M. S., Lauwers, P., & Hartsuiker, R. J. (2021). Monolingual and bilingual logical representations of quantificational scope: Evidence from priming in language comprehension. Journal of Memory and Language, 116: 104184. doi:10.1016/j.jml.2020.104184.

    Abstract

    A doubly quantified sentence, such as All hikers climbed a hill, allows two interpretations: Did all hikers climb different hills, or did they climb the same hill? Previous work has shown that comprehenders construct disambiguated logical representations of these interpretations (Raffray & Pickering, 2010). We extended this line of research by investigating whether bilingual logical representations are shared between languages or separate per language. We conducted four sentence-picture matching experiments in which we primed interpretations of doubly quantified sentences in Dutch and French monolingual and bilingual language comprehension. These experiments showed that bilinguals have fully shared logical representations and that logical representations constructed in the L1 and the L2 are comparable. Moreover, a control experiment ruled out that the priming effects were driven by visual overlap between prime and target pictures. We discuss these findings in terms of a language-dependent account of logical representations, although these findings can also be reconciled with the idea that logical representations involve conceptual mental models of sentence meaning.
  • Slivac, K., Hervais-Adelman, A., Hagoort, P., & Flecken, M. (2021). Linguistic labels cue biological motion perception and misperception. Scientific Reports, 11: 17239. doi:10.1038/s41598-021-96649-1.

    Abstract

    Linguistic labels exert a particularly strong top-down influence on perception. The potency of this influence has been ascribed to their ability to evoke category-diagnostic features of concepts. In doing this, they facilitate the formation of a perceptual template concordant with those features, effectively biasing perceptual activation towards the labelled category. In this study, we employ a cueing paradigm with moving, point-light stimuli across three experiments, in order to examine how the number of biological motion features (form and kinematics) encoded in lexical cues modulates the efficacy of lexical top-down influence on perception. We find that the magnitude of lexical influence on biological motion perception rises as a function of the number of biological motion-relevant features carried by both cue and target. When lexical cues encode multiple biological motion features, this influence is robust enough to mislead participants into reporting erroneous percepts, even when a masking level yielding high performance is used.
  • Slobin, D. I., & Bowerman, M. (2007). Interfaces between linguistic typology and child language research. Linguistic Typology, 11(1), 213-226. doi:10.1515/LINGTY.2007.015.
  • Slonimska, A., Ozyurek, A., & Capirci, O. (2021). Using depiction for efficient communication in LIS (Italian Sign Language). Language and Cognition, 13(3), 367 -396. doi:10.1017/langcog.2021.7.

    Abstract

    Meanings communicated with depictions constitute an integral part of how speakers and signers actually use language (Clark, 2016). Recent studies have argued that, in sign languages, depicting strategy like constructed action (CA), in which a signer enacts the referent, is used for referential purposes in narratives. Here, we tested the referential function of CA in a more controlled experimental setting and outside narrative context. Given the iconic properties of CA we hypothesized that this strategy could be used for efficient information transmission. Thus, we asked if use of CA increased with the increase in the information required to be communicated. Twenty-three deaf signers of LIS described unconnected images, which varied in the amount of information represented, to another player in a director–matcher game. Results revealed that participants used CA to communicate core information about the images and also increased the use of CA as images became informatively denser. The findings show that iconic features of CA can be used for referential function in addition to its depictive function outside narrative context and to achieve communicative efficiency.
  • Smeets, C. J. L. M., Ma, K. Y., Fisher, S. E., & Verbeek, D. S. (2021). Cerebellar developmental deficits underlie neurodegenerative disorder spinocerebellar ataxia type 23. Brain Pathology, 31(2), 239-252. doi:10.1111/bpa.12905.

    Abstract

    Spinocerebellar ataxia type 23 (SCA23) is a late‐onset neurodegenerative disorder characterized by slowly progressive gait and limb ataxia, for which there is no therapy available. It is caused by pathogenic variants in PDYN, which encodes prodynorphin (PDYN). PDYN is processed into the opioid peptides α‐neoendorphin and dynorphins (Dyn) A and B; inhibitory neurotransmitters that function in pain signaling, stress‐induced responses and addiction. Variants causing SCA23 mostly affect Dyn A, leading to loss of secondary structure and increased peptide stability. PDYNR212W mice express human PDYN containing the SCA23 variant p.R212W. These mice show progressive motor deficits from 3 months of age, climbing fiber (CF) deficits from 3 months of age, and Purkinje cell (PC) loss from 12 months of age. A mouse model for SCA1 showed similar CF deficits, and a recent study found additional developmental abnormalities, namely increased GABAergic interneuron connectivity and non‐cell autonomous disruption of PC function. As SCA23 mice show a similar pathology to SCA1 mice in adulthood, we hypothesized that SCA23 may also follow SCA1 pathology during development. Examining PDYNR212W cerebella during development, we uncovered developmental deficits from 2 weeks of age, namely a reduced number of GABAergic synapses on PC soma, possibly leading to the observed delay in early phase CF elimination between 2 and 3 weeks of age. Furthermore, CFs did not reach terminal height, leaving proximal PC dendrites open to be occupied by parallel fibers (PFs). The observed increase in vGlut1 protein—a marker for PF‐PC synapses—indicates that PFs indeed take over CF territory and have increased connectivity with PCs. Additionally, we detected altered expression of several critical Ca2+ channel subunits, potentially contributing to altered Ca2+ transients in PDYNR212W cerebella. These findings indicate that developmental abnormalities contribute to the SCA23 pathology and uncover a developmental role for PDYN in the cerebellum.
  • Smith, A. C., Monaghan, P., & Huettig, F. (2021). The effect of orthographic systems on the developing reading system: Typological and computational analyses. Psychological Review, 128(1), 125-159. doi:10.1037/rev0000257.

    Abstract

    Orthographic systems vary dramatically in the extent to which they encode a language’s phonological and lexico-semantic structure. Studies of the effects of orthographic transparency suggest that such variation is likely to have major implications for how the reading system operates. However, such studies have been unable to examine in isolation the contributory effect of transparency on reading due to co-varying linguistic or socio-cultural factors. We first investigated the phonological properties of languages using the range of the world’s orthographic systems (alphabetic; alphasyllabic; consonantal; syllabic; logographic), and found that, once geographical proximity is taken into account, phonological properties do not relate to orthographic system. We then explored the processing implications of orthographic variation by training a connectionist implementation of the triangle model of reading on the range of orthographic systems whilst controlling for phonological and semantic structure. We show that the triangle model is effective as a universal model of reading, able to replicate key behavioural and neuroscientific results. Importantly, the model also generates new predictions deriving from an explicit description of the effects of orthographic transparency on how reading is realised and defines the consequences of orthographic systems on reading processes.
  • Smits, R. (2000). Temporal distribution of information for human consonant recognition in VCV utterances. Journal of Phonetics, 28, 111-135. doi:10.006/jpho.2000.0107.

    Abstract

    The temporal distribution of perceptually relevant information for consonant recognition in British English VCVs is investigated. The information distribution in the vicinity of consonantal closure and release was measured by presenting initial and final portions, respectively, of naturally produced VCV utterances to listeners for categorization. A multidimensional scaling analysis of the results provided highly interpretable, four-dimensional geometrical representations of the confusion patterns in the categorization data. In addition, transmitted information as a function of truncation point was calculated for the features manner place and voicing. The effects of speaker, vowel context, stress, and distinctive feature on the resulting information distributions were tested statistically. It was found that, although all factors are significant, the location and spread of the distributions depends principally on the distinctive feature, i.e., the temporal distribution of perceptually relevant information is very different for the features manner, place, and voicing.
  • Snijders, T. M., Vosse, T., Kempen, G., Van Berkum, J. J. A., Petersson, K. M., & Hagoort, P. (2009). Retrieval and unification of syntactic structure in sentence comprehension: An fMRI study using word-category ambiguity. Cerebral Cortex, 19, 1493-1503. doi:10.1093/cercor/bhn187.

    Abstract

    Sentence comprehension requires the retrieval of single word information from long-term memory, and the integration of this information into multiword representations. The current functional magnetic resonance imaging study explored the hypothesis that the left posterior temporal gyrus supports the retrieval of lexical-syntactic information, whereas left inferior frontal gyrus (LIFG) contributes to syntactic unification. Twenty-eight subjects read sentences and word sequences containing word-category (noun–verb) ambiguous words at critical positions. Regions contributing to the syntactic unification process should show enhanced activation for sentences compared to words, and only within sentences display a larger signal for ambiguous than unambiguous conditions. The posterior LIFG showed exactly this predicted pattern, confirming our hypothesis that LIFG contributes to syntactic unification. The left posterior middle temporal gyrus was activated more for ambiguous than unambiguous conditions (main effect over both sentences and word sequences), as predicted for regions subserving the retrieval of lexical-syntactic information from memory. We conclude that understanding language involves the dynamic interplay between left inferior frontal and left posterior temporal regions.

    Additional information

    suppl1.pdf suppl2_dutch_stimulus.pdf
  • Snijders Blok, L., Goosen, Y. M., Haaften, L., Hulst, K., Fisher, S. E., Brunner, H. G., Egger, J. I. M., & Kleefstra, T. (2021). Speech‐language profiles in the context of cognitive and adaptive functioning inSATB2‐associated syndrome. Genes, Brain and Behavior, 20(7): e12761. doi:10.1111/gbb.12761.

    Abstract

    SATB2-associated syndrome (SAS) is a neurodevelopmental disorder caused by heterozygous pathogenic variants in the SATB2 gene, and is typically characterized by intellectual disability and severely impaired communication skills. The goal of this study was to contribute to the understanding of speech and language impairments in SAS, in the context of general developmental skills and cognitive and adaptive functioning. We performed detailed oral motor, speech and language profiling in combination with neuropsychological assessments in 23 individuals with a molecularly confirmed SAS diagnosis: 11 primarily verbal individuals and 12 primarily nonverbal individuals, independent of their ages. All individuals had severe receptive language delays. For all verbal individuals, we were able to define underlying speech conditions. While childhood apraxia of speech was most prevalent, oral motor problems appeared frequent as well and were more present in the nonverbal group than in the verbal group. For seven individuals, age-appropriate Wechsler indices could be derived, showing that the level of intellectual functioning of these individuals varied from moderate–mild ID to mild ID-borderline intellectual functioning. Assessments of adaptive functioning with the Vineland Screener showed relatively high scores on the domain “daily functioning” and relatively low scores on the domain “communication” in most individuals. Altogether, this study provides a detailed delineation of oral motor, speech and language skills and neuropsychological functioning in individuals with SAS, and can provide families and caregivers with information to guide diagnosis, management and treatment approaches.

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    supporting information
  • Snijders, T. M., Kooijman, V., Cutler, A., & Hagoort, P. (2007). Neurophysiological evidence of delayed segmentation in a foreign language. Brain Research, 1178, 106-113. doi:10.1016/j.brainres.2007.07.080.

    Abstract

    Previous studies have shown that segmentation skills are language-specific, making it difficult to segment continuous speech in an unfamiliar language into its component words. Here we present the first study capturing the delay in segmentation and recognition in the foreign listener using ERPs. We compared the ability of Dutch adults and of English adults without knowledge of Dutch (‘foreign listeners’) to segment familiarized words from continuous Dutch speech. We used the known effect of repetition on the event-related potential (ERP) as an index of recognition of words in continuous speech. Our results show that word repetitions in isolation are recognized with equivalent facility by native and foreign listeners, but word repetitions in continuous speech are not. First, words familiarized in isolation are recognized faster by native than by foreign listeners when they are repeated in continuous speech. Second, when words that have previously been heard only in a continuous-speech context re-occur in continuous speech, the repetition is detected by native listeners, but is not detected by foreign listeners. A preceding speech context facilitates word recognition for native listeners, but delays or even inhibits word recognition for foreign listeners. We propose that the apparent difference in segmentation rate between native and foreign listeners is grounded in the difference in language-specific skills available to the listeners.
  • Snijders Blok, L., Vino, A., Den Hoed, J., Underhill, H. R., Monteil, D., Li, H., Reynoso Santos, F. J., Chung, W. K., Amaral, M. D., Schnur, R. E., Santiago-Sim, T., Si, Y., Brunner, H. G., Kleefstra, T., & Fisher, S. E. (2021). Heterozygous variants that disturb the transcriptional repressor activity of FOXP4 cause a developmental disorder with speech/language delays and multiple congenital abnormalities. Genetics in Medicine, 23, 534-542. doi:10.1038/s41436-020-01016-6.

    Abstract

    Heterozygous pathogenic variants in various FOXP genes cause specific developmental disorders. The phenotype associated with heterozygous variants in FOXP4 has not been previously described.
    We assembled a cohort of eight individuals with heterozygous and mostly de novo variants in FOXP4: seven individuals with six different missense variants and one individual with a frameshift variant. We collected clinical data to delineate the phenotypic spectrum, and used in silico analyses and functional cell-based assays to assess pathogenicity of the variants.
    We collected clinical data for six individuals: five individuals with a missense variant in the forkhead box DNA-binding domain of FOXP4, and one individual with a truncating variant. Overlapping features included speech and language delays, growth abnormalities, congenital diaphragmatic hernia, cervical spine abnormalities, and ptosis. Luciferase assays showed loss-of-function effects for all these variants, and aberrant subcellular localization patterns were seen in a subset. The remaining two missense variants were located outside the functional domains of FOXP4, and showed transcriptional repressor capacities and localization patterns similar to the wild-type protein.
    Collectively, our findings show that heterozygous loss-of-function variants in FOXP4 are associated with an autosomal dominant neurodevelopmental disorder with speech/language delays, growth defects, and variable congenital abnormalities.
  • Snowdon, C. T., & Cronin, K. A. (2007). Cooperative breeders do cooperate. Behavioural Processes, 76, 138-141. doi:10.1016/j.beproc.2007.01.016.

