Publications

Displaying 601 - 661 of 661
  • Van Bergen, G., & Bosker, H. R. (2018). Linguistic expectation management in online discourse processing: An investigation of Dutch inderdaad 'indeed' and eigenlijk 'actually'. Journal of Memory and Language, 103, 191-209. doi:10.1016/j.jml.2018.08.004.

    Abstract

    Interpersonal discourse particles (DPs), such as Dutch inderdaad (≈‘indeed’) and eigenlijk (≈‘actually’) are highly frequent in everyday conversational interaction. Despite extensive theoretical descriptions of their polyfunctionality, little is known about how they are used by language comprehenders. In two visual world eye-tracking experiments involving an online dialogue completion task, we asked to what extent inderdaad, confirming an inferred expectation, and eigenlijk, contrasting with an inferred expectation, influence real-time understanding of dialogues. Answers in the dialogues contained a DP or a control adverb, and a critical discourse referent was replaced by a beep; participants chose the most likely dialogue completion by clicking on one of four referents in a display. Results show that listeners make rapid and fine-grained situation-specific inferences about the use of DPs, modulating their expectations about how the dialogue will unfold. Findings further specify and constrain theories about the conversation-managing function and polyfunctionality of DPs.
  • Van Leeuwen, E. J. C., Cronin, K. A., Haun, D. B. M., Mundry, R., & Bodamer, M. D. (2012). Neighbouring chimpanzee communities show different preferences in social grooming behaviour. Proceedings of the Royal Society B: Biological Sciences, 279, 4362-4367. doi:10.1098/rspb.2012.1543.

    Abstract

    Grooming handclasp (GHC) behaviour was originally advocated as the first evidence of social culture in chimpanzees owing to the finding that some populations engaged in the behaviour and others do not. To date, however, the validity of this claim and the extent to which this social behaviour varies between groups is unclear. Here, we measured (i) variation, (ii) durability and (iii) expansion of the GHC behaviour in four chimpanzee communities that do not systematically differ in their genetic backgrounds and live in similar ecological environments. Ninety chimpanzees were studied for a total of 1029 h; 1394 GHC bouts were observed between 2010 and 2012. Critically, GHC style (defined by points of bodily contact) could be systematically linked to the chimpanzee’s group identity, showed temporal consistency both withinand between-groups, and could not be accounted for by the arm-length differential between partners. GHC has been part of the behavioural repertoire of the chimpanzees under study for more than 9 years (surpassing durability criterion) and spread across generations (surpassing expansion criterion). These results strongly indicate that chimpanzees’ social behaviour is not only motivated by innate predispositions and individual inclinations, but may also be partly cultural in nature.
  • Van Campen, A. D., Kunert, R., Van den Wildenberg, W. P. M., & Ridderinkhof, K. R. (2018). Repetitive transcranial magnetic stimulation over inferior frontal cortex impairs the suppression (but not expression) of action impulses during action conflict. Psychophysiology, 55(3): e13003. doi:10.1111/psyp.13003.

    Abstract

    In the recent literature, the effects of noninvasive neurostimulation on cognitive functioning appear to lack consistency and replicability. We propose that such effects may be concealed unless dedicated, sensitive, and process-specific dependent measures are used. The expression and subsequent suppression of response capture are often studied using conflict tasks. Response-time distribution analyses have been argued to provide specific measures of the susceptibility to make fast impulsive response errors, as well as the proficiency of the selective suppression of these impulses. These measures of response capture and response inhibition are particularly sensitive to experimental manipulations and clinical deficiencies that are typically obfuscated in commonly used overall performance analyses. Recent work using structural and functional imaging techniques links these behavioral outcome measures to the integrity of frontostriatal networks. These studies suggest that the presupplementary motor area (pre-SMA) is linked to the susceptibility to response capture whereas the right inferior frontal cortex (rIFC) is associated with the selective suppression of action impulses. Here, we used repetitive transcranial magnetic stimulation (rTMS) to test the causal involvement of these two cortical areas in response capture and inhibition in the Simon task. Disruption of rIFC function specifically impaired selective suppression of conflicting action tendencies, whereas the anticipated increase of fast impulsive errors after perturbing pre-SMA function was not confirmed. These results provide a proof of principle of the notion that the selection of appropriate dependent measures is perhaps crucial to establish the effects of neurostimulation on specific cognitive functions.
  • Van Leeuwen, E. J. C., Cohen, E., Collier-Baker, E., Rapold, C. J., Schäfer, M., Schütte, S., & Haun, D. B. M. (2018). The development of human social learning across seven societies. Nature Communications, 9: 2076. doi:10.1038/s41467-018-04468-2.

    Abstract

    Social information use is a pivotal characteristic of the human species. Avoiding the cost of individual exploration, social learning confers substantial fitness benefits under a wide variety of environmental conditions, especially when the process is governed by biases toward relative superiority (e.g., experts, the majority). Here, we examine the development of social information use in children aged 4–14 years (n = 605) across seven societies in a standardised social learning task. We measured two key aspects of social information use: general reliance on social information and majority preference. We show that the extent to which children rely on social information depends on children’s cultural background. The extent of children’s majority preference also varies cross-culturally, but in contrast to social information use, the ontogeny of majority preference follows a U-shaped trajectory across all societies. Our results demonstrate both cultural continuity and diversity in the realm of human social learning.

    Additional information

    VanLeeuwen_etal_2018sup.pdf
  • 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 Alphen, P. M., & Van Berkum, J. J. A. (2012). Semantic involvement of initial and final lexical embeddings during sense-making: The advantage of starting late. Frontiers in Psychology, 3, 190. doi:10.3389/fpsyg.2012.00190.

    Abstract

    During spoken language interpretation, listeners rapidly relate the meaning of each individual word to what has been said before. However, spoken words often contain spurious other words, like 'day' in 'daisy', or 'dean' in 'sardine'. Do listeners also relate the meaning of such unintended, spurious words to the prior context? We used ERPs to look for transient meaning-based N400 effects in sentences that were completely plausible at the level of words intended by the speaker, but contained an embedded word whose meaning clashed with the context. Although carrier words with an initial embedding ('day' in 'daisy') did not elicit an embedding-related N400 effect relative to matched control words without embedding, carrier words with a final embedding ('dean' in 'sardine') did elicit such an effect. Together with prior work from our lab and the results of a Shortlist B simulation, our findings suggest that listeners do semantically interpret embedded words, albeit not under all conditions. We explain the latter by assuming that the sense-making system adjusts its hypothesis for how to interpret the external input at every new syllable, in line with recent ideas of active sampling in perception.
  • Van Donkelaar, M. M. J., Hoogman, M., Pappa, I., Tiemeier, H., Buitelaar, J. K., Franke, B., & Bralten, J. (2018). Pleiotropic Contribution of MECOM and AVPR1A to Aggression and Subcortical Brain Volumes. Frontiers in Behavioral Neuroscience, 12: 61. doi:10.3389/fnbeh.2018.00061.