    Abstract

    Bergmuller et al. (2007) make an important contribution to studies of cooperative breeding and provide a theoretical basis for linking the evolution of cooperative breeding with cooperative behavior.We have long been involved in empirical research on the only family of nonhuman primates to exhibit cooperative breeding, the Callitrichidae, which includes marmosets and tamarins, with studies in both field and captive contexts. In this paper we expand on three themes from Bergm¨uller et al. (2007) with empirical data. First we provide data in support of the importance of helpers and the specific benefits that helpers can gain in terms of fitness. Second, we suggest that mechanisms of rewarding helpers are more common and more effective in maintaining cooperative breeding than punishments. Third, we present a summary of our own research on cooperative behavior in cotton-top tamarins (Saguinus oedipus) where we find greater success in cooperative problem solving than has been reported for non-cooperatively breeding species.
  • Soares, S. M. P., & Rothman, J. (2021). Cognitive states in third language acquisition and beyond: theoretical and methodological paths forward. Linguistic Approaches to Bilingualism, 11(1), 89-95. doi:10.1075/lab.20080.per.
  • Sønderby, I. E., Van der Meer, D., Moreau, C., Kaufmann, T., Walters, G. B., Ellegaard, M., Abdellaoui, A., Ames, D., Amunts, K., Andersson, M., Armstrong, N. J., Bernard, M., Blackburn, N. B., Blangero, J., Boomsma, D. I., Brodaty, H., Brouwer, R. M., Bülow, R., Bøen, R., Cahn, W. and 125 moreSønderby, I. E., Van der Meer, D., Moreau, C., Kaufmann, T., Walters, G. B., Ellegaard, M., Abdellaoui, A., Ames, D., Amunts, K., Andersson, M., Armstrong, N. J., Bernard, M., Blackburn, N. B., Blangero, J., Boomsma, D. I., Brodaty, H., Brouwer, R. M., Bülow, R., Bøen, R., Cahn, W., Calhoun, V. D., Caspers, S., Ching, C. R. K., Cichon, S., Ciufolini, S., Crespo-Facorro, B., Curran, J. E., Dale, A. M., Dalvie, S., Dazzan, P., De Geus, E. J. C., De Zubicaray, G. I., De Zwarte, S. M. C., Desrivieres, S., Doherty, J. L., Donohoe, G., Draganski, B., Ehrlich, S., Eising, E., Espeseth, T., Fejgin, K., Fisher, S. E., Fladby, T., Frei, O., Frouin, V., Fukunaga, M., Gareau, T., Ge, T., Glahn, D. C., Grabe, H. J., Groenewold, N. A., Gústafsson, Ó., Haavik, J., Haberg, A. K., Hall, J., Hashimoto, R., Hehir-Kwa, J. Y., Hibar, D. P., Hillegers, M. H. J., Hoffmann, P., Holleran, L., Holmes, A. J., Homuth, G., Hottenga, J.-J., Hulshoff Pol, H. E., Ikeda, M., Jahanshad, N., Jockwitz, C., Johansson, S., Jönsson, E. G., Jørgensen, N. R., Kikuchi, M., Knowles, E. E. M., Kumar, K., Le Hellard, S., Leu, C., Linden, D. E., Liu, J., Lundervold, A., Lundervold, A. J., Maillard, A. M., Martin, N. G., Martin-Brevet, S., Mather, K. A., Mathias, S. R., McMahon, K. L., McRae, A. F., Medland, S. E., Meyer-Lindenberg, A., Moberget, T., Modenato, C., Monereo Sánchez, J., Morris, D. W., Mühleisen, T. W., Murray, R. M., Nielsen, J., Nordvik, J. E., Nyberg, L., Olde Loohuis, L. M., Ophoff, R. A., Owen, M. J., Paus, T., Pausova, Z., Peralta, J. M., Pike, B., Prieto, C., Quinlan, E. B., Reinbold, C. S., Reis Marques, T., Rucker, J. J. H., Sachdev, P. S., Sando, S. B., Schofield, P. R., Schork, A. J., Schumann, G., Shin, J., Shumskaya, E., Silva, A. I., Sisodiya, S. M., Steen, V. M., Stein, D. J., Strike, L. T., Suzuki, I. K., Tamnes, C. K., Teumer, A., Thalamuthu, A., Tordesillas-Gutiérrez, D., Uhlmann, A., Úlfarsson, M. Ö., Van 't Ent, D., Van den Bree, M. B. M., Vanderhaeghen, P., Vassos, E., Wen, W., Wittfeld, K., Wright, M. J., Agartz, I., Djurovic, S., Westlye, L. T., Stefánsson, H., Stefánsson, K., Jacquemont, S., Thompson, P. M., Andreassen, O. A., & the ENIGMA-CNV working group (2021). 1q21.1 distal copy number variants are associated with cerebral and cognitive alterations in humans. Translational Psychiatry, 11: 182. doi:10.1038/s41398-021-01213-0.

    Abstract

    Low-frequency 1q21.1 distal deletion and duplication copy number variant (CNV) carriers are predisposed to multiple neurodevelopmental disorders, including schizophrenia, autism and intellectual disability. Human carriers display a high prevalence of micro- and macrocephaly in deletion and duplication carriers, respectively. The underlying brain structural diversity remains largely unknown. We systematically called CNVs in 38 cohorts from the large-scale ENIGMA-CNV collaboration and the UK Biobank and identified 28 1q21.1 distal deletion and 22 duplication carriers and 37,088 non-carriers (48% male) derived from 15 distinct magnetic resonance imaging scanner sites. With standardized methods, we compared subcortical and cortical brain measures (all) and cognitive performance (UK Biobank only) between carrier groups also testing for mediation of brain structure on cognition. We identified positive dosage effects of copy number on intracranial volume (ICV) and total cortical surface area, with the largest effects in frontal and cingulate cortices, and negative dosage effects on caudate and hippocampal volumes. The carriers displayed distinct cognitive deficit profiles in cognitive tasks from the UK Biobank with intermediate decreases in duplication carriers and somewhat larger in deletion carriers—the latter potentially mediated by ICV or cortical surface area. These results shed light on pathobiological mechanisms of neurodevelopmental disorders, by demonstrating gene dose effect on specific brain structures and effect on cognitive function.

    Additional information

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  • Speed, L., Chen, J., Huettig, F., & Majid, A. (2021). Classifier categories reflect, but do not affect conceptual organization. Journal of Experimental Psychology: Learning, Memory, and Cognition, 47(4), 625-640. doi:10.1037/xlm0000967.

    Abstract

    Do we structure object-related conceptual information according to real-world sensorimotor experience, or can it also be shaped by linguistic information? This study investigates whether a feature of language coded in grammar—numeral classifiers—affects the conceptual representation of objects. We compared speakers of Mandarin (a classifier language) with speakers of Dutch (a language without classifiers) on how they judged object similarity in four studies. In the first three studies, participants had to rate how similar a target object was to four comparison objects, one of which shared a classifier with the target. Objects were presented as either words or pictures. Overall, the target object was always rated as most similar to the object with the shared classifier, but this was the case regardless of the language of the participant. In a final study employing a successive pile-sorting task, we also found that the underlying object concepts were similar for speakers of Mandarin and Dutch. Speakers of a non-classifier language are therefore sensitive to the same conceptual similarities that underlie classifier systems in a classifier language. Classifier systems may therefore reflect conceptual structure, rather than shape it.
  • Spiteri, E., Konopka, G., Coppola, G., Bomar, J., Oldham, M., Ou, J., Vernes, S. C., Fisher, S. E., Ren, B., & Geschwind, D. (2007). Identification of the transcriptional targets of FOXP2, a gene linked to speech and language, in developing human brain. American Journal of Human Genetics, 81(6), 1144-1157. doi:10.1086/522237.

    Abstract

    Mutations in FOXP2, a member of the forkhead family of transcription factor genes, are the only known cause of developmental speech and language disorders in humans. To date, there are no known targets of human FOXP2 in the nervous system. The identification of FOXP2 targets in the developing human brain, therefore, provides a unique tool with which to explore the development of human language and speech. Here, we define FOXP2 targets in human basal ganglia (BG) and inferior frontal cortex (IFC) by use of chromatin immunoprecipitation followed by microarray analysis (ChIP-chip) and validate the functional regulation of targets in vitro. ChIP-chip identified 285 FOXP2 targets in fetal human brain; statistically significant overlap of targets in BG and IFC indicates a core set of 34 transcriptional targets of FOXP2. We identified targets specific to IFC or BG that were not observed in lung, suggesting important regional and tissue differences in FOXP2 activity. Many target genes are known to play critical roles in specific aspects of central nervous system patterning or development, such as neurite outgrowth, as well as plasticity. Subsets of the FOXP2 transcriptional targets are either under positive selection in humans or differentially expressed between human and chimpanzee brain. This is the first ChIP-chip study to use human brain tissue, making the FOXP2-target genes identified in these studies important to understanding the pathways regulating speech and language in the developing human brain. These data provide the first insight into the functional network of genes directly regulated by FOXP2 in human brain and by evolutionary comparisons, highlighting genes likely to be involved in the development of human higher-order cognitive processes.
  • Spychalska, M., Reimer, L., Schumacher, P. B., & Werning, M. (2021). The cost of the epistemic step: Investigating scalar implicatures in full and partial information contexts. Frontiers in Psychology, 12: 679491. doi:10.3389/fpsyg.2021.679491.

    Abstract

    We present the first ERP experiments that test the online processing of the scalar implicature some ⇝ not all in contexts where the speaker competence assumption is violated. Participants observe game scenarios with four open cards on the table and two closed cards outside of the table, while listening to statements made by a virtual player. In the full access context, the player makes a fully informed statement by referring only to the open cards, as cards on the table; in the partial access context, she makes a partially informed statement by referring to the whole set of cards, as cards in the game. If all of the open cards contain a given object X (Fullset condition), then some cards on the table contain Xs is inconsistent with the not all reading, whereas it is unknown whether some cards in the game contain X is consistent with this reading. If only a subset of the open cards contains X (Subset condition), then both utterances are known to be consistent with the not all implicature. Differential effects are observed depending on the quantifier reading adopted by the participant: For those participants who adopt the not all reading in the full access context, but not in the partial access context (weak pragmatic reading), a late posterior negativity effect is observed in the partial access context for the Fullset relative to the Subset condition. This effect is argued to reflect inference-driven context retrieval and monitoring processes related to epistemic reasoning involved in evaluating the competence assumption. By contrast, for participants who adopt the logical interpretation of some (some and possibly all), an N400 effect is observed in the partial access context, when comparing the Subset against the Fullset condition, which is argued to result from the competition between the two quantifying expressions some cards on the table and some cards in the game functioning in the experiment as scalar alternatives.

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  • Stark, B. C., Clough, S., & Duff, M. (2021). Suggestions for improving the investigation of gesture in aphasia. Journal of Speech, Language, and Hearing Research, 64(10), 4004-4013. doi:10.1044/2021_JSLHR-21-00125.

    Abstract

    Purpose: When we speak, we gesture, and indeed, persons with aphasia gesture more frequently. The reason(s) for this is still being investigated, spurring an increase in the number of studies of gesture in persons with aphasia. As the number of studies increases, so too does the need for a shared set of best practices for gesture research in aphasia. After briefly reviewing the importance and use of gesture in persons with aphasia, this viewpoint puts forth methodological and design considerations when evaluating gesture in persons with aphasia.
    Method & Results: We explore several different design and methodological considerations for gesture research specific to persons with aphasia, such as video angle specifications, data collection techniques, and analysis considerations. The goal of these suggestions is to develop transparent and reproducible methods for evaluating gesture in aphasia to build a solid foundation for continued work in this area.
    Conclusions: We have proposed that it is critical to evaluate multimodal communication in a methodologically robust way to facilitate increased knowledge about the relationship of gesture to spoken language, cognition, and to other aspects of living with aphasia and recovery from aphasia. We conclude by postulating future directions for gesture research in aphasia.
  • Stevelink, R., Luykx, J. J., Lin, B. D., Leu, C., Lal, D., Smith, A. W., Schijven, D., Carpay, J. A., Rademaker, K., Baldez, R., A., R., Devinsky, O., Braun, K. P. J., Jansen, F. E., Smit, D. J. A., Koeleman, B. P. C., International League Against Epilepsy Consortium on Complex Epilepsies, & Epi25 Collaborative (2021). Shared genetic basis between genetic generalized epilepsy and background electroencephalographic oscillations. Epilepsia, 62(7), 1518-1527. doi:10.1111/epi.16922.

    Abstract

    Abstract Objective Paroxysmal epileptiform abnormalities on electroencephalography (EEG) are the hallmark of epilepsies, but it is uncertain to what extent epilepsy and background EEG oscillations share neurobiological underpinnings. Here, we aimed to assess the genetic correlation between epilepsy and background EEG oscillations. Methods Confounding factors, including the heterogeneous etiology of epilepsies and medication effects, hamper studies on background brain activity in people with epilepsy. To overcome this limitation, we compared genetic data from a genome-wide association study (GWAS) on epilepsy (n = 12 803 people with epilepsy and 24 218 controls) with that from a GWAS on background EEG (n = 8425 subjects without epilepsy), in which background EEG oscillation power was quantified in four different frequency bands: alpha, beta, delta, and theta. We replicated our findings in an independent epilepsy replication dataset (n = 4851 people with epilepsy and 20 428 controls). To assess the genetic overlap between these phenotypes, we performed genetic correlation analyses using linkage disequilibrium score regression, polygenic risk scores, and Mendelian randomization analyses. Results Our analyses show strong genetic correlations of genetic generalized epilepsy (GGE) with background EEG oscillations, primarily in the beta frequency band. Furthermore, we show that subjects with higher beta and theta polygenic risk scores have a significantly higher risk of having generalized epilepsy. Mendelian randomization analyses suggest a causal effect of GGE genetic liability on beta oscillations. Significance Our results point to shared biological mechanisms underlying background EEG oscillations and the susceptibility for GGE, opening avenues to investigate the clinical utility of background EEG oscillations in the diagnostic workup of epilepsy.

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  • Stewart, A. J., Kidd, E., & Haigh, M. (2009). Early sensitivity to discourse-level anomalies: Evidence from self-paced reading. Discourse Processes, 46(1), 46-69. doi:10.1080/01638530802629091.

    Abstract

    Two word-by-word, self-paced reading experiments investigated the speed with which readers were sensitive to discourse-level anomalies. An account arguing for delayed sensitivity (Guzman & Klin, 2000 Guzman, A. E. and Klin, C. M. 2000. Maintaining global coherence in reading: The role of sentence boundaries.. Memory & Cognition, 28: 722–730. [PubMed], [Web of Science ®], [Google Scholar]) was contrasted with one allowing for rapid sensitivity (Myers & O'Brien, 1998 Myers, J. L. and O'Brien, E. J. 1998. Accessing the discourse representation during reading.. Discourse Processes, 26: 131–157. [Taylor & Francis Online], [Web of Science ®], [Google Scholar]). Anomalies related to spatial information (Experiment 1) and character-attribute information (Experiment 2) were examined. Both experiments found that readers displayed rapid sensitivity to the anomalous information. A reading time penalty was observed for the region of text containing the anomalous information. This finding is most compatible with an account of text processing whereby incoming words are rapidly evaluated with respect to prior context. They are not consistent with an account that argues for delayed integration. Results are discussed in light of their implications for competing models of text processing.
  • Stewart, A. J., Haigh, M., & Kidd, E. (2009). An investigation into the online processing of counterfactual and indicative conditionals. Quarterly Journal of Experimental Psychology, 62(11), 2113-2125. doi:10.1080/17470210902973106.

    Abstract

    The ability to represent conditional information is central to human cognition. In two self-paced reading experiments we investigated how readers process counterfactual conditionals (e.g., If Darren had been athletic, he could probably have played on the rugby team ) and indicative conditionals (e.g., If Darren is athletic, he probably plays on the rugby team ). In Experiment 1 we focused on how readers process counterfactual conditional sentences. We found that processing of the antecedent of counterfactual conditionals was rapidly constrained by prior context (i.e., knowing whether Darren was or was not athletic). A reading-time penalty was observed for the critical region of text comprising the last word of the antecedent and the first word of the consequent when the information in the antecedent did not fit with prior context. In Experiment 2 we contrasted counterfactual conditionals with indicative conditionals. For counterfactual conditionals we found the same effect on the critical region as we found in Experiment 1. In contrast, however, we found no evidence that processing of the antecedent of indicative conditionals was constrained by prior context. For indicative conditionals (but not for counterfactual conditionals), the results we report are consistent with the suppositional account of conditionals. We propose that current theories of conditionals need to be able to account for online processing differences between indicative and counterfactual conditionals
  • Stewart, A., Holler, J., & Kidd, E. (2007). Shallow processing of ambiguous pronouns: Evidence for delay. Quarterly Journal of Experimental Psychology, 60, 1680-1696. doi:10.1080/17470210601160807.
  • Stivers, T., & Majid, A. (2007). Questioning children: Interactional evidence of implicit bias in medical interviews. Social Psychology Quarterly, 70(4), 424-441.

    Abstract

    Social psychologists have shown experimentally that implicit race bias can influence an individual's behavior. Implicit bias has been suggested to be more subtle and less subject to cognitive control than more explicit forms of racial prejudice. Little is known about how implicit bias is manifest in naturally occurring social interaction. This study examines the factors associated with physicians selecting children rather than parents to answer questions in pediatric interviews about routine childhood illnesses. Analysis of the data using a Generalized Linear Latent and Mixed Model demonstrates a significant effect of parent race and education on whether physicians select children to answer questions. Black children and Latino children of low-education parents are less likely to be selected to answer questions than their same aged white peers irrespective of education. One way that implicit bias manifests itself in naturally occurring interaction may be through the process of speaker selection during questioning.
  • Stivers, T., Enfield, N. J., Brown, P., Englert, C., Hayashi, M., Heinemann, T., Hoymann, G., Rossano, F., De Ruiter, J. P., Yoon, K.-E., & Levinson, S. C. (2009). Universals and cultural variation in turn-taking in conversation. Proceedings of the National Academy of Sciences of the United States of America, 106 (26), 10587-10592. doi:10.1073/pnas.0903616106.

    Abstract

    Informal verbal interaction is the core matrix for human social life. A mechanism for coordinating this basic mode of interaction is a system of turn-taking that regulates who is to speak and when. Yet relatively little is known about how this system varies across cultures. The anthropological literature reports significant cultural differences in the timing of turn-taking in ordinary conversation. We test these claims and show that in fact there are striking universals in the underlying pattern of response latency in conversation. Using a worldwide sample of 10 languages drawn from traditional indigenous communities to major world languages, we show that all of the languages tested provide clear evidence for a general avoidance of overlapping talk and a minimization of silence between conversational turns. In addition, all of the languages show the same factors explaining within-language variation in speed of response. We do, however, find differences across the languages in the average gap between turns, within a range of 250 ms from the cross-language mean. We believe that a natural sensitivity to these tempo differences leads to a subjective perception of dramatic or even fundamental differences as offered in ethnographic reports of conversational style. Our empirical evidence suggests robust human universals in this domain, where local variations are quantitative only, pointing to a single shared infrastructure for language use with likely ethological foundations.

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  • Suomi, K., McQueen, J. M., & Cutler, A. (1997). Vowel harmony and speech segmentation in Finnish. Journal of Memory and Language, 36, 422-444. doi:10.1006/jmla.1996.2495.

    Abstract

    Finnish vowel harmony rules require that if the vowel in the first syllable of a word belongs to one of two vowel sets, then all subsequent vowels in that word must belong either to the same set or to a neutral set. A harmony mismatch between two syllables containing vowels from the opposing sets thus signals a likely word boundary. We report five experiments showing that Finnish listeners can exploit this information in an on-line speech segmentation task. Listeners found it easier to detect words likehymyat the end of the nonsense stringpuhymy(where there is a harmony mismatch between the first two syllables) than in the stringpyhymy(where there is no mismatch). There was no such effect, however, when the target words appeared at the beginning of the nonsense string (e.g.,hymypuvshymypy). Stronger harmony effects were found for targets containing front harmony vowels (e.g.,hymy) than for targets containing back harmony vowels (e.g.,paloinkypaloandkupalo). The same pattern of results appeared whether target position within the string was predictable or unpredictable. Harmony mismatch thus appears to provide a useful segmentation cue for the detection of word onsets in Finnish speech.
  • Sutcliffe, D. J., Dinasarapu, A. R., Visser, J. E., Den Hoed, J., Seifar, F., Joshi, P., Ceballos-Picot, I., Sardar, T., Hess, E. J., Sun, Y. V., Wen, Z., Zwick, M. E., & Jinnah, H. A. (2021). Induced pluripotent stem cells from subjects with Lesch-Nyhan disease. Scientific Reports, 11: 8523. doi:10.1038/s41598-021-87955-9.

    Abstract

    Lesch-Nyhan disease (LND) is an inherited disorder caused by pathogenic variants in the HPRT1 gene, which encodes the purine recycling enzyme hypoxanthine–guanine phosphoribosyltransferase (HGprt). We generated 6 induced pluripotent stem cell (iPSC) lines from 3 individuals with LND, along with 6 control lines from 3 normal individuals. All 12 lines had the characteristics of pluripotent stem cells, as assessed by immunostaining for pluripotency markers, expression of pluripotency genes, and differentiation into the 3 primary germ cell layers. Gene expression profiling with RNAseq demonstrated significant heterogeneity among the lines. Despite this heterogeneity, several anticipated abnormalities were readily detectable across all LND lines, including reduced HPRT1 mRNA. Several unexpected abnormalities were also consistently detectable across the LND lines, including decreases in FAR2P1 and increases in RNF39. Shotgun proteomics also demonstrated several expected abnormalities in the LND lines, such as absence of HGprt protein. The proteomics study also revealed several unexpected abnormalities across the LND lines, including increases in GNAO1 decreases in NSE4A. There was a good but partial correlation between abnormalities revealed by the RNAseq and proteomics methods. Finally, functional studies demonstrated LND lines had no HGprt enzyme activity and resistance to the toxic pro-drug 6-thioguanine. Intracellular purines in the LND lines were normal, but they did not recycle hypoxanthine. These cells provide a novel resource to reveal insights into the relevance of heterogeneity among iPSC lines and applications for modeling LND.

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  • Swaab, T. Y., Brown, C. M., & Hagoort, P. (1997). Spoken sentence comprehension in aphasia: Event-related potential evidence for a lexical integration deficit. Journal of Cognitive Neuroscience, 9(1), 39-66.

    Abstract

    In this study the N400 component of the event-related potential was used to investigate spoken sentence understanding in Broca's and Wernicke's aphasics. The aim of the study was to determine whether spoken sentence comprehension problems in these patients might result from a deficit in the on-line integration of lexical information. Subjects listened to sentences spoken at a normal rate. In half of these sentences, the meaning of the final word of the sentence matched the semantic specifications of the preceding sentence context. In the other half of the sentences, the sentence-final word was anomalous with respect to the preceding sentence context. The N400 was measured to the sentence-final words in both conditions. The results for the aphasic patients (n = 14) were analyzed according to the severity of their comprehension deficit and compared to a group of 12 neurologically unimpaired age-matched controls, as well as a group of 6 nonaphasic patients with a lesion in the right hemisphere. The nonaphasic brain damaged patients and the aphasic patients with a light comprehension deficit (high comprehenders, n = 7) showed an N400 effect that was comparable to that of the neurologically unimpaired subjects. In the aphasic patients with a moderate to severe comprehension deficit (low comprehenders, n = 7), a reduction and delay of the N400 effect was obtained. In addition, the P300 component was measured in a classical oddball paradigm, in which subjects were asked to count infrequent low tones in a random series of high and low tones. No correlation was found between the occurrence of N400 and P300 effects, indicating that changes in the N400 results were related to the patients' language deficit. Overall, the pattern of results was compatible with the idea that aphasic patients with moderate to severe comprehension problems are impaired in the integration of lexical information into a higher order representation of the preceding sentence context.
  • Swingley, D., & Aslin, R. N. (2007). Lexical competition in young children's word learning. Cognitive Psychology, 54(2), 99-132.

    Abstract

    In two experiments, 1.5-year-olds were taught novel words whose sound patterns were phonologically similar to familiar words (novel neighbors) or were not (novel nonneighbors). Learning was tested using a picture-fixation task. In both experiments, children learned the novel nonneighbors but not the novel neighbors. In addition, exposure to the novel neighbors impaired recognition performance on familiar neighbors. Finally, children did not spontaneously use phonological differences to infer that a novel word referred to a novel object. Thus, lexical competition—inhibitory interaction among words in speech comprehension—can prevent children from using their full phonological sensitivity in judging words as novel. These results suggest that word learning in young children, as in adults, relies not only on the discrimination and identification of phonetic categories, but also on evaluating the likelihood that an utterance conveys a new word.
  • Swingley, D. (2007). Lexical exposure and word-from encoding in 1.5-year-olds. Developmental Psychology, 43(2), 454-464. doi:10.1037/0012-1649.43.2.454.

    Abstract

    In this study, 1.5-year-olds were taught a novel word. Some children were familiarized with the word's phonological form before learning the word's meaning. Fidelity of phonological encoding was tested in a picture-fixation task using correctly pronounced and mispronounced stimuli. Only children with additional exposure in familiarization showed reduced recognition performance given slight mispronunciations relative to correct pronunciations; children with fewer exposures did not. Mathematical modeling of vocabulary exposure indicated that children may hear thousands of words frequently enough for accurate encoding. The results provide evidence compatible with partial failure of phonological encoding at 19 months of age, demonstrate that this limitation in learning does not always hinder word recognition, and show the value of infants' word-form encoding in early lexical development.
  • Swingley, D., & Aslin, R. N. (2000). Spoken word recognition and lexical representation in very young children. Cognition, 76, 147-166. doi:10.1016/S0010-0277(00)00081-0.