    Abstract

    Reactive and proactive subtypes of aggression have been recognized to help parse etiological heterogeneity of this complex phenotype. With a heritability of about 50%, genetic factors play a role in the development of aggressive behavior. Imaging studies implicate brain structures related to social behavior in aggression etiology, most notably the amygdala and striatum. This study aimed to gain more insight into the pathways from genetic risk factors for aggression to aggression phenotypes. To this end, we conducted genome-wide gene-based cross-trait meta-analyses of aggression with the volumes of amygdala, nucleus accumbens and caudate nucleus to identify genes influencing both aggression and aggression-related brain volumes. We used data of large-scale genome-wide association studies (GWAS) of: (a) aggressive behavior in children and adolescents (EAGLE, N = 18,988); and (b) Magnetic Resonance Imaging (MRI)-based volume measures of aggression-relevant subcortical brain regions (ENIGMA2, N = 13,171). Second, the identified genes were further investigated in a sample of healthy adults (mean age (SD) = 25.28 (4.62) years; 43% male) who had genome-wide genotyping data and questionnaire data on aggression subtypes available (Brain Imaging Genetics, BIG, N = 501) to study their effect on reactive and proactive subtypes of aggression. Our meta-analysis identified two genes, MECOM and AVPR1A, significantly associated with both aggression risk and nucleus accumbens (MECOM) and amygdala (AVPR1A) brain volume. Subsequent in-depth analysis of these genes in healthy adults (BIG), including sex as an interaction term in the model, revealed no significant subtype-specific gene-wide associations. Using cross-trait meta-analysis of brain measures and psychiatric phenotypes, this study generated new hypotheses about specific links between genes, the brain and behavior. Results indicate that MECOM and AVPR1A may exert an effect on aggression through mechanisms involving nucleus accumbens and amygdala volumes, respectively.
  • Van Leeuwen, E. J. C., Cronin, K. A., & Haun, D. B. M. (2018). Population-specific social dynamics in chimpanzees. Proceedings of the National Academy of Sciences of the United States of America, 115(45), 11393-11400. doi:10.1073/pnas.1722614115.

    Abstract

    Understanding intraspecific variation in sociality is essential for characterizing the flexibility and evolution of social systems, yet its study in nonhuman animals is rare. Here, we investigated whether chimpanzees exhibit population-level differences in sociality that cannot be easily explained by differences in genetics or ecology. We compared social proximity and grooming tendencies across four semiwild populations of chimpanzees living in the same ecological environment over three consecutive years, using both linear mixed models and social network analysis. Results indicated temporally stable, population-level differences in dyadic-level sociality. Moreover, group cohesion measures capturing network characteristics beyond dyadic interactions (clustering, modularity, and social differentiation) showed population-level differences consistent with the dyadic indices. Subsequently, we explored whether the observed intraspecific variation in sociality could be attributed to cultural processes by ruling out alternative sources of variation including the influences of ecology, genetics, and differences in population demographics. We conclude that substantial variation in social behavior exists across neighboring populations of chimpanzees and that this variation is in part shaped by cultural processes.

    Additional information

    pnas.1722614115.sapp.pdf
  • Van Ackeren, M. J., Casasanto, D., Bekkering, H., Hagoort, P., & Rueschemeyer, S.-A. (2012). Pragmatics in action: Indirect requests engage theory of mind areas and the cortical motor network. Journal of Cognitive Neuroscience, 24, 2237-2247. doi:10.1162/jocn_a_00274.

    Abstract

    Research from the past decade has shown that understanding the meaning of words and utterances (i.e., abstracted symbols) engages the same systems we used to perceive and interact with the physical world in a content-specific manner. For example, understanding the word “grasp” elicits activation in the cortical motor network, that is, part of the neural substrate involved in planned and executing a grasping action. In the embodied literature, cortical motor activation during language comprehension is thought to reflect motor simulation underlying conceptual knowledge [note that outside the embodied framework, other explanations for the link between action and language are offered, e.g., Mahon, B. Z., & Caramazza, A. A critical look at the embodied cognition hypothesis and a new proposal for grouding conceptual content. Journal of Physiology, 102, 59–70, 2008; Hagoort, P. On Broca, brain, and binding: A new framework. Trends in Cognitive Sciences, 9, 416–423, 2005]. Previous research has supported the view that the coupling between language and action is flexible, and reading an action-related word form is not sufficient for cortical motor activation [Van Dam, W. O., van Dijk, M., Bekkering, H., & Rueschemeyer, S.-A. Flexibility in embodied lexical–semantic representations. Human Brain Mapping, doi: 10.1002/hbm.21365, 2011]. The current study goes one step further by addressing the necessity of action-related word forms for motor activation during language comprehension. Subjects listened to indirect requests (IRs) for action during an fMRI session. IRs for action are speech acts in which access to an action concept is required, although it is not explicitly encoded in the language. For example, the utterance “It is hot here!” in a room with a window is likely to be interpreted as a request to open the window. However, the same utterance in a desert will be interpreted as a statement. The results indicate (1) that comprehension of IR sentences activates cortical motor areas reliably more than comprehension of sentences devoid of any implicit motor information. This is true despite the fact that IR sentences contain no lexical reference to action. (2) Comprehension of IR sentences also reliably activates substantial portions of the theory of mind network, known to be involved in making inferences about mental states of others. The implications of these findings for embodied theories of language are discussed.
  • Van de Ven, M., Ernestus, M., & Schreuder, R. (2012). Predicting acoustically reduced words in spontaneous speech: The role of semantic/syntactic and acoustic cues in context. Laboratory Phonology, 3, 455-481. doi:10.1515/lp-2012-0020.

    Abstract

    In spontaneous speech, words may be realised shorter than in formal speech (e.g., English yesterday may be pronounced like [jɛʃeɩ]). Previous research has shown that context is required to understand highly reduced pronunciation variants. We investigated the extent to which listeners can predict low predictability reduced words on the basis of the semantic/syntactic and acoustic cues in their context. In four experiments, participants were presented with either the preceding context or the preceding and following context of reduced words, and either heard these fragments of conversational speech, or read their orthographic transcriptions. Participants were asked to predict the missing reduced word on the basis of the context alone, choosing from four plausible options. Participants made use of acoustic cues in the context, although casual speech typically has a high speech rate, and acoustic cues are much more unclear than in careful speech. Moreover, they relied on semantic/syntactic cues. Whenever there was a conflict between acoustic and semantic/syntactic contextual cues, measured as the word's probability given the surrounding words, listeners relied more heavily on acoustic cues. Further, context appeared generally insufficient to predict the reduced words, underpinning the significance of the acoustic characteristics of the reduced words themselves.
  • Van de Ven, M., & Ernestus, M. (2018). The role of segmental and durational cues in the processing of reduced words. Language and Speech, 61(3), 358-383. doi:10.1177/0023830917727774.

    Abstract

    In natural conversations, words are generally shorter and they often lack segments. It is unclear to what extent such durational and segmental reductions affect word recognition. The present study investigates to what extent reduction in the initial syllable hinders word comprehension, which types of segments listeners mostly rely on, and whether listeners use word duration as a cue in word recognition. We conducted three experiments in Dutch, in which we adapted the gating paradigm to study the comprehension of spontaneously uttered conversational speech by aligning the gates with the edges of consonant clusters or vowels. Participants heard the context and some segmental and/or durational information from reduced target words with unstressed initial syllables. The initial syllable varied in its degree of reduction, and in half of the stimuli the vowel was not clearly present. Participants gave too short answers if they were only provided with durational information from the target words, which shows that listeners are unaware of the reductions that can occur in spontaneous speech. More importantly, listeners required fewer segments to recognize target words if the vowel in the initial syllable was absent. This result strongly suggests that this vowel hardly plays a role in word comprehension, and that its presence may even delay this process. More important are the consonants and the stressed vowel.
  • Van Berkum, J. J. A. (2012). Zonder gevoel geen taal. Neerlandistiek.nl. Wetenschappelijk tijdschrift voor de Nederlandse taal- en letterkunde, 12(01).

    Abstract

    Geïllustreerde herpublicatie van de oratie, uitgesproken bij het aanvaarden van de leeropdracht Discourse, cognitie en communicatie op 30 september 2011 (Universiteit Utrecht). In tegenstelling tot de oorspronkelijke oratie-tekst bevat deze herpublicatie ook diverse illustraties en links. Daarnaast is er in twee aansluitende artikelen door vakgenoten op gereageerd (zie http://www.neerlandistiek.nl/12.01a/ en http://www.neerlandistiek.nl/12.01b/)
  • Vanderauwera, J., De Vos, A., Forkel, S. J., Catani, M., Wouters, J., Vandermosten, M., & Ghesquière, P. (2018). Neural organization of ventral white matter tracts parallels the initial steps of reading development: A DTI tractography study. Brain and Language, 183, 32-40. doi:10.1016/j.bandl.2018.05.007.

    Abstract

    Insight in the developmental trajectory of the neuroanatomical reading correlates is important to understand related cognitive processes and disorders. In adults, a dual pathway model has been suggested encompassing a dorsal phonological and a ventral orthographic white matter system. This dichotomy seems not present in pre-readers, and the specific role of ventral white matter in reading remains unclear. Therefore, the present longitudinal study investigated the relation between ventral white matter and cognitive processes underlying reading in children with a broad range of reading skills (n = 61). Ventral pathways of the reading network were manually traced using diffusion tractography: the inferior fronto-occipital fasciculus (IFOF), inferior longitudinal fasciculus (ILF) and uncinate fasciculus (UF). Pathways were examined pre-reading (5–6 years) and after two years of reading acquisition (7–8 years). Dimension reduction for the cognitive measures resulted in one component for pre-reading cognitive measures and a separate phonological and orthographic component for the early reading measures. Regression analyses revealed a relation between the pre-reading cognitive component and bilateral IFOF and left ILF. Interestingly, exclusively the left IFOF was related to the orthographic component, whereas none of the pathways was related to the phonological component. Hence, the left IFOF seems to serve as the lexical reading route, already in the earliest reading stages.
  • Vanlangendonck, F., Takashima, A., Willems, R. M., & Hagoort, P. (2018). Distinguishable memory retrieval networks for collaboratively and non-collaboratively learned information. Neuropsychologia, 111, 123-132. doi:10.1016/j.neuropsychologia.2017.12.008.

    Abstract

    Learning often occurs in communicative and collaborative settings, yet almost all research into the neural basis of memory relies on participants encoding and retrieving information on their own. We investigated whether learning linguistic labels in a collaborative context at least partly relies on cognitively and neurally distinct representations, as compared to learning in an individual context. Healthy human participants learned labels for sets of abstract shapes in three different tasks. They came up with labels with another person in a collaborative communication task (collaborative condition), by themselves (individual condition), or were given pre-determined unrelated labels to learn by themselves (arbitrary condition). Immediately after learning, participants retrieved and produced the labels aloud during a communicative task in the MRI scanner. The fMRI results show that the retrieval of collaboratively generated labels as compared to individually learned labels engages brain regions involved in understanding others (mentalizing or theory of mind) and autobiographical memory, including the medial prefrontal cortex, the right temporoparietal junction and the precuneus. This study is the first to show that collaboration during encoding affects the neural networks involved in retrieval.
  • Vanlangendonck, F., Willems, R. M., & Hagoort, P. (2018). Taking common ground into account: Specifying the role of the mentalizing network in communicative language production. PLoS One, 13(10): e0202943. doi:10.1371/journal.pone.0202943.
  • Varma, S., Daselaar, S. M., Kessels, R. P. C., & Takashima, A. (2018). Promotion and suppression of autobiographical thinking differentially affect episodic memory consolidation. PLoS One, 13(8): e0201780. doi:10.1371/journal.pone.0201780.

    Abstract

    During a post-encoding delay period, the ongoing consolidation of recently acquired memories can suffer interference if the delay period involves encoding of new memories, or sensory stimulation tasks. Interestingly, two recent independent studies suggest that (i) autobiographical thinking also interferes markedly with ongoing consolidation of recently learned wordlist material, while (ii) a 2-Back task might not interfere with ongoing consolidation, possibly due to the suppression of autobiographical thinking. In this study, we directly compare these conditions against a quiet wakeful rest baseline to test whether the promotion (via familiar sound-cues) or suppression (via a 2-Back task) of autobiographical thinking during the post-encoding delay period can affect consolidation of studied wordlists in a negative or a positive way, respectively. Our results successfully replicate previous studies and show a significant interference effect (as compared to the rest condition) when learning is followed by familiar sound-cues that promote autobiographical thinking, whereas no interference effect is observed when learning is followed by the 2-Back task. Results from a post-experimental experience-sampling questionnaire further show significant differences in the degree of autobiographical thinking reported during the three post-encoding periods: highest in the presence of sound-cues and lowest during the 2-Back task. In conclusion, our results suggest that varying levels of autobiographical thought during the post-encoding period may modulate episodic memory consolidation.
  • Verdonschot, R. G., & Kinoshita, S. (2018). Mora or more? The phonological unit of Japanese word production in the Stroop color naming task. Memory & Cognition, 46(3), 410-425. doi:10.3758/s13421-017-0774-4.

    Abstract

    In English, Dutch, and other European languages, it is well established that the fundamental phonological unit in word production is the phoneme; in contrast, recent studies have shown that in Chinese it is the (atonal) syllable and in Japanese the mora. The present study investigated whether this cross-language variation in the size of the unit of word production is due to the type of script used in the language (i.e., alphabetic, morphosyllabic, or moraic). Capitalizing on the multiscriptal nature of Japanese, and using the Stroop color naming task, we show that the overlap in the initial mora between the color name and the written distractor facilitates color naming independent of script type. These results confirm the mora as the phonological unit of word production in Japanese, and establish the Stroop color naming task as a useful task for investigating the fundamental (or "proximate") phonological unit used in speech production.
  • Verdonschot, R. G., Middelburg, R., Lensink, S. E., & Schiller, N. O. (2012). Morphological priming survives a language switch. Cognition, 124(3), 343-349. doi:10.1016/j.cognition.2012.05.019.

    Abstract

    In a long-lag morphological priming experiment, Dutch (L1)-English (L2) bilinguals were asked to name pictures and read aloud words. A design using non-switch blocks, consisting solely of Dutch stimuli, and switch-blocks, consisting of Dutch primes and targets with intervening English trials, was administered. Target picture naming was facilitated by morphologically related primes in both non-switch and switch blocks with equal magnitude. These results contrast some assumptions of sustained reactive inhibition models. However, models that do not assume bilinguals having to reactively suppress all activation of the non-target language can account for these data. (C) 2012 Elsevier B.V. All rights reserved.
  • Verheijen, J., Van der Zee, J., Gijselinck, I., Van den Bossche, T., Dillen, L., Heeman, B., Gómez-Tortosa, E., Lladó, A., Sanchez-Valle, R., Graff, C., Pastor, P., Pastor, M. A., Benussi, L., Ghidoni, R., Binetti, G., Clarimon, J., De Mendonça, A., Gelpi, E., Tsolaki, M., Diehl-Schmid, J. and 12 moreVerheijen, J., Van der Zee, J., Gijselinck, I., Van den Bossche, T., Dillen, L., Heeman, B., Gómez-Tortosa, E., Lladó, A., Sanchez-Valle, R., Graff, C., Pastor, P., Pastor, M. A., Benussi, L., Ghidoni, R., Binetti, G., Clarimon, J., De Mendonça, A., Gelpi, E., Tsolaki, M., Diehl-Schmid, J., Nacmias, B., Almeida, M. R., Borroni, B., Matej, R., Ruiz, A., Engelborghs, S., Vandenberghe, R., De Deyn, P. P., Cruts, M., Van Broeckhoven, C., Sleegers, K., BELNEU Consortium, & EU EOD Consortium (2018). Common and rare TBK1 variants in early-onset Alzheimer disease in a European cohort. Neurobiology of Aging, 62, 245.e1-245.e7. doi:10.1016/j.neurobiolaging.2017.10.012.

    Abstract

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

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

    Abstract

    Over 25 genes are known to affect the risk of developing Alzheimer disease (AD), the most common neurodegenerative dementia. However, mechanistic insights and improved disease management remains limited, due to difficulties in determining the functional consequences of genetic associations. Transcriptomics is increasingly being used to corroborate or enhance interpretation of genetic discoveries. These approaches, which include second and third generation sequencing, single-cell sequencing, and bioinformatics, reveal allele-specific events connecting AD risk genes to expression profiles, and provide converging evidence of pathophysiological pathways underlying AD. Simultaneously, they highlight brain region- and cell-type-specific expression patterns, and alternative splicing events that affect the straightforward relation between a genetic variant and AD, re-emphasizing the need for an integrated approach of genetics and transcriptomics in understanding AD. © 2018 The Authors
  • Viebahn, M., McQueen, J. M., Ernestus, M., Frauenfelder, U. H., & Bürki, A. (2018). How much does orthography influence the processing of reduced word forms? Evidence from novel-word learning about French schwa deletion. The Quarterly Journal of Experimental Psychology, 71(11), 2378-2394. doi:10.1177/1747021817741859.