    Abstract

    Although children's knowledge of the sound patterns of words has been a focus of debate for many years, little is known about the lexical representations very young children use in word recognition. In particular, researchers have questioned the degree of specificity encoded in early lexical representations. The current study addressed this issue by presenting 18–23-month-olds with object labels that were either correctly pronounced, or mispronounced. Mispronunciations involved replacement of one segment with a similar segment, as in ‘baby–vaby’. Children heard sentences containing these words while viewing two pictures, one of which was the referent of the sentence. Analyses of children's eye movements showed that children recognized the spoken words in both conditions, but that recognition was significantly poorer when words were mispronounced. The effects of mispronunciation on recognition were unrelated to age or to spoken vocabulary size. The results suggest that children's representations of familiar words are phonetically well-specified, and that this specification may not be a consequence of the need to differentiate similar words in production.
  • Tagliapietra, L., Fanari, R., De Candia, C., & Tabossi, P. (2009). Phonotactic regularities in the segmentation of spoken Italian. Quarterly Journal of Experimental Psychology, 62(2), 392 -415. doi:10.1080/17470210801907379.

    Abstract

    Five word-spotting experiments explored the role of consonantal and vocalic phonotactic cues in the segmentation of spoken Italian. The first set of experiments tested listeners' sensitivity to phonotactic constraints cueing syllable boundaries. Participants were slower in spotting words in nonsense strings when target onsets were misaligned (e.g., lago in ri.blago) than when they were aligned (e.g., lago in rin.lago) with phonotactically determined syllabic boundaries. This effect held also for sequences that occur only word-medially (e.g., /tl/ in ri.tlago), and competition effects could not account for the disadvantage in the misaligned condition. Similarly, target detections were slower when their offsets were misaligned (e.g., cittaacute in cittaacuteu.ba) than when they were aligned (e.g., cittaacute in cittaacute.oba) with a phonotactic syllabic boundary. The second set of experiments tested listeners' sensitivity to phonotactic cues, which specifically signal lexical (and not just syllable) boundaries. Results corroborate the role of syllabic information in speech segmentation and suggest that Italian listeners make little use of additional phonotactic information that specifically cues word boundaries.

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  • Tagliapietra, L., Fanari, R., Collina, S., & Tabossi, P. (2009). Syllabic effects in Italian lexical access. Journal of Psycholinguistic Research, 38(6), 511-526. doi:10.1007/s10936-009-9116-4.

    Abstract

    Two cross-modal priming experiments tested whether lexical access is constrained by syllabic structure in Italian. Results extend the available Italian data on the processing of stressed syllables showing that syllabic information restricts the set of candidates to those structurally consistent with the intended word (Experiment 1). Lexical access, however, takes place as soon as possible and it is not delayed till the incoming input corresponds to the first syllable of the word. And, the initial activated set includes candidates whose syllabic structure does not match the intended word (Experiment 2). The present data challenge the early hypothesis that in Romance languages syllables are the units for lexical access during spoken word recognition. The implications of the results for our understanding of the role of syllabic information in language processing are discussed.
  • Takashima, A., Nieuwenhuis, I. L. C., Rijpkema, M., Petersson, K. M., Jensen, O., & Fernández, G. (2007). Memory trace stabilization leads to large-scale changes in the retrieval network: A functional MRI study on associative memory. Learning & Memory, 14, 472-479. doi:10.1101/lm.605607.

    Abstract

    Spaced learning with time to consolidate leads to more stabile memory traces. However, little is known about the neural correlates of trace stabilization, especially in humans. The present fMRI study contrasted retrieval activity of two well-learned sets of face-location associations, one learned in a massed style and tested on the day of learning (i.e., labile condition) and another learned in a spaced scheme over the course of one week (i.e., stabilized condition). Both sets of associations were retrieved equally well, but the retrieval of stabilized association was faster and accompanied by large-scale changes in the network supporting retrieval. Cued recall of stabilized as compared with labile associations was accompanied by increased activity in the precuneus, the ventromedial prefrontal cortex, the bilateral temporal pole, and left temporo–parietal junction. Conversely, memory representational areas such as the fusiform gyrus for faces and the posterior parietal cortex for locations did not change their activity with stabilization. The changes in activation in the precuneus, which also showed increased connectivity with the fusiform area, are likely to be related to the spatial nature of our task. The activation increase in the ventromedial prefrontal cortex, on the other hand, might reflect a general function in stabilized memory retrieval. This area might succeed the hippocampus in linking distributed neocortical representations.
  • Tanenhaus, M. K., Magnuson, J. S., Dahan, D., & Chaimbers, G. (2000). Eye movements and lexical access in spoken-language comprehension: evaluating a linking hypothesis between fixations and linguistic processing. Journal of Psycholinguistic Research, 29, 557-580. doi:10.1023/A:1026464108329.

    Abstract

    A growing number of researchers in the sentence processing community are using eye movements to address issues in spoken language comprehension. Experiments using this paradigm have shown that visually presented referential information, including properties of referents relevant to specific actions, influences even the earliest moments of syntactic processing. Methodological concerns about task-specific strategies and the linking hypothesis between eye movements and linguistic processing are identified and discussed. These concerns are addressed in a review of recent studies of spoken word recognition which introduce and evaluate a detailed linking hypothesis between eye movements and lexical access. The results provide evidence about the time course of lexical activation that resolves some important theoretical issues in spoken-word recognition. They also demonstrate that fixations are sensitive to properties of the normal language-processing system that cannot be attributed to task-specific strategies
  • Tartaro, G., Takashima, A., & McQueen, J. M. (2021). Consolidation as a mechanism for word learning in sequential bilinguals. Bilingualism: Language and Cognition, 24(5), 864-878. doi:10.1017/S1366728921000286.

    Abstract

    First-language research suggests that new words, after initial episodic-memory encoding, are consolidated and hence become lexically integrated. We asked here if lexical consolidation, about word forms and meanings, occurs in a second language. Italian–English sequential bilinguals learned novel English-like words (e.g., apricon, taught to mean “stapler”). fMRI analyses failed to reveal a predicted shift, after consolidation time, from hippocampal to temporal neocortical activity. In a pause-detection task, responses to existing phonological competitors of learned words (e.g., apricot for apricon) were slowed down if the words had been learned two days earlier (i.e., after consolidation time) but not if they had been learned the same day. In a lexical-decision task, new words primed responses to semantically-related existing words (e.g., apricon-paper) whether the words were learned that day or two days earlier. Consolidation appears to support integration of words into the bilingual lexicon, possibly more rapidly for meanings than for forms.

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  • Ten Oever, S., & Martin, A. E. (2021). An oscillating computational model can track pseudo-rhythmic speech by using linguistic predictions. eLife, 10: e68066. doi:10.7554/eLife.68066.

    Abstract

    Neuronal oscillations putatively track speech in order to optimize sensory processing. However, it is unclear how isochronous brain oscillations can track pseudo-rhythmic speech input. Here we propose that oscillations can track pseudo-rhythmic speech when considering that speech time is dependent on content-based predictions flowing from internal language models. We show that temporal dynamics of speech are dependent on the predictability of words in a sentence. A computational model including oscillations, feedback, and inhibition is able to track pseudo-rhythmic speech input. As the model processes, it generates temporal phase codes, which are a candidate mechanism for carrying information forward in time. The model is optimally sensitive to the natural temporal speech dynamics and can explain empirical data on temporal speech illusions. Our results suggest that speech tracking does not have to rely only on the acoustics but could also exploit ongoing interactions between oscillations and constraints flowing from internal language models.
  • Ten Oever, S., Sack, A. T., Oehrn, C. R., & Axmacher, N. (2021). An engram of intentionally forgotten information. Nature Communications, 12: 6443. doi:10.1038/s41467-021-26713-x.

    Abstract

    Successful forgetting of unwanted memories is crucial for goal-directed behavior and mental wellbeing. While memory retention strengthens memory traces, it is unclear what happens to memory traces of events that are actively forgotten. Using intracranial EEG recordings from lateral temporal cortex, we find that memory traces for actively forgotten information are partially preserved and exhibit unique neural signatures. Memory traces of successfully remembered items show stronger encoding-retrieval similarity in gamma frequency patterns. By contrast, encoding-retrieval similarity of item-specific memory traces of actively forgotten items depend on activity at alpha/beta frequencies commonly associated with functional inhibition. Additional analyses revealed selective modification of item-specific patterns of connectivity and top-down information flow from dorsolateral prefrontal cortex to lateral temporal cortex in memory traces of intentionally forgotten items. These results suggest that intentional forgetting relies more on inhibitory top-down connections than intentional remembering, resulting in inhibitory memory traces with unique neural signatures and representational formats.

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  • Tendolkar, I., Arnold, J., Petersson, K. M., Weis, S., Brockhaus-Dumke, A., Van Eijndhoven, P., Buitelaar, J., & Fernández, G. (2007). Probing the neural correlates of associative memory formation: A parametrically analyzed event-related functional MRI study. Brain Research, 1142, 159-168. doi:10.1016/j.brainres.2007.01.040.

    Abstract

    The medial temporal lobe (MTL) is crucial for declarative memory formation, but the function of its subcomponents in associative memory formation remains controversial. Most functional imaging studies on this topic are based on a stepwise approach comparing a condition with and one without associative encoding. Extending this approach we applied additionally a parametric analysis by varying the amount of associative memory formation. We found a hippocampal subsequent memory effect of almost similar magnitude regardless of the amount of associations formed. By contrast, subsequent memory effects in rhinal and parahippocampal cortices were parametrically and positively modulated by the amount of associations formed. Our results indicate that the parahippocampal region supports associative memory formation as tested here and the hippocampus adds a general mnemonic operation. This pattern of results might suggest a new interpretation. Instead of having either a fixed division of labor between the hippocampus (associative memory formation) and the rhinal cortex (non-associative memory formation) or a functionally unitary MTL system, in which all substructures are contributing to memory formation in a similar way, we propose that the location where associations are formed within the MTL depends on the kind of associations bound: If visual single-dimension associations, as used here, can already be integrated within the parahippocampal region, the hippocampus might add a general purpose mnemonic operation only. In contrast, if associations have to be formed across widely distributed neocortical representations, the hippocampus may provide a binding operation in order to establish a coherent memory.
  • Ter Avest, I. J., & Mulder, K. (2009). The Acquisition of Gender Agreement in the Determiner Phrase by Bilingual Children. Toegepaste Taalwetenschap in Artikelen, 81(1), 133-142.
  • Terrill, A. (2007). [Review of ‘Andrew Pawley, Robert Attenborough, Jack Golson, and Robin Hide, eds. 2005. Papuan pasts: Cultural, linguistic and biological histories of Papuan-speaking people]. Oceanic Linguistics, 46(1), 313-321. doi:10.1353/ol.2007.0025.
  • Terrill, A. (2009). [Review of Felix K. Ameka, Alan Dench, and Nicholas Evans (eds). 2006. Catching language: The standing challenge of grammar writing]. Language Documentation & Conservation, 3(1), 132-137. Retrieved from http://hdl.handle.net/10125/4432.
  • Tesink, C. M. J. Y., Buitelaar, J. K., Petersson, K. M., Van der Gaag, R. J., Kan, C. C., Tendolkar, I., & Hagoort, P. (2009). Neural correlates of pragmatic language comprehension in autism disorders. Brain, 132, 1941-1952. doi:10.1093/brain/awp103.

    Abstract

    Difficulties with pragmatic aspects of communication are universal across individuals with autism spectrum disorders (ASDs). Here we focused on an aspect of pragmatic language comprehension that is relevant to social interaction in daily life: the integration of speaker characteristics inferred from the voice with the content of a message. Using functional magnetic resonance imaging (fMRI), we examined the neural correlates of the integration of voice-based inferences about the speaker’s age, gender or social background, and sentence content in adults with ASD and matched control participants. Relative to the control group, the ASD group showed increased activation in right inferior frontal gyrus (RIFG; Brodmann area 47) for speakerincongruent sentences compared to speaker-congruent sentences. Given that both groups performed behaviourally at a similar level on a debriefing interview outside the scanner, the increased activation in RIFG for the ASD group was interpreted as being compensatory in nature. It presumably reflects spill-over processing from the language dominant left hemisphere due to higher task demands faced by the participants with ASD when integrating speaker characteristics and the content of a spoken sentence. Furthermore, only the control group showed decreased activation for speaker-incongruent relative to speaker-congruent sentences in right ventral medial prefrontal cortex (vMPFC; Brodmann area 10), including right anterior cingulate cortex (ACC; Brodmann area 24/32). Since vMPFC is involved in self-referential processing related to judgments and inferences about self and others, the absence of such a modulation in vMPFC activation in the ASD group possibly points to atypical default self-referential mental activity in ASD. Our results show that in ASD compensatory mechanisms are necessary in implicit, low-level inferential processes in spoken language understanding. This indicates that pragmatic language problems in ASD are not restricted to high-level inferential processes, but encompass the most basic aspects of pragmatic language processing.
  • Tesink, C. M. J. Y., Petersson, K. M., Van Berkum, J. J. A., Van den Brink, D., Buitelaar, J. K., & Hagoort, P. (2009). Unification of speaker and meaning in language comprehension: An fMRI study. Journal of Cognitive Neuroscience, 21, 2085-2099. doi:10.1162/jocn.2008.21161.