    Abstract

    This study examines the influence of orthography on the processing of reduced word forms. For this purpose, we compared the impact of phonological variation with the impact of spelling-sound consistency on the processing of words that may be produced with or without the vowel schwa. Participants learnt novel French words in which the vowel schwa was present or absent in the first syllable. In Experiment 1, the words were consistently produced without schwa or produced in a variable manner (i.e., sometimes produced with and sometimes produced without schwa). In Experiment 2, words were always produced in a consistent manner, but an orthographic exposure phase was included in which words that were produced without schwa were either spelled with or without the letter . Results from naming and eye-tracking tasks suggest that both phonological variation and spelling-sound consistency influence the processing of spoken novel words. However, the influence of phonological variation outweighs the effect of spelling-sound consistency. Our findings therefore suggest that the influence of orthography on the processing of reduced word forms is relatively small.
  • von Stutterheim, C., Andermann, M., Carroll, M., Flecken, M., & Schmiedtova, B. (2012). How grammaticized concepts shape event conceptualization in language production: Insights from linguistic analysis, eye tracking data, and memory performance. Linguistics, 50(4), 833-867. doi:10.1515/ling-2012-0026.

    Abstract

    The role of grammatical systems in profiling particular conceptual categories is used as a key in exploring questions concerning language specificity during the conceptualization phase in language production. This study focuses on the extent to which crosslinguistic differences in the concepts profiled by grammatical means in the domain of temporality (grammatical aspect) affect event conceptualization and distribution of attention when talking about motion events. The analyses, which cover native speakers of Standard Arabic, Czech, Dutch, English, German, Russian and Spanish, not only involve linguistic evidence, but also data from an eye tracking experiment and a memory test. The findings show that direction of attention to particular parts of motion events varies to some extent with the existence of grammaticized means to express imperfective/progressive aspect. Speakers of languages that do not have grammaticized aspect of this type are more likely to take a holistic view when talking about motion events and attend to as well as refer to endpoints of motion events, in contrast to speakers of aspect languages.

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  • Vonk, W., Hustinx, L. G., & Simons, W. H. (1992). The use of referential expressions in structuring discourse. Language and Cognitive Processes, 301-333. doi:10.1080/01690969208409389.

    Abstract

    Referential expressions that refer to entities that occur in a text differ in lexical specificity. It is claimed that if these anaphoric expressions are more specific than necessary for their identificational function, they not only relate the current information to the intended referent, but also contribute to the expression of the thematic structure of the discourse and to the comprehension of the thematic structure. In two controlled production experiments, it is demonstrated that thematic shifts are produced when one has to make use of such an overspecified expression, and that overspecified referential expressions are produced when one has to formulate a thematic shift. In two comprehension experiments, using a probe recognition technique, it is shown that an overspecified referential expression decreases the availability of information contained in a sentence that precedes the overspecification. This finding is interpreted in terms of the thematic structuring function of referential expressions in the understanding of discourse.
  • De Vos, J., Schriefers, H., Nivard, M. C., & Lemhöfer, K. (2018). A meta‐analysis and meta‐regression of incidental second language word learning from spoken input. Language Learning, 68(4), 906-941. doi:10.1111/lang.12296.

    Abstract

    We meta‐analyzed the effectiveness of incidental second language word learning from spoken input. Our sample contained 105 effect sizes from 32 primary studies employing meaning‐focused word‐learning activities with 1,964 participants with typical cognitive functioning. The random‐effects meta‐analysis yielded a mean effect size of g = 1.05, reflecting generally large vocabulary gains from spoken input in meaning‐focused activities. A meta‐regression with three substantive and two methodological predictors also revealed that adult participants outperformed children in terms of word learning and that interactive learning tasks were more effective than noninteractive ones. Furthermore, learning scores were higher when measured with recognition than with recall tests. Methodologically, the use of a no‐input control group seemed to protect against an overestimation of learning effects, evidenced by smaller effect sizes. Finally, whether a pretest–posttest design was used did not influence effect sizes. All data and the analysis script are publicly available.
  • De Vos, C., & Palfreyman, N. (2012). [Review of the book Deaf around the World: The impact of language / ed. by Mathur & Napoli]. Journal of Linguistics, 48, 731 -735.

    Abstract

    First paragraph. Since its advent half a century ago, the field of sign language linguistics has had close ties to education and the empowerment of deaf communities, a union that is fittingly celebrated by Deaf around the world: The impact of language. With this fruitful relationship in mind, sign language researchers and deaf educators gathered in Philadelphia in 2008, and in the volume under review, Gaurav Mathur & Donna Jo Napoli (henceforth M&N) present a selection of papers from this conference, organised in two parts: ‘Sign languages: Creation, context, form’, and ‘Social issues/civil rights ’. Each of the chapters is accompanied by a response chapter on the same or a related topic. The first part of the volume focuses on the linguistics of sign languages and includes papers on the impact of language modality on morphosyntax, second language acquisition, and grammaticalisation, highlighting the fine balance that sign linguists need to strike when conducting methodologically sound research. The second part of the book includes accounts by deaf activists from countries including China, India, Japan, Kenya, South Africa and Sweden who are considered prominent figures in areas such as deaf education, politics, culture and international development.
  • De Vos, C., & Nyst, V.A.S (2018). Introduction: The time-depth and typology of rural sign languages. Sign Language Studies, 18(4), 477-487.
  • De Vos, J., Schriefers, H., & Lemhöfer, K. (2018). Noticing vocabulary holes aids incidental second language word learning: An experimental study. Bilingualism: Language and Cognition, 22(3), 500-515. doi:10.1017/S1366728918000019.

    Abstract

    Noticing the hole (NTH) occurs when speakers want to say something, but realise they do not know the right word(s). Such awareness of lacking knowledge supposedly facilitates the acquisition of the unknown word(s) from later input (Swain, 1993). We tested this claim by experimentally inducing NTH in a second language (L2) for some participants (experimental), but not others (control). Then, in a price comparison game, all participants were exposed to spoken L2 input containing the to-be-learned words. They were unaware of taking part in an L2 study. Post-tests showed that participants who had noticed holes in their vocabulary had indeed learned more words compared to participants who had not. This held both for the experimental group as well as those participants in the control group who later reported to have noticed holes. Thus, when we become aware of vocabulary holes, the first step to improve our vocabulary is already taken.
  • De Vries, C., Reijnierse, W. G., & Willems, R. M. (2018). Eye movements reveal readers’ sensitivity to deliberate metaphors during narrative reading. Scientific Study of Literature, 8(1), 135-164. doi:10.1075/ssol.18008.vri.

    Abstract

    Metaphors occur frequently in literary texts. Deliberate Metaphor Theory (DMT; e.g., Steen, 2017) proposes that metaphors that serve a communicative function as metaphor are radically different from metaphors that do not have this function. We investigated differences in processing between deliberate and non-deliberate metaphors, compared to non-metaphorical words in literary reading. Using the Deliberate Metaphor Identification Procedure (Reijnierse et al., 2018), we identified metaphors in two literary stories. Then, eye-tracking was used to investigate participants’ (N = 72) reading behavior. Deliberate metaphors were read slower than non-deliberate metaphors, and both metaphor types were read slower than non-metaphorical words. Differences were controlled for several psycholinguistic variables. Differences in reading behavior were related to individual differences in reading experience and absorption and appreciation of the story. These results are in line with predictions from DMT and underline the importance of distinguishing between metaphor types in the experimental study of literary reading.
  • De Vries, M. H., Petersson, K. M., Geukes, S., Zwitserlood, P., & Christiansen, M. H. (2012). Processing multiple non-adjacent dependencies: Evidence from sequence learning. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 367, 2065-2076. doi:10.1098/rstb.2011.0414.

    Abstract

    Processing non-adjacent dependencies is considered to be one of the hallmarks of human language. Assuming that sequence-learning tasks provide a useful way to tap natural-language-processing mechanisms, we cross-modally combined serial reaction time and artificial-grammar learning paradigms to investigate the processing of multiple nested (A1A2A3B3B2B1) and crossed dependencies (A1A2A3B1B2B3), containing either three or two dependencies. Both reaction times and prediction errors highlighted problems with processing the middle dependency in nested structures (A1A2A3B3_B1), reminiscent of the ‘missing-verb effect’ observed in English and French, but not with crossed structures (A1A2A3B1_B3). Prior linguistic experience did not play a major role: native speakers of German and Dutch—which permit nested and crossed dependencies, respectively—showed a similar pattern of results for sequences with three dependencies. As for sequences with two dependencies, reaction times and prediction errors were similar for both nested and crossed dependencies. The results suggest that constraints on the processing of multiple non-adjacent dependencies are determined by the specific ordering of the non-adjacent dependencies (i.e. nested or crossed), as well as the number of non-adjacent dependencies to be resolved (i.e. two or three). Furthermore, these constraints may not be specific to language but instead derive from limitations on structured sequence learning.
  • Vromans, R. D., & Jongman, S. R. (2018). The interplay between selective and nonselective inhibition during single word production. PLoS One, 13(5): e0197313. doi:10.1371/journal.pone.0197313.

    Abstract

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

    Additional information

    Data available (link to Figshare)
  • Wagensveld, B., Segers, E., Van Alphen, P. M., Hagoort, P., & Verhoeven, L. (2012). A neurocognitive perspective on rhyme awareness: The N450 rhyme effect. Brain Research, 1483, 63-70. doi:10.1016/j.brainres.2012.09.018.

    Abstract

    Rhyme processing is reflected in the electrophysiological signals of the brain as a negative deflection for non-rhyming as compared to rhyming stimuli around 450 ms after stimulus onset. Studies have shown that this N450 component is not solely sensitive to rhyme but also responds to other types of phonological overlap. In the present study, we examined whether the N450 component can be used to gain insight into the global similarity effect, indicating that rhyme judgment skills decrease when participants are presented with word pairs that share a phonological overlap but do not rhyme (e.g., bell–ball). We presented 20 adults with auditory rhyming, globally similar overlapping and unrelated word pairs. In addition to measuring behavioral responses by means of a yes/no button press, we also took EEG measures. The behavioral data showed a clear global similarity effect; participants judged overlapping pairs more slowly than unrelated pairs. However, the neural outcomes did not provide evidence that the N450 effect responds differentially to globally similar and unrelated word pairs, suggesting that globally similar and dissimilar non-rhyming pairs are processed in a similar fashion at the stage of early lexical access.
  • Wagensveld, B., Van Alphen, P. M., Segers, E., & Verhoeven, L. (2012). The nature of rhyme processing in preliterate children. British Journal of Educational Psychology, 82, 672-689. doi:10.1111/j.2044-8279.2011.02055.x.

    Abstract

    Background. Rhyme awareness is one of the earliest forms of phonological awareness to develop and is assessed in many developmental studies by means of a simple rhyme task. The influence of more demanding experimental paradigms on rhyme judgment performance is often neglected. Addressing this issue may also shed light on whether rhyme processing is more global or analytical in nature. Aims. The aim of the present study was to examine whether lexical status and global similarity relations influenced rhyme judgments in kindergarten children and if so, if there is an interaction between these two factors. Sample. Participants were 41 monolingual Dutch-speaking preliterate kindergartners (average age 6.0 years) who had not yet received any formal reading education. Method. To examine the effects of lexical status and phonological similarity processing, the kindergartners were asked to make rhyme judgements on (pseudo) word targets that rhymed, phonologically overlapped or were unrelated to (pseudo) word primes. Results. Both a lexicality effect (pseudo-words were more difficult than words) and a global similarity effect (globally similar non-rhyming items were more difficult to reject than unrelated items) were observed. In addition, whereas in words the global similarity effect was only present in accuracy outcomes, in pseudo-words it was also observed in the response latencies. Furthermore, a large global similarity effect in pseudo-words correlated with a low score on short-term memory skills and grapheme knowledge. Conclusions. Increasing task demands led to a more detailed assessment of rhyme processing skills. Current assessment paradigms should therefore be extended with more demanding conditions. In light of the views on rhyme processing, we propose that a combination of global and analytical strategies is used to make a correct rhyme judgment.
  • Walker, R. M., Hill, A. E., Newman, A. C., Hamilton, G., Torrance, H. S., Anderson, S. M., Ogawa, F., Derizioti, P., Nicod, J., Vernes, S. C., Fisher, S. E., Thomson, P. A., Porteous, D. J., & Evans, K. L. (2012). The DISC1 promoter: Characterization and regulation by FOXP2. Human Molecular Genetics, 21, 2862-2872. doi:10.1093/hmg/dds111.

    Abstract

    Disrupted in schizophrenia 1 (DISC1) is a leading candidate susceptibility gene for schizophrenia, bipolar disorder, and recurrent major depression, which has been implicated in other psychiatric illnesses of neurodevelopmental origin, including autism. DISC1 was initially identified at the breakpoint of a balanced chromosomal translocation, t(1;11) (q42.1;14.3), in a family with a high incidence of psychiatric illness. Carriers of the translocation show a 50% reduction in DISC1 protein levels, suggesting altered DISC1 expression as a pathogenic mechanism in psychiatric illness. Altered DISC1 expression in the post-mortem brains of individuals with psychiatric illness and the frequent implication of non-coding regions of the gene by association analysis further support this assertion. Here, we provide the first characterisation of the DISC1 promoter region. Using dual luciferase assays, we demonstrate that a region -300bp to -177bp relative to the transcription start site (TSS) contributes positively to DISC1 promoter activity, whilst a region -982bp to -301bp relative to the TSS confers a repressive effect. We further demonstrate inhibition of DISC1 promoter activity and protein expression by FOXP2, a transcription factor implicated in speech and language function. This inhibition is diminished by two distinct FOXP2 point mutations, R553H and R328X, which were previously found in families affected by developmental verbal dyspraxia (DVD). Our work identifies an intriguing mechanistic link between neurodevelopmental disorders that have traditionally been viewed as diagnostically distinct but which do share varying degrees of phenotypic overlap.
  • Wang, L., Jensen, O., Van den Brink, D., Weder, N., Schoffelen, J.-M., Magyari, L., Hagoort, P., & Bastiaansen, M. C. M. (2012). Beta oscillations relate to the N400m during language comprehension. Human Brain Mapping, 33, 2898-2912. doi:10.1002/hbm.21410.

    Abstract

    The relationship between the evoked responses (ERPs/ERFs) and the event-related changes in EEG/MEG power that can be observed during sentence-level language comprehension is as yet unclear. This study addresses a possible relationship between MEG power changes and the N400m component of the event-related field. Whole-head MEG was recorded while subjects listened to spoken sentences with incongruent (IC) or congruent (C) sentence endings. A clear N400m was observed over the left hemisphere, and was larger for the IC sentences than for the C sentences. A time–frequency analysis of power revealed a decrease in alpha and beta power over the left hemisphere in roughly the same time range as the N400m for the IC relative to the C condition. A linear regression analysis revealed a positive linear relationship between N400m and beta power for the IC condition, not for the C condition. No such linear relation was found between N400m and alpha power for either condition. The sources of the beta decrease were estimated in the LIFG, a region known to be involved in semantic unification operations. One source of the N400m was estimated in the left superior temporal region, which has been related to lexical retrieval. We interpret our data within a framework in which beta oscillations are inversely related to the engagement of task-relevant brain networks. The source reconstructions of the beta power suppression and the N400m effect support the notion of a dynamic communication between the LIFG and the left superior temporal region during language comprehension.
  • Wang, L., Hagoort, P., & Jensen, O. (2018). Language prediction is reflected by coupling between frontal gamma and posterior alpha oscillations. Journal of Cognitive Neuroscience, 30(3), 432-447. doi:10.1162/jocn_a_01190.