    Abstract

    When interpreting a message, a listener takes into account several sources of linguistic and extralinguistic information. Here we focused on one particular form of extralinguistic information, certain speaker characteristics as conveyed by the voice. Using functional magnetic resonance imaging, we examined the neural structures involved in the unification of sentence meaning and voice-based inferences about the speaker's age, sex, or social background. We found enhanced activation in the inferior frontal gyrus bilaterally (BA 45/47) during listening to sentences whose meaning was incongruent with inferred speaker characteristics. Furthermore, our results showed an overlap in brain regions involved in unification of speaker-related information and those used for the unification of semantic and world knowledge information [inferior frontal gyrus bilaterally (BA 45/47) and left middle temporal gyrus (BA 21)]. These findings provide evidence for a shared neural unification system for linguistic and extralinguistic sources of information and extend the existing knowledge about the role of inferior frontal cortex as a crucial component for unification during language comprehension.
  • Theakston, A., & Rowland, C. F. (2009). Introduction to Special Issue: Cognitive approaches to language acquisition. Cognitive Linguistics, 20(3), 477-480. doi:10.1515/COGL.2009.021.
  • Theakston, A. L., & Rowland, C. F. (2009). The acquisition of auxiliary syntax: A longitudinal elicitation study. Part 1: Auxiliary BE. Journal of Speech, Language, and Hearing Research, 52, 1449-1470. doi:10.1044/1092-4388(2009/08-0037).

    Abstract

    Purpose: The question of how and when English-speaking children acquire auxiliaries is the subject of extensive debate. Some researchers posit the existence of innately given Universal Grammar principles to guide acquisition, although some aspects of the auxiliary system must be learned from the input. Others suggest that auxiliaries can be learned without Universal Grammar, citing evidence of piecemeal learning in their support. This study represents a unique attempt to trace the development of auxiliary syntax by using a longitudinal elicitation methodology. Method: Twelve English-speaking children participated in 3 tasks designed to elicit auxiliary BE in declaratives and yes/no and wh-questions. They completed each task 6 times in total between the ages of 2;10 (years;months) and 3;6. Results: The children’s levels of correct use of 2 forms of BE (is,are) differed according to auxiliary form and sentence structure, and these relations changed over development. An analysis of the children’s errors also revealed complex interactions between these factors. Conclusion: These data are problematic for existing accounts of auxiliary acquisition and highlight the need for researchers working within both generativist and constructivist frameworks to develop more detailed theories of acquisition that directly predict the pattern of acquisition observed.
  • Tilmatine, M., Hubers, F., & Hintz, F. (2021). Exploring individual differences in recognizing idiomatic expressions in context. Journal of Cognition, 4(1): 37. doi:10.5334/joc.183.

    Abstract

    Written language comprehension requires readers to integrate incoming information with stored mental knowledge to construct meaning. Literally plausible idiomatic expressions can activate both figurative and literal interpretations, which convey different meanings. Previous research has shown that contexts biasing the figurative or literal interpretation of an idiom can facilitate its processing. Moreover, there is evidence that processing of idiomatic expressions is subject to individual differences in linguistic knowledge and cognitive-linguistic skills. It is therefore conceivable that individuals vary in the extent to which they experience context-induced facilitation in processing idiomatic expressions. To explore the interplay between reader-related variables and contextual facilitation, we conducted a self-paced reading experiment. We recruited participants who had recently completed a battery of 33 behavioural tests measuring individual differences in linguistic knowledge, general cognitive skills and linguistic processing skills. In the present experiment, a subset of these participants read idiomatic expressions that were either presented in isolation or preceded by a figuratively or literally biasing context. We conducted analyses on the reading times of idiom-final nouns and the word thereafter (spill-over region) across the three conditions, including participants’ scores from the individual differences battery. Our results showed no main effect of the preceding context, but substantial variation in contextual facilitation between readers. We observed main effects of participants’ word reading ability and non-verbal intelligence on reading times as well as an interaction between condition and linguistic knowledge. We encourage interested researchers to exploit the present dataset for follow-up studies on individual differences in idiom processing.
  • Tilot, A. K., Khramtsova, E. A., Liang, D., Grasby, K. L., Jahanshad, N., Painter, J., Colodro-Conde, L., Bralten, J., Hibar, D. P., Lind, P. A., Liu, S., Brotman, S. M., Thompson, P. M., Medland, S. E., Macciardi, F., Stranger, B. E., Davis, L. K., Fisher, S. E., & Stein, J. L. (2021). The evolutionary history of common genetic variants influencing human cortical surface area. Cerebral Cortex, 31(4), 1873-1887. doi:10.1093/cercor/bhaa327.

    Abstract

    Structural brain changes along the lineage leading to modern Homo sapiens contributed to our distinctive cognitive and social abilities. However, the evolutionarily relevant molecular variants impacting key aspects of neuroanatomy are largely unknown. Here, we integrate evolutionary annotations of the genome at diverse timescales with common variant associations from large-scale neuroimaging genetic screens. We find that alleles with evidence of recent positive polygenic selection over the past 2000–3000 years are associated with increased surface area (SA) of the entire cortex, as well as specific regions, including those involved in spoken language and visual processing. Therefore, polygenic selective pressures impact the structure of specific cortical areas even over relatively recent timescales. Moreover, common sequence variation within human gained enhancers active in the prenatal cortex is associated with postnatal global SA. We show that such variation modulates the function of a regulatory element of the developmentally relevant transcription factor HEY2 in human neural progenitor cells and is associated with structural changes in the inferior frontal cortex. These results indicate that non-coding genomic regions active during prenatal cortical development are involved in the evolution of human brain structure and identify novel regulatory elements and genes impacting modern human brain structure.
  • Timpson, N. J., Tobias, J. H., Richards, J. B., Soranzo, N., Duncan, E. L., Sims, A.-M., Whittaker, P., Kumanduri, V., Zhai, G., Glaser, B., Eisman, J., Jones, G., Nicholson, G., Prince, R., Seeman, E., Spector, T. D., Brown, M. A., Peltonen, L., Smith, G. D., Deloukas, P. and 1 moreTimpson, N. J., Tobias, J. H., Richards, J. B., Soranzo, N., Duncan, E. L., Sims, A.-M., Whittaker, P., Kumanduri, V., Zhai, G., Glaser, B., Eisman, J., Jones, G., Nicholson, G., Prince, R., Seeman, E., Spector, T. D., Brown, M. A., Peltonen, L., Smith, G. D., Deloukas, P., & Evans, D. M. (2009). Common variants in the region around Osterix are associated with bone mineral density and growth in childhood. Human Molecular Genetics, 18(8), 1510-1517. doi:10.1093/hmg/ddp052.

    Abstract

    Peak bone mass achieved in adolescence is a determinant of bone mass in later life. In order to identify genetic variants affecting bone mineral density (BMD), we performed a genome-wide association study of BMD and related traits in 1518 children from the Avon Longitudinal Study of Parents and Children (ALSPAC). We compared results with a scan of 134 adults with high or low hip BMD. We identified associations with BMD in an area of chromosome 12 containing the Osterix (SP7) locus, a transcription factor responsible for regulating osteoblast differentiation (ALSPAC: P = 5.8 x 10(-4); Australia: P = 3.7 x 10(-4)). This region has previously shown evidence of association with adult hip and lumbar spine BMD in an Icelandic population, as well as nominal association in a UK population. A meta-analysis of these existing studies revealed strong association between SNPs in the Osterix region and adult lumbar spine BMD (P = 9.9 x 10(-11)). In light of these findings, we genotyped a further 3692 individuals from ALSPAC who had whole body BMD and confirmed the association in children as well (P = 5.4 x 10(-5)). Moreover, all SNPs were related to height in ALSPAC children, but not weight or body mass index, and when height was included as a covariate in the regression equation, the association with total body BMD was attenuated. We conclude that genetic variants in the region of Osterix are associated with BMD in children and adults probably through primary effects on growth.
  • Tomasello, M., Carpenter, M., & Liszkowski, U. (2007). A new look at infant pointing. Child Development, 78, 705-722. doi:10.1111/j.1467-8624.2007.01025.x.

    Abstract

    The current article proposes a new theory of infant pointing involving multiple layers of intentionality and shared intentionality. In the context of this theory, evidence is presented for a rich interpretation of prelinguistic communication, that is, one that posits that when 12-month-old infants point for an adult they are in some sense trying to influence her mental states. Moreover, evidence is also presented for a deeply social view in which infant pointing is best understood—on many levels and in many ways—as depending on uniquely human skills and motivations for cooperation and shared intentionality (e.g., joint intentions and attention with others). Children's early linguistic skills are built on this already existing platform of prelinguistic communication.
  • Torres Borda, L., Jadoul, Y., Rasilo, H., Salazar-Casals, A., & Ravignani, A. (2021). Vocal plasticity in harbour seal pups. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376(1840): 20200456. doi:10.1098/rstb.2020.0456.

    Abstract

    Vocal plasticity can occur in response to environmental and biological factors, including conspecifics' vocalizations and noise. Pinnipeds are one of the few mammalian groups capable of vocal learning, and are therefore relevant to understanding the evolution of vocal plasticity in humans and other animals. Here, we investigate the vocal plasticity of harbour seals (Phoca vitulina), a species with vocal learning abilities observed in adulthood but not puppyhood. To evaluate early mammalian vocal development, we tested 1–3 weeks-old seal pups. We tailored noise playbacks to this species and age to induce seal pups to shift their fundamental frequency (f0), rather than adapt call amplitude or temporal characteristics. We exposed individual pups to low- and high-intensity bandpass-filtered noise, which spanned—and masked—their typical range of f0; simultaneously, we recorded pups' spontaneous calls. Unlike most mammals, pups modified their vocalizations by lowering their f0 in response to increased noise. This modulation was precise and adapted to the particular experimental manipulation of the noise condition. In addition, higher levels of noise induced less dispersion around the mean f0, suggesting that pups may have actively focused their phonatory efforts to target lower frequencies. Noise did not seem to affect call amplitude. However, one seal showed two characteristics of the Lombard effect known for human speech in noise: significant increase in call amplitude and flattening of spectral tilt. Our relatively low noise levels may have favoured f0 modulation while inhibiting amplitude adjustments. This lowering of f0 is unusual, as most animals commonly display no such f0 shift. Our data represent a relatively rare case in mammalian neonates, and have implications for the evolution of vocal plasticity and vocal learning across species, including humans.

    Additional information

    supplement
  • Tourtouri, E. N., Delogu, F., & Crocker, M. W. (2021). Rational Redundancy in Referring Expressions: Evidence from Event-related Potentials. Cognitive Science, 45(12): e13071. doi:10.1111/cogs.13071.

    Abstract

    In referential communication, Grice's Maxim of Quantity is thought to imply that utterances conveying unnecessary information should incur comprehension difficulties. There is, however, considerable evidence that speakers frequently encode redundant information in their referring expressions, raising the question as to whether such overspecifications hinder listeners' processing. Evidence from previous work is inconclusive, and mostly comes from offline studies. In this article, we present two event-related potential (ERP) experiments, investigating the real-time comprehension of referring expressions that contain redundant adjectives in complex visual contexts. Our findings provide support for both Gricean and bounded-rational accounts. We argue that these seemingly incompatible results can be reconciled if common ground is taken into account. We propose a bounded-rational account of overspecification, according to which even redundant words can be beneficial to comprehension to the extent that they facilitate the reduction of listeners' uncertainty regarding the target referent.
  • Trilsbeek, P., & Van Uytvanck, D. (2009). Regional archives and community portals. IASA Journal, 32, 69-73.
  • Trompenaars, T., Kaluge, T. A., Sarabi, R., & De Swart, P. (2021). Cognitive animacy and its relation to linguistic animacy: Evidence from Japanese and Persian. Language Sciences, 86: 101399. doi:10.1016/j.langsci.2021.101399.

    Abstract

    Animacy, commonly defined as the distinction between living and non-living entities, is a useful notion in cognitive science and linguistics employed to describe and predict variation in psychological and linguistic behaviour. In the (psycho)linguistics literature we find linguistic animacy dichotomies which are (implicitly) assumed to correspond to biological dichotomies. We argue this is problematic, as it leaves us without a cognitively grounded, universal description for non-prototypical cases. We show that ‘animacy’ in language can be better understood as universally emerging from a gradual, cognitive property by collecting animacy ratings for a great range of nouns from Japanese and Persian. We used these cognitive ratings in turn to predict linguistic variation in these languages traditionally explained through dichotomous distinctions. We show that whilst (speakers of) languages may subtly differ in their conceptualisation of animacy, universality may be found in the process of mapping conceptual animacy to linguistic variation.
  • Trujillo, J. P., & Holler, J. (2021). The kinematics of social action: Visual signals provide cues for what interlocutors do in conversation. Brain Sciences, 11: 996. doi:10.3390/brainsci11080996.

    Abstract

    During natural conversation, people must quickly understand the meaning of what the other speaker is saying. This concerns not just the semantic content of an utterance, but also the social action (i.e., what the utterance is doing—requesting information, offering, evaluating, checking mutual understanding, etc.) that the utterance is performing. The multimodal nature of human language raises the question of whether visual signals may contribute to the rapid processing of such social actions. However, while previous research has shown that how we move reveals the intentions underlying instrumental actions, we do not know whether the intentions underlying fine-grained social actions in conversation are also revealed in our bodily movements. Using a corpus of dyadic conversations combined with manual annotation and motion tracking, we analyzed the kinematics of the torso, head, and hands during the asking of questions. Manual annotation categorized these questions into six more fine-grained social action types (i.e., request for information, other-initiated repair, understanding check, stance or sentiment, self-directed, active participation). We demonstrate, for the first time, that the kinematics of the torso, head and hands differ between some of these different social action categories based on a 900 ms time window that captures movements starting slightly prior to or within 600 ms after utterance onset. These results provide novel insights into the extent to which our intentions shape the way that we move, and provide new avenues for understanding how this phenomenon may facilitate the fast communication of meaning in conversational interaction, social action, and conversation

    Additional information

    analyses scripts
  • Trujillo, J. P., Ozyurek, A., Holler, J., & Drijvers, L. (2021). Speakers exhibit a multimodal Lombard effect in noise. Scientific Reports, 11: 16721. doi:10.1038/s41598-021-95791-0.

    Abstract

    In everyday conversation, we are often challenged with communicating in non-ideal settings, such as in noise. Increased speech intensity and larger mouth movements are used to overcome noise in constrained settings (the Lombard effect). How we adapt to noise in face-to-face interaction, the natural environment of human language use, where manual gestures are ubiquitous, is currently unknown. We asked Dutch adults to wear headphones with varying levels of multi-talker babble while attempting to communicate action verbs to one another. Using quantitative motion capture and acoustic analyses, we found that (1) noise is associated with increased speech intensity and enhanced gesture kinematics and mouth movements, and (2) acoustic modulation only occurs when gestures are not present, while kinematic modulation occurs regardless of co-occurring speech. Thus, in face-to-face encounters the Lombard effect is not constrained to speech but is a multimodal phenomenon where the visual channel carries most of the communicative burden.

    Additional information

    supplementary material
  • Trujillo, J. P., Ozyurek, A., Kan, C. C., Sheftel-Simanova, I., & Bekkering, H. (2021). Differences in the production and perception of communicative kinematics in autism. Autism Research, 14(12), 2640-2653. doi:10.1002/aur.2611.

    Abstract

    In human communication, social intentions and meaning are often revealed in the way we move. In this study, we investigate the flexibility of human communication in terms of kinematic modulation in a clinical population, namely, autistic individuals. The aim of this study was twofold: to assess (a) whether communicatively relevant kinematic features of gestures differ between autistic and neurotypical individuals, and (b) if autistic individuals use communicative kinematic modulation to support gesture recognition. We tested autistic and neurotypical individuals on a silent gesture production task and a gesture comprehension task. We measured movement during the gesture production task using a Kinect motion tracking device in order to determine if autistic individuals differed from neurotypical individuals in their gesture kinematics. For the gesture comprehension task, we assessed whether autistic individuals used communicatively relevant kinematic cues to support recognition. This was done by using stick-light figures as stimuli and testing for a correlation between the kinematics of these videos and recognition performance. We found that (a) silent gestures produced by autistic and neurotypical individuals differ in communicatively relevant kinematic features, such as the number of meaningful holds between movements, and (b) while autistic individuals are overall unimpaired at recognizing gestures, they processed repetition and complexity, measured as the amount of submovements perceived, differently than neurotypicals do. These findings highlight how subtle aspects of neurotypical behavior can be experienced differently by autistic individuals. They further demonstrate the relationship between movement kinematics and social interaction in high-functioning autistic individuals.

    Additional information

    supporting information
  • Tsoukala, C., Frank, S. L., Van Den Bosch, A., Valdés Kroff, J., & Broersma, M. (2021). Modeling the auxiliary phrase asymmetry in code-switched Spanish–English. Bilingualism: Language and Cognition, 24(2), 271-280. doi:10.1017/S1366728920000449.

    Abstract

    Spanish–English bilinguals rarely code-switch in the perfect structure between the Spanish auxiliary haber (“to have”) and the participle (e.g., “Ella ha voted”; “She has voted”). However, they are somewhat likely to switch in the progressive structure between the Spanish auxiliary estar (“to be”) and the participle (“Ella está voting”; “She is voting”). This phenomenon is known as the “auxiliary phrase asymmetry”. One hypothesis as to why this occurs is that estar has more semantic weight as it also functions as an independent verb, whereas haber is almost exclusively used as an auxiliary verb. To test this hypothesis, we employed a connectionist model that produces spontaneous code-switches. Through simulation experiments, we showed that i) the asymmetry emerges in the model and that ii) the asymmetry disappears when using haber also as a main verb, which adds semantic weight. Therefore, the lack of semantic weight of haber may indeed cause the asymmetry.
  • Tsoukala, C., Broersma, M., Van den Bosch, A., & Frank, S. L. (2021). Simulating code-switching using a neural network model of bilingual sentence production. Computational Brain & Behavior, 4, 87-100. doi:10.1007/s42113-020-00088-6.

    Abstract

    Code-switching is the alternation from one language to the other during bilingual speech. We present a novel method of researching this phenomenon using computational cognitive modeling. We trained a neural network of bilingual sentence production to simulate early balanced Spanish–English bilinguals, late speakers of English who have Spanish as a dominant native language, and late speakers of Spanish who have English as a dominant native language. The model produced code-switches even though it was not exposed to code-switched input. The simulations predicted how code-switching patterns differ between early balanced and late non-balanced bilinguals; the balanced bilingual simulation code-switches considerably more frequently, which is in line with what has been observed in human speech production. Additionally, we compared the patterns produced by the simulations with two corpora of spontaneous bilingual speech and identified noticeable commonalities and differences. To our knowledge, this is the first computational cognitive model simulating the code-switched production of non-balanced bilinguals and comparing the simulated production of balanced and non-balanced bilinguals with that of human bilinguals.

    Additional information

    dual-path model
  • Tyler, M., & Cutler, A. (2009). Cross-language differences in cue use for speech segmentation. Journal of the Acoustical Society of America, 126, 367-376. doi:10.1121/1.3129127.

    Abstract

    Two artificial-language learning experiments directly compared English, French, and Dutch listeners’ use of suprasegmental cues for continuous-speech segmentation. In both experiments, listeners heard unbroken sequences of consonant-vowel syllables, composed of recurring three- and four-syllable “words.” These words were demarcated by(a) no cue other than transitional probabilities induced by their recurrence, (b) a consistent left-edge cue, or (c) a consistent right-edge cue. Experiment 1 examined a vowel lengthening cue. All three listener groups benefited from this cue in right-edge position; none benefited from it in left-edge position. Experiment 2 examined a pitch-movement cue. English listeners used this cue in left-edge position, French listeners used it in right-edge position, and Dutch listeners used it in both positions. These findings are interpreted as evidence of both language-universal and language-specific effects. Final lengthening is a language-universal effect expressing a more general (non-linguistic) mechanism. Pitch movement expresses prominence which has characteristically different placements across languages: typically at right edges in French, but at left edges in English and Dutch. Finally, stress realization in English versus Dutch encourages greater attention to suprasegmental variation by Dutch than by English listeners, allowing Dutch listeners to benefit from an informative pitch-movement cue even in an uncharacteristic position.
  • Vágvölgyi, R., Bergström, K., Bulajić, A., Klatte, M., Fernandes, T., Grosche, M., Huettig, F., Rüsseler, J., & Lachmann, T. (2021). Functional illiteracy and developmental dyslexia: Looking for common roots. A systematic review. Journal of Cultural Cognitive Science, 5, 159-179. doi:10.1007/s41809-021-00074-9.

    Abstract

    A considerable amount of the population in more economically developed countries are functionally illiterate (i.e., low literate). Despite some years of schooling and basic reading skills, these individuals cannot properly read and write and, as a consequence have problems to understand even short texts. An often-discussed approach (Greenberg et al., 1997) assumes weak phonological processing skills coupled with untreated developmental dyslexia as possible causes of functional illiteracy. Although there is some data suggesting commonalities between low literacy and developmental dyslexia, it is still not clear, whether these reflect shared consequences (i.e., cognitive and behavioral profile) or shared causes. The present systematic review aims at exploring the similarities and differences identified in empirical studies investigating both functional illiterate and developmental dyslexic samples. Nine electronic databases were searched in order to identify all quantitative studies published in English or German. Although a broad search strategy and few limitations were applied, only 5 studies have been identified adequate from the resulting 9269 references. The results point to the lack of studies directly comparing functional illiterate with developmental dyslexic samples. Moreover, a huge variance has been identified between the studies in how they approached the concept of functional illiteracy, particularly when it came to critical categories such the applied definition, terminology, criteria for inclusion in the sample, research focus, and outcome measures. The available data highlight the need for more direct comparisons in order to understand what extent functional illiteracy and dyslexia share common characteristics.

    Additional information

    supplementary materials
  • Van Berkum, J. J. A., Holleman, B., Nieuwland, M. S., Otten, M., & Murre, J. (2009). Right or wrong? The brain's fast response to morally objectionable statements. Psychological Science, 20, 1092 -1099. doi:10.1111/j.1467-9280.2009.02411.x.

    Abstract

    How does the brain respond to statements that clash with a person's value system? We recorded event-related brain potentials while respondents from contrasting political-ethical backgrounds completed an attitude survey on drugs, medical ethics, social conduct, and other issues. Our results show that value-based disagreement is unlocked by language extremely rapidly, within 200 to 250 ms after the first word that indicates a clash with the reader's value system (e.g., "I think euthanasia is an acceptable/unacceptable…"). Furthermore, strong disagreement rapidly influences the ongoing analysis of meaning, which indicates that even very early processes in language comprehension are sensitive to a person's value system. Our results testify to rapid reciprocal links between neural systems for language and for valuation.

    Additional information

    Critical survey statements (in Dutch)
  • Van Bergen, G., & Hogeweg, L. (2021). Managing interpersonal discourse expectations: a comparative analysis of contrastive discourse particles in Dutch. Linguistics, 59(2), 333-360. doi:10.1515/ling-2021-0020.

    Abstract

    In this article we investigate how speakers manage discourse expectations in dialogue by comparing the meaning and use of three Dutch discourse particles, i.e. wel, toch and eigenlijk, which all express a contrast between their host utterance and a discourse-based expectation. The core meanings of toch, wel and eigenlijk are formally distinguished on the basis of two intersubjective parameters: (i) whether the particle marks alignment or misalignment between speaker and addressee discourse beliefs, and (ii) whether the particle requires an assessment of the addressee’s representation of mutual discourse beliefs. By means of a quantitative corpus study, we investigate to what extent the intersubjective meaning distinctions between wel, toch and eigenlijk are reflected in statistical usage patterns across different social situations. Results suggest that wel, toch and eigenlijk are lexicalizations of distinct generalized politeness strategies when expressing contrast in social interaction. Our findings call for an interdisciplinary approach to discourse particles in order to enhance our understanding of their functions in language.
  • Van Heukelum, S., Tulva, K., Geers, F. E., van Dulm, S., Ruisch, I. H., Mill, J., Viana, J. F., Beckmann, C. F., Buitelaar, J. K., Poelmans, G., Glennon, J. C., Vogt, B. A., Havenith, M. N., & França, A. S. (2021). A central role for anterior cingulate cortex in the control of pathological aggression. Current Biology, 31, 2321-2333.e5. doi:10.1016/j.cub.2021.03.062.

    Abstract

    Controlling aggression is a crucial skill in social species like rodents and humans and has been associated with anterior cingulate cortex (ACC). Here, we directly link the failed regulation of aggression in BALB/cJ mice to ACC hypofunction. We first show that ACC in BALB/cJ mice is structurally degraded: neuron density is decreased, with pervasive neuron death and reactive astroglia. Gene-set enrichment analysis suggested that this process is driven by neuronal degeneration, which then triggers toxic astrogliosis. cFos expression across ACC indicated functional consequences: during aggressive encounters, ACC was engaged in control mice, but not BALB/cJ mice. Chemogenetically activating ACC during aggressive encounters drastically suppressed pathological aggression but left species-typical aggression intact. The network effects of our chemogenetic perturbation suggest that this behavioral rescue is mediated by suppression of amygdala and hypothalamus and activation of mediodorsal thalamus. Together, these findings highlight the central role of ACC in curbing pathological aggression.
  • Ip, H. F., Van der Laan, C. M., Krapohl, E. M. L., Brikell, I., Sánchez-Mora, C., Nolte, I. M., St Pourcain, B., Bolhuis, K., Palviainen, T., Zafarmand, H., Colodro-Conde, L., Gordon, S., Zayats, T., Aliev, F., Jiang, C., Wang, C. A., Saunders, G., Karhunen, V., Hammerschlag, A. R., Adkins, D. E. and 129 moreIp, H. F., Van der Laan, C. M., Krapohl, E. M. L., Brikell, I., Sánchez-Mora, C., Nolte, I. M., St Pourcain, B., Bolhuis, K., Palviainen, T., Zafarmand, H., Colodro-Conde, L., Gordon, S., Zayats, T., Aliev, F., Jiang, C., Wang, C. A., Saunders, G., Karhunen, V., Hammerschlag, A. R., Adkins, D. E., Border, R., Peterson, R. E., Prinz, J. A., Thiering, E., Seppälä, I., Vilor-Tejedor, N., Ahluwalia, T. S., Day, F. R., Hottenga, J.-J., Allegrini, A. G., Rimfeld, K., Chen, Q., Lu, Y., Martin, J., Soler Artigas, M., Rovira, P., Bosch, R., Español, G., Ramos Quiroga, J. A., Neumann, A., Ensink, J., Grasby, K., Morosoli, J. J., Tong, X., Marrington, S., Middeldorp, C., Scott, J. G., Vinkhuyzen, A., Shabalin, A. A., Corley, R., Evans, L. M., Sugden, K., Alemany, S., Sass, L., Vinding, R., Ruth, K., Tyrrell, J., Davies, G. E., Ehli, E. A., Hagenbeek, F. A., De Zeeuw, E., Van Beijsterveldt, T. C., Larsson, H., Snieder, H., Verhulst, F. C., Amin, N., Whipp, A. M., Korhonen, T., Vuoksimaa, E., Rose, R. J., Uitterlinden, A. G., Heath, A. C., Madden, P., Haavik, J., Harris, J. R., Helgeland, Ø., Johansson, S., Knudsen, G. P. S., Njolstad, P. R., Lu, Q., Rodriguez, A., Henders, A. K., Mamun, A., Najman, J. M., Brown, S., Hopfer, C., Krauter, K., Reynolds, C., Smolen, A., Stallings, M., Wadsworth, S., Wall, T. L., Silberg, J. L., Miller, A., Keltikangas-Järvinen, L., Hakulinen, C., Pulkki-Råback, L., Havdahl, A., Magnus, P., Raitakari, O. T., Perry, J. R. B., Llop, S., Lopez-Espinosa, M.-J., Bønnelykke, K., Bisgaard, H., Sunyer, J., Lehtimäki, T., Arseneault, L., Standl, M., Heinrich, J., Boden, J., Pearson, J., Horwood, L. J., Kennedy, M., Poulton, R., Eaves, L. J., Maes, H. H., Hewitt, J., Copeland, W. E., Costello, E. J., Williams, G. M., Wray, N., Järvelin, M.-R., McGue, M., Iacono, W., Caspi, A., Moffitt, T. E., Whitehouse, A., Pennell, C. E., Klump, K. L., Burt, S. A., Dick, D. M., Reichborn-Kjennerud, T., Martin, N. G., Medland, S. E., Vrijkotte, T., Kaprio, J., Tiemeier, H., Davey Smith, G., Hartman, C. A., Oldehinkel, A. J., Casas, M., Ribasés, M., Lichtenstein, P., Lundström, S., Plomin, R., Bartels, M., Nivard, M. G., & Boomsma, D. I. (2021). Genetic association study of childhood aggression across raters, instruments, and age. Translational Psychiatry, 11: 413. doi:10.1038/s41398-021-01480-x.
  • van der Burght, C. L., Friederici, A. D., Goucha, T., & Hartwigsen, G. (2021). Pitch accents create dissociable syntactic and semantic expectations during sentence processing. Cognition, 212: 104702. doi:10.1016/j.cognition.2021.104702.