    Abstract

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

    Abstract

    Using magnetoencephalography, the current study examined gamma activity associated with language prediction. Participants read high- and low-constraining sentences in which the final word of the sentence was either expected or unexpected. Although no consistent gamma power difference induced by the sentence-final words was found between the expected and unexpected conditions, the correlation of gamma power during the prediction and activation intervals of the sentence-final words was larger when the presented words matched with the prediction compared with when the prediction was violated or when no prediction was available. This suggests that gamma magnitude relates to the match between predicted and perceived words. Moreover, the expected words induced activity with a slower gamma frequency compared with that induced by unexpected words. Overall, the current study establishes that prediction is related to gamma power correlations and a slowing of the gamma frequency.
  • Wang, L., Bastiaansen, M. C. M., Yang, Y., & Hagoort, P. (2012). Information structure influences depth of syntactic processing: Event-related potential evidence for the Chomsky illusion. PLoS One, 7(10), e47917. doi:10.1371/journal.pone.0047917.

    Abstract

    Information structure facilitates communication between interlocutors by highlighting relevant information. It has previously been shown that information structure modulates the depth of semantic processing. Here we used event-related potentials to investigate whether information structure can modulate the depth of syntactic processing. In question-answer pairs, subtle (number agreement) or salient (phrase structure) syntactic violations were placed either in focus or out of focus through information structure marking. P600 effects to these violations reflect the depth of syntactic processing. For subtle violations, a P600 effect was observed in the focus condition, but not in the non-focus condition. For salient violations, comparable P600 effects were found in both conditions. These results indicate that information structure can modulate the depth of syntactic processing, but that this effect depends on the salience of the information. When subtle violations are not in focus, they are processed less elaborately. We label this phenomenon the Chomsky illusion.
  • Wang, L., Zhu, Z., & Bastiaansen, M. C. M. (2012). Integration or predictability? A further specification of the functional role of gamma oscillations in language comprehension. Frontiers in Psychology, 3, 187. doi:10.3389/fpsyg.2012.00187.

    Abstract

    Gamma-band neuronal synchronization during sentence-level language comprehension has previously been linked with semantic unification. Here, we attempt to further narrow down the functional significance of gamma during language comprehension, by distinguishing between two aspects of semantic unification: successful integration of word meaning into the sentence context, and prediction of upcoming words. We computed event-related potentials (ERPs) and frequency band-specific electroencephalographic (EEG) power changes while participants read sentences that contained a critical word (CW) that was (1) both semantically congruent and predictable (high cloze, HC), (2) semantically congruent but unpredictable (low cloze, LC), or (3) semantically incongruent (and therefore also unpredictable; semantic violation, SV). The ERP analysis showed the expected parametric N400 modulation (HC < LC < SV). The time-frequency analysis showed qualitatively different results. In the gamma-frequency range, we observed a power increase in response to the CW in the HC condition, but not in the LC and the SV conditions. Additionally, in the theta frequency range we observed a power increase in the SV condition only. Our data provide evidence that gamma power increases are related to the predictability of an upcoming word based on the preceding sentence context, rather than to the integration of the incoming word’s semantics into the preceding context. Further, our theta band data are compatible with the notion that theta band synchronization in sentence comprehension might be related to the detection of an error in the language input.
  • Wang, M., Shao, Z., Chen, Y., & Schiller, N. O. (2018). Neural correlates of spoken word production in semantic and phonological blocked cyclic naming. Language, Cognition and Neuroscience, 33(5), 575-586. doi:10.1080/23273798.2017.1395467.

    Abstract

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

    Abstract

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

    Additional information

    1-s2.0-S0006322317321911-mmc1.pdf
  • Wassenaar, M., Hagoort, P., & Brown, C. M. (1997). Syntactic ERP effects in Broca's aphasics with agrammatic comprehension. Brain and Language, 60, 61-64. doi:10.1006/brln.1997.1911.
  • Watson, L. M., Wong, M. M. K., Vowles, J., Cowley, S. A., & Becker, E. B. E. (2018). A simplified method for generating purkinje cells from human-induced pluripotent stem cells. The Cerebellum, 17(4), 419-427. doi:10.1007/s12311-017-0913-2.

    Abstract

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

    Additional information

    12311_2017_913_MOESM1_ESM.docx
  • Weber, A., & Scharenborg, O. (2012). Models of spoken-word recognition. Wiley Interdisciplinary Reviews: Cognitive Science, 3, 387-401. doi:10.1002/wcs.1178.

    Abstract

    All words of the languages we know are stored in the mental lexicon. Psycholinguistic models describe in which format lexical knowledge is stored and how it is accessed when needed for language use. The present article summarizes key findings in spoken-word recognition by humans and describes how models of spoken-word recognition account for them. Although current models of spoken-word recognition differ considerably in the details of implementation, there is general consensus among them on at least three aspects: multiple word candidates are activated in parallel as a word is being heard, activation of word candidates varies with the degree of match between the speech signal and stored lexical representations, and activated candidate words compete for recognition. No consensus has been reached on other aspects such as the flow of information between different processing levels, and the format of stored prelexical and lexical representations. WIREs Cogn Sci 2012
  • Weber, A., & Crocker, M. W. (2012). On the nature of semantic constraints on lexical access. Journal of Psycholinguistic Research, 41, 195-214. doi:10.1007/s10936-011-9184-0.

    Abstract

    We present two eye-tracking experiments that investigate lexical frequency and semantic context constraints in spoken-word recognition in German. In both experiments, the pivotal words were pairs of nouns overlapping at onset but varying in lexical frequency. In Experiment 1, German listeners showed an expected frequency bias towards high-frequency competitors (e.g., Blume, ‘flower’) when instructed to click on low-frequency targets (e.g., Bluse, ‘blouse’). In Experiment 2, semantically constraining context increased the availability of appropriate low-frequency target words prior to word onset, but did not influence the availability of semantically inappropriate high-frequency competitors at the same time. Immediately after target word onset, however, the activation of high-frequency competitors was reduced in semantically constraining sentences, but still exceeded that of unrelated distractor words significantly. The results suggest that (1) semantic context acts to downgrade activation of inappropriate competitors rather than to exclude them from competition, and (2) semantic context influences spoken-word recognition, over and above anticipation of upcoming referents.
  • Weekes, B. S., Abutalebi, J., Mak, H.-K.-F., Borsa, V., Soares, S. M. P., Chiu, P. W., & Zhang, L. (2018). Effect of monolingualism and bilingualism in the anterior cingulate cortex: a proton magnetic resonance spectroscopy study in two centers. Letras de Hoje, 53(1), 5-12. doi:10.15448/1984-7726.2018.1.30954.