    Abstract

    The language system uses syntactic, semantic, as well as prosodic cues to efficiently guide auditory sentence comprehension. Prosodic cues, such as pitch accents, can build expectations about upcoming sentence elements. This study investigates to what extent syntactic and semantic expectations generated by pitch accents can be dissociated and if so, which cues take precedence when contradictory information is present. We used sentences in which one out of two nominal constituents was placed in contrastive focus with a third one. All noun phrases carried overt syntactic information (case-marking of the determiner) and semantic information (typicality of the thematic role of the noun). Two experiments (a sentence comprehension and a sentence completion task) show that focus, marked by pitch accents, established expectations in both syntactic and semantic domains. However, only the syntactic expectations, when violated, were strong enough to interfere with sentence comprehension. Furthermore, when contradictory cues occurred in the same sentence, the local syntactic cue (case-marking) took precedence over the semantic cue (thematic role), and overwrote previous information cued by prosody. The findings indicate that during auditory sentence comprehension the processing system integrates different sources of information for argument role assignment, yet primarily relies on syntactic information.
  • Van Paridon, J., Ostarek, M., Arunkumar, M., & Huettig, F. (2021). Does neuronal recycling result in destructive competition? The influence of learning to read on the recognition of faces. Psychological Science, 32, 459-465. doi:10.1177/0956797620971652.

    Abstract

    Written language, a human cultural invention, is far too recent for dedicated neural
    infrastructure to have evolved in its service. Culturally newly acquired skills (e.g. reading) thus ‘recycle’ evolutionarily older circuits that originally evolved for different, but similar functions (e.g. visual object recognition). The destructive competition hypothesis predicts that this neuronal recycling has detrimental behavioral effects on the cognitive functions a cortical network originally evolved for. In a study with 97 literate, low-literate, and illiterate participants from the same socioeconomic background we find that even after adjusting for cognitive ability and test-taking familiarity, learning to read is associated with an increase, rather than a decrease, in object recognition abilities. These results are incompatible with the claim that neuronal recycling results in destructive competition and consistent with the possibility that learning to read instead fine-tunes general object recognition mechanisms, a hypothesis that needs further neuroscientific investigation.

    Additional information

    supplemental material
  • Van Leeuwen, T. M., Wilsson, L., Norrman, H. N., Dingemanse, M., Bölte, S., & Neufeld, J. (2021). Perceptual processing links autism and synesthesia: A co-twin control study. Cortex, 145, 236-249. doi:10.1016/j.cortex.2021.09.016.
  • Van Turennout, M., Hagoort, P., & Brown, C. M. (1997). Electrophysiological evidence on the time course of semantic and phonological processes in speech production. Journal of Experimental Psychology: Learning, Memory, and Cognition, 23(4), 787-806.

    Abstract

    The temporal properties of semantic and phonological processes in speech production were investigated in a new experimental paradigm using movement-related brain potentials. The main experimental task was picture naming. In addition, a 2-choice reaction go/no-go procedure was included, involving a semantic and a phonological categorization of the picture name. Lateralized readiness potentials (LRPs) were derived to test whether semantic and phonological information activated motor processes at separate moments in time. An LRP was only observed on no-go trials when the semantic (not the phonological) decision determined the response hand. Varying the position of the critical phoneme in the picture name did not affect the onset of the LRP but rather influenced when the LRP began to differ on go and no-go trials and allowed the duration of phonological encoding of a word to be estimated. These results provide electrophysiological evidence for early semantic activation and later phonological encoding.
  • Van de Geer, J. P., & Levelt, W. J. M. (1963). Detection of visual patterns disturbed by noise: An exploratory study. Quarterly Journal of Experimental Psychology, 15, 192-204. doi:10.1080/17470216308416324.

    Abstract

    An introductory study of the perception of stochastically specified events is reported. The initial problem was to determine whether the perceiver can split visual input data of this kind into random and determined components. The inability of subjects to do so with the stimulus material used (a filmlike sequence of dot patterns), led to the more general question of how subjects code this kind of visual material. To meet the difficulty of defining the subjects' responses, two experiments were designed. In both, patterns were presented as a rapid sequence of dots on a screen. The patterns were more or less disturbed by “noise,” i.e. the dots did not appear exactly at their proper places. In the first experiment the response was a rating on a semantic scale, in the second an identification from among a set of alternative patterns. The results of these experiments give some insight in the coding systems adopted by the subjects. First, noise appears to be detrimental to pattern recognition, especially to patterns with little spread. Second, this shows connections with the factors obtained from analysis of the semantic ratings, e.g. easily disturbed patterns show a large drop in the semantic regularity factor, when only a little noise is added.
  • Van Berkum, J. J. A., Koornneef, A. W., Otten, M., & Nieuwland, M. S. (2007). Establishing reference in language comprehension: An electrophysiological perspective. Brain Research, 1146, 158-171. doi:10.1016/j.brainres.2006.06.091.

    Abstract

    The electrophysiology of language comprehension has long been dominated by research on syntactic and semantic integration. However, to understand expressions like "he did it" or "the little girl", combining word meanings in accordance with semantic and syntactic constraints is not enough--readers and listeners also need to work out what or who is being referred to. We review our event-related brain potential research on the processes involved in establishing reference, and present a new experiment in which we examine when and how the implicit causality associated with specific interpersonal verbs affects the interpretation of a referentially ambiguous pronoun. The evidence suggests that upon encountering a singular noun or pronoun, readers and listeners immediately inspect their situation model for a suitable discourse entity, such that they can discriminate between having too many, too few, or exactly the right number of referents within at most half a second. Furthermore, our implicit causality findings indicate that a fragment like "David praised Linda because..." can immediately foreground a particular referent, to the extent that a subsequent "he" is at least initially construed as a syntactic error. In all, our brain potential findings suggest that referential processing is highly incremental, and not necessarily contingent upon the syntax. In addition, they demonstrate that we can use ERPs to relatively selectively keep track of how readers and listeners establish reference.
  • Van Wingen, G., Van Broekhoven, F., Verkes, R. J., Petersson, K. M., Bäckström, T., Buitelaar, J., & Fernández, G. (2007). How progesterone impairs memory for biologically salient stimuli in healthy young women. Journal of Neuroscience, 27(42), 11416-11423. doi:10.1523/JNEUROSCI.1715-07.2007.

    Abstract

    Progesterone, or rather its neuroactive metabolite allopregnanolone, modulates amygdala activity and thereby influences anxiety. Cognition and, in particular, memory are also altered by allopregnanolone. In the present study, we investigated whether allopregnanolone modulates memory for biologically salient stimuli by influencing amygdala activity, which in turn may affect neural processes in other brain regions. A single progesterone dose was administered orally to healthy young women in a double-blind, placebo-controlled, crossover design, and participants were asked to memorize and recognize faces while undergoing functional magnetic resonance imaging. Progesterone decreased recognition accuracy without affecting reaction times. The imaging results show that the amygdala, hippocampus, and fusiform gyrus supported memory formation. Importantly, progesterone decreased responses to faces in the amygdala and fusiform gyrus duringmemoryencoding, whereas it increased hippocampal responses. The progesterone-induced decrease in neural activity in the amygdala and fusiform gyrus predicted the decrease in memory performance across subjects. However, progesterone did not modulate the differential activation between subsequently remembered and subsequently forgotten faces in these areas. A similar pattern of results was observed in the fusiform gyrus and prefrontal cortex during memory retrieval. These results suggest that allopregnanolone impairs memory by reducing the recruitment of those brain regions that support memory formation and retrieval. Given the important role of the amygdala in the modulation of memory, these results suggest that allopregnanolone alters memory by influencing amygdala activity, which in turn may affect memory processes in other brain regions
  • Van de Geer, J. P., Levelt, W. J. M., & Plomp, R. (1962). The connotation of musical consonance. Acta Psychologica, 20, 308-319.

    Abstract

    As a preliminary to further research on musical consonance an explanatory investigation was made on the different modes of judgment of musical intervals. This was done by way of a semantic differential. Subjects rated 23 intervals against 10 scales. In a factor analysis three factors appeared: pitch, evaluation and fusion. The relation between these factors and some physical characteristics has been investigated. The scale consonant-dissonant showed to be purely evaluative (in opposition to Stumpf's theory). This evaluative connotation is not in accordance with the musicological meaning of consonance. Suggestions to account for this difference have been given.
  • Van Valin Jr., R. D. (2007). Some thoughts on the reason for the lesser status of typology in the USA as opposed to Europe. Linguistic Typology, 11(1), 253-257. doi:10.1515/LINGTY.2007.019.

    Abstract

    This article addresses the issue of the different status that typology has in American linguistics as opposed to European linguistics. The historical roots of the difference lie in both structural and generative linguistics, in the contrasts between post-Bloomfieldian structuralism in the US vs. Praguean structuralism in Europe, and in the extent of the influence of generative grammar on the two continents.
  • Van Tiel, B., Deliens, G., Geelhand, P., Murillo Oosterwijk, A., & Kissine, M. (2021). Strategic deception in adults with autism spectrum disorder. Journal of Autism and Developmental Disorders, 51, 255-266. doi:10.1007/s10803-020-04525-0.

    Abstract

    Autism Spectrum Disorder (ASD) is often associated with impaired perspective-taking skills. Deception is an important indicator of perspective-taking, and therefore may be thought to pose difficulties to people with ASD (e.g., Baron-Cohen in J Child Psychol Psychiatry 3:1141–1155, 1992). To test this hypothesis, we asked participants with and without ASD to play a computerised deception game. We found that participants with ASD were equally likely—and in complex cases of deception even more likely—to deceive and detect deception, and learned deception at a faster rate. However, participants with ASD initially deceived less frequently, and were slower at detecting deception. These results suggest that people with ASD readily engage in deception but may do so through conscious and effortful reasoning about other people’s perspectiv
  • Van Paridon, J., & Thompson, B. (2021). subs2vec: Word embeddings from subtitles in 55 languages. Behavior Research Methods, 53(2), 629-655. doi:10.3758/s13428-020-01406-3.

    Abstract

    This paper introduces a novel collection of word embeddings, numerical representations of lexical semantics, in 55 languages, trained on a large corpus of pseudo-conversational speech transcriptions from television shows and movies. The embeddings were trained on the OpenSubtitles corpus using the fastText implementation of the skipgram algorithm. Performance comparable with (and in some cases exceeding) embeddings trained on non-conversational (Wikipedia) text is reported on standard benchmark evaluation datasets. A novel evaluation method of particular relevance to psycholinguists is also introduced: prediction of experimental lexical norms in multiple languages. The models, as well as code for reproducing the models and all analyses reported in this paper (implemented as a user-friendly Python package), are freely available at: https://github.com/jvparidon/subs2vec.

    Additional information

    https://github.com/jvparidon/subs2vec
  • Van Berkum, J. J. A. (1997). Syntactic processes in speech production: The retrieval of grammatical gender. Cognition, 64(2), 115-152. doi:10.1016/S0010-0277(97)00026-7.