    Abstract

    Reports of an advantage of bilingualism on brain structure in young adult participants
    are inconsistent. Abutalebi et al. (2012) reported more efficient monitoring of conflict during the
    Flanker task in young bilinguals compared to young monolingual speakers. The present study
    compared young adult (mean age = 24) Cantonese-English bilinguals in Hong Kong and young
    adult monolingual speakers. We expected (a) differences in metabolites in neural tissue to result
    from bilingual experience, as measured by 1H-MRS at 3T, (b) correlations between metabolic
    levels and Flanker conflict and interference effects (c) different associations in bilingual and
    monolingual speakers. We found evidence of metabolic differences in the ACC due to bilingualism,
    specifically in metabolites Cho, Cr, Glx and NAA. However, we found no significant correlations
    between metabolic levels and conflict and interference effects and no significant evidence of
    differential relationships between bilingual and monolingual speakers. Furthermore, we found no
    evidence of significant differences in the mean size of conflict and interference effects between
    groups i.e. no bilingual advantage. Lower levels of Cho, Cr, Glx and NAA in bilingual adults
    compared to monolingual adults suggest that the brains of bilinguals develop greater adaptive
    control during conflict monitoring because of their extensive bilingual experience.
  • Whitehouse, A. J., Bishop, D. V., Ang, Q., Pennell, C. E., & Fisher, S. E. (2012). Corrigendum to CNTNAP2 variants affect early language development in the general population. Genes, Brain and Behavior, 11, 501. doi:10.1111/j.1601-183X.2012.00806.x.

    Abstract

    Corrigendum to CNTNAP2 variants affect early language development in the general population A. J. O. Whitehouse, D. V. M. Bishop, Q. W. Ang, C. E. Pennell and S. E. Fisher Genes Brain Behav (2011) doi: 10.1111/j.1601-183X.2011.00684.x. The authors have detected a typographical error in the Abstract of this paper. The error is in the fifth sentence, which reads: ‘‘On the basis of these findings, we performed analyses of four-marker haplotypes of rs2710102–rs759178–rs17236239–rs2538976 and identified significant association (haplotype TTAA, P = 0.049; haplotype GCAG,P = .0014).’’ Rather than ‘‘GCAG’’, the final haplotype should read ‘‘CGAG’’. This typographical error was made in the Abstract only and this has no bearing on the results or conclusions of the study, which remain unchanged. Reference Whitehouse, A. J. O., Bishop, D. V. M., Ang, Q. W., Pennell, C. E. & Fisher, S. E. (2011) CNTNAP2 variants affect early language development in the general population. Genes Brain Behav 10, 451–456. doi: 10.1111/j.1601-183X.2011.00684.x.
  • Whitehouse, H., & Cohen, E. (2012). Seeking a rapprochement between anthropology and the cognitive sciences: A problem-driven approach. Topics in Cognitive Science, 4, 404-412. doi:10.1111/j.1756-8765.2012.01203.x.

    Abstract

    Beller, Bender, and Medin question the necessity of including social anthropology within the cognitive sciences. We argue that there is great scope for fruitful rapprochement while agreeing that there are obstacles (even if we might wish to debate some of those specifically identified by Beller and colleagues). We frame the general problem differently, however: not in terms of the problem of reconciling disciplines and research cultures, but rather in terms of the prospects for collaborative deployment of expertise (methodological and theoretical) in problem-driven research. For the purposes of illustration, our focus in this article is on the evolution of cooperation
  • Willems, R. M., & Francken, J. C. (2012). Embodied cognition: Taking the next step. Frontiers in Psychology, 3, 582. doi:10.3389/fpsyg.2012.00582.

    Abstract

    Recent years have seen a large amount of empirical studies related to ‘embodied cognition’. While interesting and valuable, there is something dissatisfying with the current state of affairs in this research domain. Hypotheses tend to be underspecified, testing in general terms for embodied versus disembodied processing. The lack of specificity of current hypotheses can easily lead to an erosion of the embodiment concept, and result in a situation in which essentially any effect is taken as positive evidence. Such erosion is not helpful to the field and does not do justice to the importance of embodiment. Here we want to take stock, and formulate directions for how it can be studied in a more fruitful fashion. As an example we will describe few example studies that have investigated the role of sensori-motor systems in the coding of meaning (‘embodied semantics’). Instead of focusing on the dichotomy between embodied and disembodied theories, we suggest that the field move forward and ask how and when sensori-motor systems and behavior are involved in cognition.
  • Winsvold, B. S., Palta, P., Eising, E., Page, C. M., The International Headache Genetics Consortium, Van den Maagdenberg, A. M. J. M., Palotie, A., & Zwart, J.-A. (2018). Epigenetic DNA methylation changes associated with headache chronification: A retrospective case-control study. Cephalalgia, 38(2), 312-322. doi:10.1177/0333102417690111.

    Abstract

    Background

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

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

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

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

    Abstract

    Researchers have suggested that the vocabularies of languages are oriented towards the communicative needs of language users. Here, we provide evidence demonstrating that the higher frequency of visual words in a large variety of English corpora is reflected in greater lexical differentiation—a greater number of unique words—for the visual domain in the English lexicon. In comparison, sensory modalities that are less frequently talked about, particularly taste and smell, show less lexical differentiation. In addition, we show that even though sensory language can be expected to change across historical time and between contexts of use (e.g., spoken language versus fiction), the pattern of visual dominance is a stable property of the English language. Thus, we show that across the board, precisely those semantic domains that are more frequently talked about are also more lexically differentiated, for perceptual experiences. This correlation between type and token frequencies suggests that the sensory lexicon of English is geared towards communicative efficiency.
  • Wong, M. M. K., Hoekstra, S. D., Vowles, J., Watson, L. M., Fuller, G., Németh, A. H., Cowley, S. A., Ansorge, O., Talbot, K., & Becker, E. B. E. (2018). Neurodegeneration in SCA14 is associated with increased PKCγ kinase activity, mislocalization and aggregation. Acta Neuropathologica Communications, 6: 99. doi:10.1186/s40478-018-0600-7.

    Abstract

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

    Additional information

    additional file
  • Xiang, H., Dediu, D., Roberts, L., Van Oort, E., Norris, D., & Hagoort, P. (2012). The structural connectivity underpinning language aptitude, working memory and IQ in the perisylvian language network. Language Learning, 62(Supplement S2), 110-130. doi:10.1111/j.1467-9922.2012.00708.x.

    Abstract

    We carried out the first study on the relationship between individual language aptitude and structural connectivity of language pathways in the adult brain. We measured four components of language aptitude (vocabulary learning, VocL; sound recognition, SndRec; sound-symbol correspondence, SndSym; and grammatical inferencing, GrInf) using the LLAMA language aptitude test (Meara, 2005). Spatial working memory (SWM), verbal working memory (VWM) and IQ were also measured as control factors. Diffusion Tensor Imaging (DTI) was employed to investigate the structural connectivity of language pathways in the perisylvian language network. Principal Component Analysis (PCA) on behavioural measures suggests that a general ability might be important to the first stages of L2 acquisition. It also suggested that VocL, SndSy and SWM are more closely related to general IQ than SndRec and VocL, and distinguished the tasks specifically designed to tap into L2 acquisition (VocL, SndRec,SndSym and GrInf) from more generic measures (IQ, SWM and VWM). Regression analysis suggested significant correlations between most of these behavioural measures and the structural connectivity of certain language pathways, i.e., VocL and BA47-Parietal pathway, SndSym and inter-hemispheric BA45 pathway, GrInf and BA45-Temporal pathway and BA6-Temporal pathway, IQ and BA44-Parietal pathway, BA47-Parietal pathway, BA47-Temporal pathway and inter-hemispheric BA45 pathway, SWM and inter-hemispheric BA6 pathway and BA47-Parietal pathway, and VWM and BA47-Temporal pathway. These results are discussed in relation to relevant findings in the literature.
  • Yang, J., Zhu, H., & Tian, X. (2018). Group-level multivariate analysis in EasyEEG toolbox: Examining the temporal dynamics using topographic responses. Frontiers in Neuroscience, 12: 468. doi:10.3389/fnins.2018.00468.