    Abstract

    Jescheniak and Levelt (Jescheniak, J.-D., Levelt, W.J.M. 1994. Journal of Experimental Psychology: Learning, Memory and Cognition 20 (4), 824–843) have suggested that the speed with which native speakers of a gender-marking language retrieve the grammatical gender of a noun from their mental lexicon may depend on the recency of earlier access to that same noun's gender, as the result of a mechanism that is dedicated to facilitate gender-marked anaphoric reference to recently introduced discourse entities. This hypothesis was tested in two picture naming experiments. Recent gender access did not facilitate the production of gender-marked adjective noun phrases (Experiment 1), nor that of gender-marked definite article noun phrases (Experiment 2), even though naming times for the latter utterances were sensitive to the gender of a written distractor word superimposed on the picture to be named. This last result replicates and extends earlier gender-specific picture-word interference results (Schriefers, H. 1993. Journal of Experimental Psychology: Learning, Memory, and Cognition 19 (4), 841–850), showing that one can selectively tap into the production of grammatical gender agreement during speaking. The findings are relevant to theories of speech production and the representation of grammatical gender for that process.
  • Van Gijn, R. (2009). The phonology of mixed languages. Journal of Pidgin and Creole Languages, 24(1), 91-117. doi:10.1075/jpcl.24.1.04gij.

    Abstract

    Mixed languages are said to be the result of a process of intertwining (e.g. Bakker & Muysken 1995, Bakker 1997), a regular process in which the grammar of one language is combined with the lexicon of another. However, the outcome of this process differs from language pair to language pair. As far as morphosyntax is concerned, people have discussed these different outcomes and the reasons for them extensively, e.g. Bakker 1997 for Michif, Mous 2003 for Ma’a, Muysken 1997a for Media Lengua and 1997b for Callahuaya. The issue of phonology, however, has not generated a large debate. This paper compares the phonological systems of the mixed languages Media Lengua, Callahuaya, Mednyj Aleut, and Michif. It will be argued that the outcome of the process of intertwining, as far as phonology is concerned, is at least partly determined by the extent to which unmixed phonological domains exist.
  • Van Berkum, J. J. A., Hagoort, P., & Brown, C. M. (2000). The use of referential context and grammatical gender in parsing: A reply to Brysbaert and Mitchell. Journal of Psycholinguistic Research, 29(5), 467-481. doi:10.1023/A:1005168025226.

    Abstract

    Based on the results of an event-related brain potentials (ERP) experiment (van Berkum, Brown, & Hagoort. 1999a, b), we have recently argued that discourse-level referential context can be taken into account extremely rapidly by the parser. Moreover, our ERP results indicated that local grammatical gender information, although available within a few hundred milliseconds from word onset, is not always used quickly enough to prevent the parser from considering a discourse-supported, but agreement-violating, syntactic analysis. In a comment on our work, Brysbaert and Mitchell (2000) have raised concerns about the methodology of our ERP experiment and have challenged our interpretation of the results. In this reply, we argue that these concerns are unwarranted and, that, in contrast to our own interpretation, the alternative explanations provided by Brysbaert and Mitchell do not account for the full pattern of ERP results.
  • Van Uytvanck, D., Dukers, A., Ringersma, J., & Wittenburg, P. (2007). Using Google Earth to access language resources. Language Archive Newsletter, (9), 4-7.

    Abstract

    Over the past ten years Geographic Information Systems (GIS) have evolved from a highly specialised niche technology to one that is used daily by a wide range of people. This article describes geographic browsing of language archives, which provides intuitive exploration of resources and permits integration and correlation of information from different archives, even across different research disciplines. In order to facilitate both exploration and management of resources, digital language archives are organised according to criteria such as language name, research topic, project information, researchers, countries, or genres. A set of such criteria can form a tree-like classification scheme, such as in the MPI-IMDI archive, which in turn forms the main method of searching and querying the archive resources. Searching for information can be difficult for occasional users because effective use of these search-fields typically requires specialised knowledge. We assume that many non-specialist users of language resources will search by language name, language family, or geographic area, so that geographic navigation would offer a very powerful search method. We also assume that such users are familiar with maps, and that geographic browsing is more intuitive than browsing classification trees, so these users would prefer to start with a large scale map and then zoom in to find the data that interests them. Therefore, classification trees and geographic maps provide complementary methods for accessing language resources to meet the needs of different user groups. We selected Google Earth (GE) as a geographic browsing system and overlaid it with linguistic information. GE was chosen because it is available via the web, it has good navigation controls, it is familiar to many web users, and because the overlaid linguistic information can be formulated in XML, making it comparatively easy to interchange with other geographic systems.
  • Varola*, M., Verga*, L., Sroka, M., Villanueva, S., Charrier, I., & Ravignani, A. (2021). Can harbor seals (Phoca vitulina) discriminate familiar conspecific calls after long periods of separation? PeerJ, 9: e12431. doi:10.7717/peerj.12431.

    Abstract

    * - indicates joint first authorship -
    The ability to discriminate between familiar and unfamiliar calls may play a key role in pinnipeds’ communication and survival, as in the case of mother-pup interactions. Vocal discrimination abilities have been suggested to be more developed in pinniped species with the highest selective pressure such as the otariids; yet, in some group-living phocids, such as harbor seals (Phoca vitulina), mothers are also able to recognize their pup’s voice. Conspecifics’ vocal recognition in pups has never been investigated; however, the repeated interaction occurring between pups within the breeding season suggests that long-term vocal discrimination may occur. Here we explored this hypothesis by presenting three rehabilitated seal pups with playbacks of vocalizations from unfamiliar or familiar pups. It is uncommon for seals to come into rehabilitation for a second time in their lifespan, and this study took advantage of these rare cases. A simple visual inspection of the data plots seemed to show more reactions, and of longer duration, in response to familiar as compared to unfamiliar playbacks in two out of three pups. However, statistical analyses revealed no significant difference between the experimental conditions. We also found no significant asymmetry in orientation (left vs. right) towards familiar and unfamiliar sounds. While statistics do not support the hypothesis of an established ability to discriminate familiar vocalizations from unfamiliar ones in harbor seal pups, further investigations with a larger sample size are needed to confirm or refute this hypothesis.

    Additional information

    dataset
  • Vartiainen, J., Aggujaro, S., Lehtonen, M., Hulten, A., Laine, M., & Salmelin, R. (2009). Neural dynamics of reading morphologically complex words. NeuroImage, 47, 2064-2072. doi:10.1016/j.neuroimage.2009.06.002.

    Abstract

    Despite considerable research interest, it is still an open issue as to how morphologically complex words such as “car+s” are represented and processed in the brain. We studied the neural correlates of the processing of inflected nouns in the morphologically rich Finnish language. Previous behavioral studies in Finnish have yielded a robust inflectional processing cost, i.e., inflected words are harder to recognize than otherwise matched morphologically simple words. Theoretically this effect could stem either from decomposition of inflected words into a stem and a suffix at input level and/or from subsequent recombination at the semantic–syntactic level to arrive at an interpretation of the word. To shed light on this issue, we used magnetoencephalography to reveal the time course and localization of neural effects of morphological structure and frequency of written words. Ten subjects silently read high- and low-frequency Finnish words in inflected and monomorphemic form. Morphological complexity was accompanied by stronger and longerlasting activation of the left superior temporal cortex from 200 ms onwards. Earlier effects of morphology were not found, supporting the view that the well-established behavioral processing cost for inflected words stems from the semantic–syntactic level rather than from early decomposition. Since the effect of morphology was detected throughout the range of word frequencies employed, the majority of inflected Finnish words appears to be represented in decomposed form and only very high-frequency inflected words may acquire full-form representations.
  • Vega-Mendoza, M., Pickering, M. J., & Nieuwland, M. S. (2021). Concurrent use of animacy and event-knowledge during comprehension: Evidence from event-related potentials. Neuropsychologia, 152: 107724. doi:10.1016/j.neuropsychologia.2020.107724.

    Abstract

    In two ERP experiments, we investigated whether readers prioritize animacy over real-world event-knowledge during sentence comprehension. We used the paradigm of Paczynski and Kuperberg (2012), who argued that animacy is prioritized based on the observations that the ‘related anomaly effect’ (reduced N400s for context-related anomalous words compared to unrelated words) does not occur for animacy violations, and that animacy violations but not relatedness violations elicit P600 effects. Participants read passive sentences with plausible agents (e.g., The prescription for the mental disorder was written by the psychiatrist) or implausible agents that varied in animacy and semantic relatedness (schizophrenic/guard/pill/fence). In Experiment 1 (with a plausibility judgment task), plausible sentences elicited smaller N400s relative to all types of implausible sentences. Crucially, animate words elicited smaller N400s than inanimate words, and related words elicited smaller N400s than unrelated words, but Bayesian analysis revealed substantial evidence against an interaction between animacy and relatedness. Moreover, at the P600 time-window, we observed more positive ERPs for animate than inanimate words and for related than unrelated words at anterior regions. In Experiment 2 (without judgment task), we observed an N400 effect with animacy violations, but no other effects. Taken together, the results of our experiments fail to support a prioritized role of animacy information over real-world event-knowledge, but they support an interactive, constraint-based view on incremental semantic processing.
  • Verdonschot, R. G., Han, J.-I., & Kinoshita, S. (2021). The proximate unit in Korean speech production: Phoneme or syllable? Quarterly Journal of Experimental Psychology, 74, 187-198. doi:10.1177/1747021820950239.

    Abstract

    We investigated the “proximate unit” in Korean, that is, the initial phonological unit selected in speech production by Korean speakers. Previous studies have shown mixed evidence indicating either a phoneme-sized or a syllable-sized unit. We conducted two experiments in which participants named pictures while ignoring superimposed non-words. In English, for this task, when the picture (e.g., dog) and distractor phonology (e.g., dark) initially overlap, typically the picture target is named faster. We used a range of conditions (in Korean) varying from onset overlap to syllabic overlap, and the results indicated an important role for the syllable, but not the phoneme. We suggest that the basic unit used in phonological encoding in Korean is different from Germanic languages such as English and Dutch and also from Japanese and possibly also Chinese. Models dealing with the architecture of language production can use these results when providing a framework suitable for all languages in the world, including Korean.
  • Verga, L., & Ravignani, A. (2021). Strange seal sounds: Claps, slaps, and multimodal pinniped rhythms. Frontiers in Ecology and Evolution, 9: 644497. doi:10.3389/fevo.2021.644497.
  • Verga, L., Schwartze, M., Stapert, S., Winkens, I., & Kotz, S. A. (2021). Dysfunctional timing in traumatic brain injury patients: Co-occurrence of cognitive, motor, and perceptual deficits. Frontiers in Psychology, 12: 731898. doi:10.3389/fpsyg.2021.731898.

    Abstract

    Timing is an essential part of human cognition and of everyday life activities, such as walking or holding a conversation. Previous studies showed that traumatic brain injury (TBI) often affects cognitive functions such as processing speed and time-sensitive abilities, causing long-term sequelae as well as daily impairments. However, the existing evidence on timing capacities in TBI is mostly limited to perception and the processing of isolated intervals. It is therefore open whether the observed deficits extend to motor timing and to continuous dynamic tasks that more closely match daily life activities. The current study set out to answer these questions by assessing audio motor timing abilities and their relationship with cognitive functioning in a group of TBI patients (n=15) and healthy matched controls. We employed a comprehensive set of tasks aiming at testing timing abilities across perception and production and from single intervals to continuous auditory sequences. In line with previous research, we report functional impairments in TBI patients concerning cognitive processing speed and perceptual timing. Critically, these deficits extended to motor timing: The ability to adjust to tempo changes in an auditory pacing sequence was impaired in TBI patients, and this motor timing deficit covaried with measures of processing speed. These findings confirm previous evidence on perceptual and cognitive timing deficits resulting from TBI and provide first evidence for comparable deficits in motor behavior. This suggests basic co-occurring perceptual and motor timing impairments that may factor into a wide range of daily activities. Our results thus place TBI into the wider range of pathologies with well-documented timing deficits (such as Parkinson’s disease) and encourage the search for novel timing-based therapeutic interventions (e.g., employing dynamic and/or musical stimuli) with high transfer potential to everyday life activities.

    Additional information

    supplementary material
  • Verhagen, J., & Schimke, S. (2009). Differences or fundamental differences? Zeitschrift für Sprachwissenschaft, 28(1), 97-106. doi:10.1515/ZFSW.2009.011.
  • Verhagen, J. (2009). Temporal adverbials, negation and finiteness in Dutch as a second language: A scope-based account. IRAL, 47(2), 209-237. doi:10.1515/iral.2009.009.

    Abstract

    This study investigates the acquisition of post-verbal (temporal) adverbials and post-verbal negation in L2 Dutch. It is based on previous findings for L2 French that post-verbal negation poses less of a problem for L2 learners than post-verbal adverbial placement (Hawkins, Towell, Bazergui, Second Language Research 9: 189-233, 1993; Herschensohn, Minimally raising the verb issue: 325-336, Cascadilla Press, 1998). The current data show that, at first sight, Moroccan and Turkish learners of Dutch also have fewer problems with post-verbal negation than with post-verbal adverbials. However, when a distinction is made between different types of adverbials, it seems that this holds for adverbials of position such as 'today' but not for adverbials of contrast such as 'again'. To account for this difference, it is argued that different types of adverbial occupy different positions in the L2 data for reasons of scope marking. Moreover, the placement of adverbials such as 'again' interacts with the acquisition of finiteness marking (resulting in post-verbal placement), while there is no such interaction between adverbials such as 'today' and finiteness marking.
  • 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.

    Additional information

    supplementary information
  • 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., 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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  • 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. (Eds.). (2021). Vocal learning in animals and humans [Special Issue]. Philosophical Transactions of the Royal Society of London, Series B: Biological Sciences, 376.

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