    Abstract

    Electroencephalography (EEG) provides high temporal resolution cognitive information from non-invasive recordings. However, one of the common practices-using a subset of sensors in ERP analysis is hard to provide a holistic and precise dynamic results. Selecting or grouping subsets of sensors may also be subject to selection bias, multiple comparison, and further complicated by individual differences in the group-level analysis. More importantly, changes in neural generators and variations in response magnitude from the same neural sources are difficult to separate, which limit the capacity of testing different aspects of cognitive hypotheses. We introduce EasyEEG, a toolbox that includes several multivariate analysis methods to directly test cognitive hypotheses based on topographic responses that include data from all sensors. These multivariate methods can investigate effects in the dimensions of response magnitude and topographic patterns separately using data in the sensor space, therefore enable assessing neural response dynamics. The concise workflow and the modular design provide user-friendly and programmer-friendly features. Users of all levels can benefit from the open-sourced, free EasyEEG to obtain a straightforward solution for efficient processing of EEG data and a complete pipeline from raw data to final results for publication.
  • You, W., Zhang, Q., & Verdonschot, R. G. (2012). Masked syllable priming effects in word and picture naming in Chinese. PLoS One, 7(10): e46595. doi:10.1371/journal.pone.0046595.

    Abstract

    Four experiments investigated the role of the syllable in Chinese spoken word production. Chen, Chen and Ferrand (2003) reported a syllable priming effect when primes and targets shared the first syllable using a masked priming paradigm in Chinese. Our Experiment 1 was a direct replication of Chen et al.'s (2003) Experiment 3 employing CV (e. g., /ba2.ying2/, strike camp) and CVG (e. g., /bai2.shou3/, white haired) syllable types. Experiment 2 tested the syllable priming effect using different syllable types: e. g., CV (/qi4.qiu2/, balloon) and CVN (/qing1.ting2/, dragonfly). Experiment 3 investigated this issue further using line drawings of common objects as targets that were preceded either by a CV (e. g., /qi3/, attempt), or a CVN (e. g., /qing2/, affection) prime. Experiment 4 further examined the priming effect by a comparison between CV or CVN priming and an unrelated priming condition using CV-NX (e. g., /mi2.ni3/, mini) and CVN-CX (e. g., /min2.ju1/, dwellings) as target words. These four experiments consistently found that CV targets were named faster when preceded by CV primes than when they were preceded by CVG, CVN or unrelated primes, whereas CVG or CVN targets showed the reverse pattern. These results indicate that the priming effect critically depends on the match between the structure of the prime and that of the first syllable of the target. The effect obtained in this study was consistent across different stimuli and different tasks (word and picture naming), and provides more conclusive and consistent data regarding the role of the syllable in Chinese speech production.
  • Zavala, R. (1997). Functional analysis of Akatek voice constructions. International Journal of American Linguistics, 63(4), 439-474.

    Abstract

    L'A. étudie les corrélations entre structure syntaxique et fonction pragmatique dans les alternances de voix en akatek, une langue maya appartenant au sous-groupe Q'anjob'ala. Les alternances pragmatiques de voix sont les mécanismes par lesquels les langues encodent les différents degrés de topicalité des deux principaux participants d'un événement sémantiquement transitif, l'agent et le patient. A l'aide d'une analyse quantitative, l'A. évalue la topicalité de ces participants et identifie les structures syntaxiques permettant d'exprimer les quatre principales fonctions de voix en akatek : active-directe, inverse, passive et antipassive
  • Zheng, X., Roelofs, A., Farquhar, J., & Lemhöfer, K. (2018). Monitoring of language selection errors in switching: Not all about conflict. PLoS One, 13(11): e0200397. doi:10.1371/journal.pone.0200397.

    Abstract

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

    Abstract

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

    Additional information

    plcp_a_1363401_sm2537.docx
  • Zhu, Z., Hagoort, P., Zhang, J. X., Feng, G., Chen, H.-C., Bastiaansen, M. C. M., & Wang, S. (2012). The anterior left inferior frontal gyrus contributes to semantic unification. NeuroImage, 60, 2230-2237. doi:10.1016/j.neuroimage.2012.02.036.

    Abstract

    Semantic unification, the process by which small blocks of semantic information are combined into a coherent utterance, has been studied with various types of tasks. However, whether the brain activations reported in these studies are attributed to semantic unification per se or to other task-induced concomitant processes still remains unclear. The neural basis for semantic unification in sentence comprehension was examined using event-related potentials (ERP) and functional Magnetic Resonance Imaging (fMRI). The semantic unification load was manipulated by varying the goodness of fit between a critical word and its preceding context (in high cloze, low cloze and violation sentences). The sentences were presented in a serial visual presentation mode. The participants were asked to perform one of three tasks: semantic congruency judgment (SEM), silent reading for comprehension (READ), or font size judgment (FONT), in separate sessions. The ERP results showed a similar N400 amplitude modulation by the semantic unification load across all of the three tasks. The brain activations associated with the semantic unification load were found in the anterior left inferior frontal gyrus (aLIFG) in the FONT task and in a widespread set of regions in the other two tasks. These results suggest that the aLIFG activation reflects a semantic unification, which is different from other brain activations that may reflect task-specific strategic processing.

    Additional information

    Zhu_2012_suppl.dot
  • Zoefel, B., Ten Oever, S., & Sack, A. T. (2018). The involvement of endogenous neural oscillations in the processing of rhythmic input: More than a regular repetition of evoked neural responses. Frontiers in Neuroscience, 12: 95. doi:10.3389/fnins.2018.00095.

    Abstract

    It is undisputed that presenting a rhythmic stimulus leads to a measurable brain response that follows the rhythmic structure of this stimulus. What is still debated, however, is the question whether this brain response exclusively reflects a regular repetition of evoked responses, or whether it also includes entrained oscillatory activity. Here we systematically present evidence in favor of an involvement of entrained neural oscillations in the processing of rhythmic input while critically pointing out which questions still need to be addressed before this evidence could be considered conclusive. In this context, we also explicitly discuss the potential functional role of such entrained oscillations, suggesting that these stimulus-aligned oscillations reflect, and serve as, predictive processes, an idea often only implicitly assumed in the literature.
  • Zwaan, R. A., Van der Stoep, N., Guadalupe, T., & Bouwmeester, S. (2012). Language comprehension in the balance: The robustness of the action-compatibility effect (ACE). PLoS One, 7(2), e31204. doi:10.1371/journal.pone.0031204.

    Abstract

    How does language comprehension interact with motor activity? We investigated the conditions under which comprehending an action sentence affects people's balance. We performed two experiments to assess whether sentences describing forward or backward movement modulate the lateral movements made by subjects who made sensibility judgments about the sentences. In one experiment subjects were standing on a balance board and in the other they were seated on a balance board that was mounted on a chair. This allowed us to investigate whether the action compatibility effect (ACE) is robust and persists in the face of salient incompatibilities between sentence content and subject movement. Growth-curve analysis of the movement trajectories produced by the subjects in response to the sentences suggests that the ACE is indeed robust. Sentence content influenced movement trajectory despite salient inconsistencies between implied and actual movement. These results are interpreted in the context of the current discussion of embodied, or grounded, language comprehension and meaning representation.
  • Zwitserlood, I., Perniss, P. M., & Ozyurek, A. (2012). An empirical investigation of expression of multiple entities in Turkish Sign Language (TİD): Considering the effects of modality. Lingua, 122, 1636 -1667. doi:10.1016/j.lingua.2012.08.010.

    Abstract

    This paper explores the expression of multiple entities in Turkish Sign Language (Türk İşaret Dili; TİD), a less well-studied sign language. It aims to provide a comprehensive description of the ways and frequencies in which entity plurality in this language is expressed, both within and outside the noun phrase. We used a corpus that includes both elicited and spontaneous data from native signers. The results reveal that most of the expressions of multiple entities in TİD are iconic, spatial strategies (i.e. localization and spatial plural predicate inflection) none of which, we argue, should be considered as genuine plural marking devices with the main aim of expressing plurality. Instead, the observed devices for localization and predicate inflection allow for a plural interpretation when multiple locations in space are used. Our data do not provide evidence that TİD employs (productive) morphological plural marking (i.e. reduplication) on nouns, in contrast to some other sign languages and many spoken languages. We relate our findings to expression of multiple entities in other signed languages and in spoken languages and discuss these findings in terms of modality effects on expression of multiple entities in human language.

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